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Supplementation Effects on Early Weaned Calves Grazing Cool- and Warm-Season Grasses

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PAGE 1

SUPPLEMENTATION EFFECTS ON EA RLY WEANED CALVES GRAZING COOLAND WARM-SEASON GRASSES By JOAO MAURICIO BUENO VENDRAMINI A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2005

PAGE 2

Copyright 2005 by Joao Mauricio Bueno Vendramini

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To the memory of my grandfather, Jose Pereira Bueno For being an inspiration for me and my brothers

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ACKNOWLEDGMENTS The author wishes to express his sincere gratitude to Dr. Lynn E. Sollenberger, chairman of the advisory committee, for his guidance, support, and patience throughout the graduate program. Appreciation and gratitude are extended to Dr. John D. Arthington, co-chair of the advisory committee, for his support, enthusiasm, and friendship. Thanks are extended to other members of the advisory committee, Dr. Carrol Chambliss (deceased), Dr. Adegbola Adesogan, Dr. Todd Thrift, and Dr. Rob Kalmbacher, for their contributions to the degree program in the form of suggestions and constructive criticism. Also, the author thanks Dr. Jerry Bennett, department chairman, and Dr. David S. Wofford, graduate coordinator, for the opportunity to study in the Agronomy Department. Special appreciation is due to Dr. John Moore, Dr. John Fike, and Dr. Bisoondat Maccon for their contribution and help with intake statistical models. The contribution of several individuals was valuable to the successful completion of the research project. To fellow agronomy graduate students Jose Dubeux, Sindy Interrante, and Lawton Stewart thanks are extended for their help and friendship during the graduate program. Appreciation is extended to fellow graduate students from different departments and institutions that collaborated in the research project, Nathan and Wimberley Krueger, Marcia Grise, Ana Claudia Rugieri, Helder Santos, Flavio Silvestre, Guto Castro, and Reinaldo Cooke. Special thanks to Agronomy Department staff Claude Jones and Dwight Thomas, at the Forage Evaluation Field Laboratory, for their valuable iv

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help in the field during the sample collection, animal weighing, and overall management of the experimental area. Also, thanks go to Richard Fethiere for his help during the sample analyses at the Forage Evaluation Support Laboratory. Thanks are due to the families of Jose, Georgia and Victor Dubeux, Marcelo and Aline Carvalho, Denis and Ann Vitorelli, Gabriel, Paolete, Julia, and Alice Couto, and others that made us feel at home during graduate school in Gainesville. Last but not least, the author is grateful to his wife, Maria Lucia, parents, Arnaldo and Lais, and brothers, Jose Olavo and Paulo Emilio (deceased), for their love, patience, and support. v

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TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................iv LIST OF TABLES .............................................................................................................ix LIST OF FIGURES ..........................................................................................................xii ABSTRACT .....................................................................................................................xiv CHAPTERS 1 INTRODUCTION........................................................................................................1 2 LITERATURE REVIEW.............................................................................................4 Early Weaning..............................................................................................................4 Forage Systems in Southern USA................................................................................8 Bermudagrass...............................................................................................................9 General..................................................................................................................9 Fertilization, Yield, and Nutritive Value.............................................................10 Grazing Studies...................................................................................................11 Cool-Season Annual Grasses......................................................................................12 General................................................................................................................12 Fertilization, Yield, and Nutritive Value.............................................................13 Grazing Studies...................................................................................................15 Supplementation.........................................................................................................18 Protein..................................................................................................................19 Energy..................................................................................................................21 Interactions between Supplements and Forages..................................................23 Behavior..............................................................................................................25 Intake..........................................................................................................................26 Measurements of Forage Intake of Grazing Animals..........................................27 N Fractioning and in situ Methodology......................................................................32 In situ Procedure..................................................................................................33 In situ Disappearance of Warmand Cool-season Grasses.................................35 Other Methods to Estimate Ruminal Protein Degradation..................................37 vi

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3 SUPPLEMENTATION EFFECTS ON FORAGE CHARACTERISTICS AND PERFORMANCE OF EARLY WEANED CALVES GRAZING RYE-RYEGRASS PASTURES..........................................................................................39 Introduction.................................................................................................................39 Material and Methods.................................................................................................41 Pasture Sampling.................................................................................................43 Animal Response Variables................................................................................45 Economic Analysis..............................................................................................47 Statistical Analysis......................................................................................................48 Results and Discussion...............................................................................................50 Pasture Responses...............................................................................................50 Animal Response Variables................................................................................59 Economic Analysis..............................................................................................65 Summary and Conclusions.........................................................................................65 4 SUPPLEMENTATION EFFECTS ON FORAGE CHARACTERISTICS AND PERFORMANCE OF EARLY WEANED CALVES GRAZING TIFTON 85 BERMUDAGRASS PASTURES..............................................................................67 Introduction.................................................................................................................67 Material and Methods.................................................................................................69 Pasture Sampling.................................................................................................71 Animal Response Variables................................................................................71 Economic Analysis..............................................................................................72 Statistical Analysis......................................................................................................72 Results and Discussion...............................................................................................73 Pasture Responses...............................................................................................73 Animal Response Variables................................................................................80 Economic Analysis..............................................................................................85 Summary and Conclusions.........................................................................................86 5 BOTANICAL COMPOSITION, HERBAGE ACCUMULATION, NUTRITIVE VALUE, AND CRUDE PROTEIN FRACTIONATION OF RYE-RYEGRASS HERBAGE AT DIFFERENT REGROWTH INTERVALS AND N FERTILIZATION LEVELS.......................................................................................87 Introduction.................................................................................................................87 Material and Methods.................................................................................................90 Laboratory Analyses............................................................................................91 In situ Disappearance Procedure.........................................................................92 Dry Matter and Crude Protein Degradation Kinetics..........................................93 Statistical Analyses..............................................................................................94 Results and Discussion...............................................................................................96 Botanical Composition, Herbage Accumulation, and Nutritive Value...............96 Protein Fractionation of the Forage.....................................................................99 vii

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Degradation Parameters of DM and CP............................................................104 Summary and Conclusions.......................................................................................106 6 HERBAGE ACCUMULATION, NUTRITIVE VALUE, AND CRUDE PROTEIN FRACTIONS OF TIFTON 85 SWARDS AT DIFFERENT CUTTING INTERVALS AND N FERTILIZATION LEVELS................................................108 Introduction...............................................................................................................108 Material and Methods...............................................................................................110 Dry Matter and Crude Protein Degradation Kinetics........................................112 Statistical Analyses............................................................................................112 Results and Discussion.............................................................................................114 Herbage Accumulation and Nutritive Value.....................................................114 Protein Fractionation of the Forage...................................................................117 Disappearance Parameters of DM and CP........................................................119 Summary and Conclusions.......................................................................................124 7 SUMMARY AND CONCLUSIONS.......................................................................126 Rye-ryegrass Grazing Study..............................................................................127 Tifton 85 Bermudagrass Grazing Study............................................................128 Rye-Ryegrass Plot Study...................................................................................130 Tifton 85 Bermudagrass Plot Study..................................................................132 Implications of the Research.............................................................................133 APPENDIX A DATA TABLES A...................................................................................................136 B DATA TABLES B...................................................................................................140 LIST OF REFERENCES.................................................................................................142 BIOGRAPHICAL SKETCH...........................................................................................159 viii

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LIST OF TABLES Table page 3-1 Ingredient and chemical composition of supplement fed to early weaned calves on pasture (DM basis)..............................................................................................44 3-2 Pregraze herbage mass and herbage accumulation of rye-ryegrass pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight)...................................................................................................51 3-3 Herbage crude protein (CP) and in vitro digestible organic matter (IVDOM) concentrations of rye-ryegrass pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight).......................52 3-4 Herbage allowance and SR of rye-ryegrass pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight)............56 3-5 Average daily gain (ADG) and liveweight gain (LWG) of early weaned calves grazing rye-ryegrass pastures and supplemented with three levels of concentrate. (BW=body weight)...................................................................................................61 3-6 Total organic matter intake (OMI) and forage OMI of rye-ryegrass pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight)...................................................................................................62 3-7 Grazing time during daylight hours of early weaned calves grazing rye-ryegrass pastures and supplemented with different levels of concentrate..............................63 3-8 Blood urea nitrogen (BUN) concentrations of early weaned calves grazing rye-ryegrass pastures supplemented with three levels of concentrate. (BW=body weight)......................................................................................................................64 3-9 Economic analysis of three concentrate levels on early weaned calves grazing rye-ryegrass pastures................................................................................................65 4-1 Pregraze herbage mass and herbage accumulation of Tifton 85 pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight)...................................................................................................73 ix

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4-2 Crude protein (CP) and in vitro digestible organic matter (IVDOM) concentration of Tifton 85 pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight)..............................................76 4-3 Herbage allowance and stocking rate of Tifton 85 pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight)......................................................................................................................79 4-4 Average daily gain (ADG) and liveweight gain (LWG) ha -1 of early weaned calves grazing Tifton 85 pastures supplemented with different levels of concentrate. (BW=body weight)..............................................................................81 4-5 Total organic matter intake (OMI) and forage OMI of Tifton 85 pastures grazed by early weaned calves supplemented with three levels of concentrate..................82 4-6 Grazing time during daylight hours of early weaned calves grazing Tifton 85 pastures supplemented with three levels of concentrate. (BW=body weight).........84 4-7 Blood urea nitrogen (BUN) concentrations of early weaned calves grazing Tifton 85 pastures supplemented with three levels of concentrate. (BW=body weight)......................................................................................................................85 4-8 Economic analysis of three concentrate levels for early weaned calves grazing Tifton 85 pastures.....................................................................................................85 5-1 Year*season interaction forage effects on botanical composition, crude protein (CP), and in vitro digestible organic matter (IVDOM) of rye-ryegrass mixtures..100 5-2 Season*regrowth interval interaction effects for herbage accumulation and crude protein (CP) and in vitro digestible organic matter (IVDOM) concentrations of rye-ryegrass forage mixtures during 2003 and 2004..............................................101 5-3 N*season interaction effects on crude protein (CP) and in vitro digestible organic matter (IVDOM) concentrations of rye-ryegrass forage...........................102 5-4 Effect of N fertilization levels on DM and CP degradation parameters of rye-ryegrass forage.......................................................................................................105 6-1 N fertilization effects on herbage accumulation and in vitro digestible organic matter (IVDOM) of Tifton 85 forage during 42-d periods....................................116 6-2 Regrowth interval*N fertilization level interaction effects on CP concentrations of Tifton 85 forage.................................................................................................117 6-3 Effect of N fertilization rates on dry matter (DM) and crude protein (CP) degradation of Tifton 85 forage.............................................................................121 x

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6-4 Effect of regrowth interval on dry matter (DM) and crude protein (CP) degradation of Tifton 85 forages............................................................................122 A-1 Weather data for Years 2002, 2003, and 2004 in Gainesville FL..........................136 A-2 Herbage mass double sampling equations and r2 for the rye-ryegrass pastures....137 A-3 Total chromium output of Captec controlled release device in early weaned calves submitted to total feces collection. Year 2003............................................138 A-4 Total chromium output of Captec controlled release device in early weaned calves submitted to total feces collection. Year 2004............................................138 B-1 Herbage evaluation double sample equations and r 2 ..............................................140 B-2 Total chromium output of Captec controlled release device in early weaned calves submitted to total feces collection. Year 2003............................................141 xi

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LIST OF FIGURES Figure page 3-1 Year*period interaction effect on pregraze herbage mass of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate...............................................................................................................53 3-2 Rainfall distribution from October to May 2002/03 and 2003/04...........................53 3-3 Year*period interaction effect on average herbage accumulation of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate...............................................................................................................54 3-4 Year*period interaction effect on crude protein (CP) concentration of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate.................................................................................................57 3-5 Year*period interaction effect on in vitro digestible organic matter (IVDOM) concentration of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate...................................................57 3-6 Year*period interaction effect on herbage allowance of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate...............................................................................................................58 3-7 Year*period interaction effect on average daily gain of early weaned calves grazing rye-ryegrass pastures and supplemented with different levels of concentrate...............................................................................................................61 4-1 Year*period interaction effect on pregraze herbage mass of Tifton 85 pastures grazed by early weaned calves supplemented with different levels of concentrate...............................................................................................................74 4-2 Year*period interaction effect on average herbage accumulation of Tifton 85 pastures grazed by early weaned calves supplemented with different levels of concentrate...............................................................................................................75 4-3 Year*period interaction effect on crude protein (CP) concentration of Tifton 85 pastures grazed by early weaned calves supplemented with different levels of concentrate...............................................................................................................77 xii

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4-4 Period effect on IVDOM concentration of Tifton 85 herbage grazed by early weaned calves supplemented with different levels of concentrate...........................78 4-5 Year*period interaction effect on average daily gain of early weaned calves grazing Tifton 85 pastures and supplemented with different levels of concentrate...............................................................................................................82 5-1 Nitrogen fertilization effects on crude protein (CP) fraction concentrations in total CP of rye-ryegrass forage...............................................................................103 5-2 N fertilization rate on crude protein (CP) disappearance of rye-ryegrass forage...106 6-1 N fertilization (Fig. A) and regrowth interval (Fig. B) effects on crude protein (CP) fraction concentrations in total CP of Tifton 85 forage.................................119 6-2 Tifton 85 crude protein (CP) disappearance at three N fertilization levels (A) and two regrowth intervals (B).....................................................................................123 xiii

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Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy SUPPLEMENTATION EFFECTS ON EARLY WEANED CALVES GRAZING COOLAND WARM-SEASON GRASSES By Joao Mauricio Bueno Vendramini August, 2005 Chair: Lynn E. Sollenberger Cochair: John D. Arthington Major Department: Agronomy Early weaning of their calves increases the likelihood that first calf heifers will rebreed, but there is little information regarding pasture-based feeding programs for the weaned calf. Research was conducted in Gainesville, FL, to evaluate the effect of supplement rates on performance of early weaned (80 d of age) beef calves grazing annual ryegrass (Lolium multiflorum)-rye (Secale cereale) mixtures from January to April (winter) and Tifton 85 bermudagrass (Cynodon sp.) from May to August (summer) 2003 and 2004. Three levels of supplement (10, 15, and 20 g kg -1 of calf liveweight) were evaluated in a completely randomized design with three replications. The supplement contained 147 and 700 g kg -1 of crude protein (CP) and total digestible nutrients, repectively. Pastures were rotationally stocked using a variable stocking rate in both seasons with 21and 14-d regrowth periods for rye-ryegrass and Tifton 85 bermudagrass, respectively. Average daily gain increased linearly during the winter (0.74-0.89 kg) and summer (0.53-0.72 kg) as supplementation rate increased. There was a linear increase in xiv

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live weight gain per hectare with increasing supplement rate during the winter (950-1320 kg) and summer (1100-1780 kg). In addition, two plot studies were conducted with the same species used in the grazing trials to evaluate the effect of N fertilization levels and forage regrowth interval on in situ CP fractionation of the forage. Treatments were the factorial combinations of three N rates (0, 40, and 80 kg ha -1 ) and two regrowth intervals, 3and 6-wk (rye-ryegrass) and 2and 4-wk (Tifton 85). Increasing N fertilization levels increased rye-ryegrass CP Fraction A (readily degradable) (410-590 g kg -1 ) and decreased linearly CP Fraction B (slowly degradable) (510-360 g kg -1 ) and Fraction C (undegradable) (74-48 g kg -1 ). On Tifton 85, increasing N fertilization increased linearly Fraction B (330-455 g kg -1 ) and decreased linearly Fraction C (283-208 g kg -1 ). In addition, greater proportions of fraction C were observed in forage of 4-wk (280 g kg -1 ) than 2-wk maturity (200 g kg -1 ). Results of these studies indicate that pasture-based feeding systems with modest levels of supplementation (10-15 g kg -1 of liveweight) are practical options for raising early weaned calves in Florida. xv

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CHAPTER 1 INTRODUCTION Grasslands cover ~ 4.5 million ha in Florida, most of which are utilized by beef cattle (Bos sp.; Chambliss, 1999). The beef industry is a very important component of Floridas agriculture industry. In 2003, Florida had a total of 950 000 beef cows that accounted for $348 million of income to Florida farmers and ranchers (Florida Agriculture Statistics, 2004). The profitability of the Florida cow-calf enterprise is largely dependent on cow productivity. The low nutritive value of Floridas tropical forages in combination with reduced pasture production during winter reduces cow reproductive performance. Low body condition is responsible for lower conception rates and overall cow herd productivity (Arthington, 2002). These effects are most pronounced in young cows and heifers which have higher nutritional demands to support both lactation and growth (Arthington, 2002). Early weaning calves from young cows and first-calf heifers is a management strategy that has potential to address this problem. At early weaning, cows stop lactating and their requirement for total digestible nutrients (TDN) decreases by as much as 49%. As a result they are able to restore body condition rapidly (Arthington, 2002). Bishop and Wettemann (1990), cited by Lusby (1995), reported that 100% of cows with body condition scores > 5 had begun cycling within 25 d after their calves were weaned at 45 d of age compared to only 43% of cows with condition score < 5. Although the benefits of early weaning, e.g., improving reproduction and reducing nutrient requirements of the cow, have been recognized for many years, producer 1

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2 adoption of early weaning has been limited by lack of management information about the weaned calves (Lusby, 1995). The major expense in raising these calves is feed cost (Lusby and Donald, 1994). Therefore, low-cost alternatives to high concentrate diets are needed. Forage intake of early weaned calves has not been well documented and varies greatly between the time of early and normal weaning due to increasing development of rumen function. The quantity of concentrate supplementation required during this period for calves on pasture is likewise not well defined. Muehlmann et al. (1997) studied different species of tropical pastures as the sole source of feed to early weaned calves. Daily gains were low, averaging 0.2 kg d -1 The National Research Council NRC (1996) reported that calves of 100-kg live-weight must consume 2.6 kg of dry matter d -1 with 250 g kg -1 of protein and 820 g kg -1 of TDN to achieve 1.0 kg of daily weight gain. Considering that most forages and all of the C4 grasses are unable to supply these CP and TDN concentrations, providing the appropriate amount of concentrate supplement to early weaned calves is critical to achieving high performance. Research to quantify the effect of various levels of concentrate on forage intake and calf performance is needed to improve the management of early weaned calves. The research described in this dissertation consists of four experiments. The first two experiments were grazing trials with early weaned calves fed three levels of concentrate supplement while grazing rye (Secale cereale L.)-annual ryegrass (Lolium multiflorum Lam.) pastures during winter (Chapter 3) and Tifton 85 bermudagrass (Cynodon spp.) pastures during summer (Chapter 4). The objectives of these studies were to measure animal responses including average daily gain, stocking rate, live weight gain per unit of pasture area, and forage intake. Pastures were characterized in terms of

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3 herbage mass, accumulation, allowance, and nutritive value. Economic analyses for the different rates of concentrate were performed. These data will allow producers to better assess the profitability of early weaning of beef calves in North-Central Florida. The last two experiments were plot studies with the same forage species used in the grazing trials, rye-ryegrass (Chapter 5) during the winter-spring and Tifton 85 bermudagrass (Chapter 6) during the spring-summer. The primary objective was to characterize the forage N fractions using an in situ technique. This is an important tool for characterizing the rate and extent of N release in the rumen, potentially resulting in better supplementation programs that increase animal performance and decrease loss of N to the environment. Information regarding the N profile of forages is also useful to guide future research and development of animal nutrition models. Additional forage responses, including botanical composition and herbage accumulation, were measured in these studies.

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CHAPTER 2 LITERATURE REVIEW Early Weaning Early weaning is a management practice used by dairy and beef cattle (Bos sp.) producers. The definition of early weaning varies, but typically calves weaned < 150 d of age are considered early weaned. Several authors have reported that increasing the fertility rates of beef cows is a benefit of early weaning. Lusby and Wetteman (1980) compared reproductive performance in first-calf heifers following early and normal weaning of their calves. Conception rate was greater among heifers whose calves were early weaned (EW) (97%) than among those whose calves were normal weaned (NW) (59%) during the 64-d breeding season. The average interval from parturition to conception was reduced from 91 to 73 d by early weaning. According to Arthington and Kalmbacher (2003), early weaning resulted in heavier first-calf heifers with greater body condition score at the time of normal weaning. Heifers with EW calves had higher pregnancy rates than those with NW calves (93 vs. 65%) during a 2-yr study in South Florida. Despite improved pregnancy rates, the practice of early weaning calves is still a challenge for beef cattle producers, in part because of few management options for the weaned calves. Producers that practice EW could possibly send these animals directly to the feedlots; however, this procedure has not been the most profitable. The necessity of keeping the calves during the winter to target the historically greater values of the spring calf market has resulted in recent studies evaluating feed management of EW calves. 4

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5 The ability of the young ruminant to sustain itself nutritionally is highly dependent on the development of a functional rumen. During the first 3 wk of life, the calves are limited to milk feeding. This limitation directly related to the presence or absence and activity of digestive enzymes during early development of the gastrointestinal system. From 3 to 8 wk of age, dry feed consumption may increase to the level at which it contributes significantly to the energy and protein requirements to the animal (Davis and Drackley, 1998). There is a large variability among calves for dry-feed intake after 8 wk of age. This variability probably is related to the rate at which the calf develops physiologically and its health. However, the calves are able to the majority of their energy requirement from solid feed after 8 wk (Davis and Clark, 1981).Based on their physiological development and in order to maintain a 365-d calving interval, calves should be EW at less than 80 d of age. About 40 d of age may be a practical minimum for early weaning in beef herds. Calves of at least 40 d of age do not require milk replacers in the ration and are old enough to eat dry feed. Mild winters in the southern USA offer an opportunity to raise calves on pasture-based systems using high nutritive value cool-season annual forages. According to Arthington (2002), EW calves grazing high quality pastures can grow as fast as NW calves. The practice of raising EW calves on winter pastures is a feasible alternative, especially if pasture quality and quantity are superior to hay available for winter feeding (Barnes et al., 1996). Coffey et al. (2000) stated that calves fed hay plus supplement gained less weight than calves that grazed winter-annual forages. Weder (1997) reported an average daily gain of 0.8 kg d -1 for EW calves grazing annual ryegrass. According to Rouquette et al. (1997), annual ryegrass resulted in individual live weight gains of 1.25

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6 kg d -1 for suckling calves and 1.05 kg d -1 for stocker calves. Early weaned calves grazing wheat (Triticum aestivum L.) pastures consumed 27% less forage dry matter (DM) during the first 20 d than during the subsequent 50 d (Paisley et al., 1998). The authors suggested that initial performance of EW calves grazing winter wheat is limited by low forage intake. Average daily gain for EW calves grazing wheat pasture was 0.86 kg d -1 during a study with fall-born calves in Oklahoma (Purvis et al., 1996). Few studies have been conducted with EW calves grazing summer pastures. Muehlmann et al. (1997) studied tropical pastures as the sole source of feed to EW calves. Calf performance was low with daily gains averaging 0.2 kg d -1 Smeaton et al. (1996) reported that concentrate supplementation gave a substantial improvement in live weight gain (+19 kg) in EW calves grazing on low nutritive value summer pastures. In south Florida, supplementing EW calves with concentrate at 10 g kg -1 of body weight resulted in gains of 0.59 and 0.44 kg d -1 on stargrass (Cynodon nlemfuensis Vanderyst) and atrapaspalum (Paspalum atratum Swallen), respectively (Vendramini et al., 2003). In a study with different forage species, Harvey and Burns (1989) evaluated gain of EW calves grazing hill-land pastures (control), Tillman white clover (Trifolium repens L.), Tillman white clover and tall fescue (Festuca arundinacea Schreb.) mixtures, and pearl millet [Pennisetum glaucum (L.) R. Brown]. Calves on each forage treatment were provided ground ear maize (Zea mays L.) ad libitum. Calf daily gains were 0.72, 0.82, 0.57, and 0.92 kg d -1 respectively. Calves grazing millet had the greatest response in gain and conversion of concentrate to liveweight gain. According to some authors, EW calves have better feed efficiency than NW calves. Myers et al. (1999b) conducted a study comparing EW to NW with or without creep

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7 feeding for beef steers. Early weaned steers had greater average daily gains (ADG) during the entire study than the average of creep-fed or normal-fed steers. Early weaned steers had a lesser intake (7.29 vs. 7.68 kg d -1 ) and better feed conversion (0.16 vs. 0.14 kg gain kg -1 feed) than the average of NW calves with and without creep feeding. Lusby (1995) reported fall EW calves consumed 20% less hay and were 43% more efficient in converting total digestible nutrients (TDN) to weight gain than NW calves. In a study to determine the effect of age at feedlot entry, Schoonmaker et al. (2001) reported that EW calves (110 d of age) were more efficient than NW (202 d of age) and yearling steers (371 d of age) (0.23, 0.21, and 0.18 kg gain kg -1 feed, respectively). Ringwall (2002) evaluated EW calves with an average weight at weaning of 70 kg. They gained 0.9 kg d -1 and the feed efficiency was 5.3 kg of feed kg -1 of gain. When weaned at 70 d, EW calves were 14 kg heavier, had a greater average daily gain, and were 11-d younger at slaughter than NW calves (Grimes and Turner, 1991). Considering meat quality traits at slaughter, EW calves had a higher quality grade than NW calves (Fluharty et al., 2000; Myers et al., 1999b). In a similar study Myers et al. (1999a) did not find differences for carcass weight, longissimus muscle area, or yield grade among calves weaned at 90, 152, and 215 d of age. Moreover, there was no difference in marbling score or percentage of steers grading greater than or equal to Choice or Average Choice. Fluharty et al. (2000) reported that EW calves fed either 100 or 90% concentrate diets were heavier than NW calves at 210 d. The NW calves had greater backfat thickness at 210 d but no difference in longissimus muscle area when compared to EW calves fed a 60% concentrate diet.

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8 Forage Systems in Southern USA Hoveland (1992) estimated that out of ~ 39 million ha of perennial pastures grown in the eastern USA almost 30 million ha are located in the South. These pastures are mainly used for beef cows and calves and stockering of weaned calves. Seasonal variation in amount and nutritive value of forage are the main challenges over the course of the growing season. The predominantly warm-season grasses in Florida are bahiagrass (Paspalum notatum Flgge) and bermudagrass [Cynodon dactylon (L.) Pers.]. These perennial, warm-season forages adapted to the Southeast are typically of lower nutritive value than either cool-season perennials or warm-season annuals (NRC, 1989). Warm-season grasses are more efficient in light conversion than cool-season species, but the major adaptation advantage is their better water-use efficiency, resulting in higher total forage production when compared with cool-season grasses in tropical and subtropical climates (Brown and Simmons, 1979). While the division of physiological activity between mesophyll and bundle sheath cells in warm-season grasses is beneficial for CO 2 fixation and adaptation, the anatomical organization required for warm-season grasses causes forage quality to generally be lower than for cool-season grasses (Akin, 1986). Warm-season grasses have relatively larger proportions of vascular tissues, bundle sheath, and sclerenchyma than cool-season grasses. Some of these tissues in warm-season grasses are potentially digestible, but because of their chemical composition and plant anatomical structure the rate of degradation and fermentation is relatively slow (Coleman et al., 2004). Although warm-season grasses grow well in the summer, their production is seasonal, being interrupted by cold or dry weather.

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9 Cool-season grass species generally are considered to have high nutritive value than warm-season grasses mainly because of the amount and arrangement of tissues (Akin, 1986). Mild winters in Florida give producers the opportunity to cultivate cool-season annual forages during a period when productivity of warm-season grasses is low. The importance of the cool-season grasses in the southern USA can be appreciated by considering the seed market. The annual sale of seeds of winter annuals is enough to plant 6 million ha of winter pastures (Moser and Hoveland, 1996). Bermudagrass General Cynodon spp. are geographically widely distributed. They occur in greatest abundance in tropical and subtemperate environments (Taliaferro et al., 2004). Bermudagrass is one of the most important species in the southeastern USA with ~ 10 to 12 million ha of planted areas for livestock grazing and hay. Many Cynodons have been used as herbage for livestock, but several are of minor value because of narrow distribution or characteristics limiting their adaptation. In 1937, USDA-ARS geneticist Glenn W. Burton initiated a bermudagrass breeding program at the Coastal Plain Experiment Station in Tifton, GA, with the objective of developing cultivars that were more productive than the common strains and were capable of supplying highly nutritious and palatable forage during a greater portion of the year (Burton, 1947). In 1943, the release of Coastal bermudagrass was a landmark in bermudagrass breeding, because it was widely accepted by livestock producers in the southeastern USA. In 1993, Tifton 85, a hybrid between a South African bermudagrass and Tifton 68 stargrass, was released. Burton et al. (1993) described it as taller, with larger culms, broader leaves, and

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10 darker color than other bermudagrass hybrids. Compared with Coastal, Tifton 85 yielded 26% more DM, was 11% more digestible, and was more succulent at harvest. Fertilization, Yield, and Nutritive Value Bermudagrasses require relatively high soil nutrient availability to maintain good production performance. The major determinants of fertilizer response are climate, native soil nutrient status, source and rate of applied nutrients, season of application(s), cultivar, and defoliation regimen and method. Nitrogen has the greatest influence on biomass yield and accordingly influences the amount of other nutrients required to sustain production at specific N levels (Taliaferro et al., 2004). Bermudagrass hay yield and quality are maximized when N is applied in split applications at annual levels higher than 400 kg ha yr (Overman et al., 1992). According to Wilkinson and Langdale (1974), cited by Taliaferro et al. (2004), Coastal and other select bermudagrass cultivars produced maximum yields of about 27 Mg ha -1 at N rates of 1200 kg ha -1 with near linear yield response to N up to about 600 to 700 kg N ha -1 Prine and Burton (1956) evaluated the effects of different N rates and harvest frequencies on Coastal bermudagrass biomass production and nutritive value. There was a curvilinear increase in herbage accumulation as N rate increased from 0 to 1008 kg N ha -1 with maximum DM accumulation of 18.4 Mg ha -1 with a 6to 8-wk harvest interval. Bermudagrass showed the highest efficiency in production (kg DM kg -1 N ha -1 ) at 336 kg N ha -1 The forage CP concentration ranged from 97 to 190 g kg -1 with N fertilization rates from 0 to 1008 kg ha -1 Typically, increased CP concentrations have been reported with increasing N application. Stallcup et al. (1986) reported that CP concentrations in bermudagrass fertilized with 0 and 50 kg N ha -1 were 114 to 143 g kg -1 Burton et al.

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11 (1963) studied different defoliation intervals of Coastal fertilized with 660 kg N ha -1 The maximum herbage accumulation was 22 Mg ha -1 using a 6-wk interval. According to the same author, annual DM production of 20 Mg ha -1 removed 400 to 450 kg N, 30 to 50 kg P, 260 to 320 kg K, 30 kg S, 30 kg Mg, and 60 kg Ca ha -1 from the soil. Johnson et al. (2001b) compared different species of warm-season grasses in Florida. Bermudagrass produced more forage DM (1,540 43 kg ha -1 cutting -1 ) than stargrass (1,400 43 kg ha -1 cutting -1 ) or bahiagrass (1,300 43 kg ha -1 cutting -1 ) when harvested every 28 d. Peak herbage accumulation for all species occurred in late June and July. A quadratic response to N fertilization was noted for in vitro digestible organic matter digestibility (IVDOM) of bermudagrass, whereas bahiagrass was not affected. Bermudagrass was more digestible (575 4 g kg -1 ) than stargrass (546 4 g kg -1 ) and bahiagrass (519 4 g kg -1 ). As fertilization level increased, neutral detergent fiber (NDF) decreased linearly in all three forages. Total N concentration and the concentration of all protein fractions for all forages increased as N fertilization increased. In summary, digestibility, CP concentration, and yield of bermudagrass increased with increasing N application when plants were harvested every 28 d. Grazing Studies Bermudagrass is the most popular warm-season perennial grass for hay production and pastures in the southern USA (Taliaferro et al., 2004). In North-Central Florida, Pedreira et al. (1998) compared Florakirk and Tifton 85 bermudagrass in a 3-yr grazing study. The pastures were continuously stocked and fertilized with 210 kg N ha -1 in four applications yr -1 Although the average daily gain was similar between species (0.6 kg d -1 ), Tifton 85 pastures supported higher average stocking rates (6.0 vs. 4.0 heifers ha -1 ), resulting in greater gain (648 vs. 371 kg ha -1 ). Examination of 3-yr, total-season (169 d)

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12 gains of steers grazing high quality bermudagrass pastures revealed that steer ADGs were 0.90 and 0.88 kg, respectively, from April to July, but only 0.30 and 0.43 kg, respectively, for Tifton 78 and Tifton 85 pastures from July to October (Hill et al., 1993). The authors suggested that decreased ADG was the consequence of increased maintenance requirement of heavier steers and lower nutritive value later in the season. Rouquette et al., (2003) reported that stocker steers gained 1.03 kg d -1 on Tifton 85 compared to 0.75 kg d -1 on Coastal without supplementation. With supplementation of 1.4 kg d -1 of a 3:1 maize:soybean (Glycine max L.) meal concentrate, steer ADGs were 1.25 kg d -1 Carnevalli et al. (1999) maintained Tifton 85 swards heights of 5, 10, 15, or 20 cm with grazing Santa Inez crossbred lambs (Ovis aries L.) paired by sex and body weight. Nutritive value of forage was different for sward heights; short swards (5 cm) had the highest CP and in vitro DM digestibility (IVDMD), but lamb performance was more dependent on carrying capacity and forage allowance, with lambs performing better on taller swards (15 to 20 cm). Cool-Season Annual Grasses General Although warm-season grasses dominate most of the pastures in the southeastern USA, climatic conditions during winter allow for use of cool-season annual grasses, either sod-seeded or in prepared seedbeds. Reasons for use of cool-season annual grasses include the extending the grazing period, high nutritive value, compatibility and ease of establishment in warm-season perennial grass pastures, and tolerance to different defoliation regimens and stocking rates (Rouquette et al., 1997). Small grains are often a component of cool-season grass mixtures because they improve early-season forage

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13 production. Of the options available, annual ryegrass-small grain mixtures provide the earliest and greatest amount of forage if planted early on a well-prepared seedbed; however, they also are expensive and are best used by lactating dairy cows, stocker calves, replacement heifers, or limit grazed by beef cows nursing fall calves (Evers et al., 1997). Annual ryegrass, a high-yielding, high quality grass, is the most commonly grown cool-season pasture forage in the southern and southeastern USA from November to May (Evers et al., 1997). Common annual ryegrass was successfully grown in the Gulf Coast region of the USA in the 1940s and 1950s. However, crown rust (Puccinia coronata f. sp. avenae) was a serious problem and reduced forage yields and quality. Plant breeding efforts in Texas, Mississippi, and Florida resulted in the release of Florida Rust Resistant annual ryegrass (Chapman and Webb, 1965). Hectarage of annual ryegrass planted area has increased due to improved crown rust resistance as well as increased forage and seed yield, and significant gains in winter hardiness (Griffith and Chastain, 1997). Among several cultivars released for southern areas, the tretaploid Jumbo (Reg. no. CV-220, PI 614099) is noted for its high seed yield, high resistance to crown rust and gray leaf spot (Pyricularia grisea [Magnaporthe grisea]) and moderate resistance to Helminthosporium leaf spot disease (Drechslera). Jumbo is late maturing, has larger stems, leaves, seed heads, and seeds than the diploid Surrey, with CP concentration and IVDOM comparing favorably with Surrey and other ryegrass cultivars (Prine et al., 2002). Fertilization, Yield, and Nutritive Value Robinson et al. (1987) reported annual ryegrass yields in the southern states have generally increased with N application rates up to 450 kg ha -1 Hovermale (1993)

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14 obtained maximum ryegrass yields with 380 kg N ha -1 however, the maximum yield was only 6.7 Mg ha -1 Allen et al. (1974) tested different levels of N in oat (Avena sativa L.)-ryegrass mixed swards. Forage yield was 4.3 Mg ha -1 and 9.0 Mg ha -1 with 0 and 280 kg N ha -1 Morris et al. (1994) reported herbage accumulation of 10.5 Mg ha -1 when 280 kg N ha -1 was applied. In Florida, Dunavan (1975) showed that three equal treatments with ammonium nitrate (133 kg ha -1 N application -1 ) applied at planting and in January and March produced higher total yields than a single application of 400 kg N ha -1 Comparing three N fertilization amounts, the rates of N recovery were 46, 45, and 26% for a single application of ammonium nitrate to a rye-ryegrass mixture at N rates of 100, 200, and 400 kg ha -1 respectively. The CP concentration of cool-season grasses is strongly influenced by the available soil N. Application of N fertilizer to grasses usually increases CP concentration as well as crop growth. The majority of the increased CP is non-protein N (NPN) in the form of nitrate and free amino acids (Van Soest, 1982). Crude protein concentrations in annual ryegrass tend to be high (Haby and Robinson, 1997), commonly averaging 150 to 200 g kg -1 with no N applied and increasing to 280 g kg -1 at N rates of 448 kg ha -1 The CP concentrations of ryegrass samples collected in mid-January in different locations in Texas were 228, 321, and 311 g kg -1 for Angleton, Overton, and Knippa, respectively, (Lippke 1993, cited by Lippke and Ellis, 1997). Redfearn et al. (2002) analyzed nutritive value of different cultivars of ryegrass. The CP and IVDOM results in January were 234, 229, and 232 g CP kg -1 and 849, 849, and 846 g IVDOM kg -1 for Gulf, Jackson, and

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15 Marshall, respectively. In general, CP concentrations of cool-season grasses range from 200 to 300 g kg -1 when well fertilized and in vegetative growth stage. Lippke and Evers (1986) reported that ryegrass forage in vegetative stages of growth usually has IVDOM > 700 g kg -1 approaching and sometimes exceeding 800 g kg -1 in the first weeks of the grazing season (Ulyatt, 1981). In vitro digestibility of annual ryegrass grown at different N rates has been evaluated in Louisiana (Allen et al., 1974). The authors reported a 60 g kg -1 increase in digestibility at 448 kg N ha -1 compared with that at 224 kg N ha -1 The cell wall digestibility and leaf/stem ratio of cool-season grasses decreases with maturity. Data from Buxton and Russell (1988) for forages sampled following the beginning of reproductive development indicate a decline in IVDOM from 761 to 552 g kg -1 in orchardgrass (Dactylis glomerata L.) leaves from May to July. Buxton and Merten (1989) reported that herbage IVDOM of four cool-season grass species decreased linearly with time during the spring. Plants with a moderate number of reproductive tillers had an average rate of decline in IVDOM of 3.9 g kg -1 d -1 Grazing Studies Cool-season grasses in the southeastern USA usually are productive from late December to early May. The use of small grains such as oat, wheat, and rye (Secale cereale L.) in mixtures with ryegrass generally provides forage for grazing during the late fall as well as during the winter (Rouquette et al., 1997). Small grains grow better from late December to mid-February and ryegrass results in rapid forage growth during March to late May, often requiring frequent increases in stocking rate to efficiently use ryegrass pastures. According to Rouquette et al. (1997), annual ryegrass is very tolerant of frequent and severe defoliation and often supports 2000 kg ha -1 body weight for a 75to 100-d

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16 period in the spring. Depending upon management and climate, stocker calf gain may range from 450 to 900 kg ha -1 from annual ryegrass pastures during 100 d of grazing. In early studies, Riewe et al. (1963) compared Gulf ryegrass with tall fescue at different stocking rates. The stocking rates ranged from 2.2 to 3.9 steers ha -1 for ryegrass, and 1.9 to 4.9 steers ha -1 for tall fescue. The average daily gains for ryegrass were 0.67, 0.49, and 0.28 kg d -1 respectively, for low, medium, and high stocking rates. Maximum gain per steer and gain per hectare were higher for ryegrass pastures than for tall fescue. Rye is more productive than other annual cool-season grasses on sandy, well drained soil of low fertility, while oat, wheat, and barley (Hordeum vulgare L.) do not grow as well on very sandy soils. Rye produces more forage in fall than spring, while oat DM production is variable (Phillips et al., 1996). Bransby and Gamble (1993) compared ADG and LWG from pastures planted to Bonel rye only or to Bonel rye plus ryegrass. The inclusion of ryegrass resulted in greater stocking rates and ADGs, resulting in greater LWG. At a stocking rate of five, 350-kg steers ha -1 LWG were ~ 330 and 520 kg ha -1 for rye and rye-ryegrass, respectively. In Louisiana, Feazel (1986) evaluated Marshall and Gulf ryegrass in pure stand and an Elbon rye-Gulf ryegrass mixtures. The average stocking rates were similar among the three treatments, 3.5 head ha -1 The LWG ranged from 522 kg ha -1 for heifers grazing Gulf ryegrass to 688 kg ha -1 for Marshall ryegrass. In the 3-yr study, there was no advantage to including Elbon rye in mixture with ryegrass. In North Florida, Bertrand and Dunavan (1974) studied length of the grazing season of calves (170 kg liveweight) grazing ryegrass and ryegrass-triticale (Triticale hexaploide Lart.). The inclusion of ryegrass in the mixtures extended the grazing season from 533 to 650 animal days ha -1

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17 Feazel (1986) evaluated two methods of establishment of annual ryegrass, prepared seedbed and sod-seeded, on performance of beef steers. Steers grazing prepared seedbeds gained 1.05 kg d -1 and those grazing sod-seeded stands gained 1.25 kg d -1 ; however, because initiation of grazing was 36 d later on sod-seeded pastures, total gain per steer was 180 and 157 kg for prepared seedbed and sod-seeded pastures, respectively. In field experiments in Louisiana, rye and ryegrass were sown alone or together into bermudagrass stubble, or into a prepared seedbed. Dry matter forage yields over seven cuts between December and May were ~ 6.0 and 7.3 Mg ha -1 for ryegrass and rye-ryegrass sown into a prepared seedbed and 3.9 and 4.8 Mg ha -1 for ryegrass and rye-ryegrass sown into a bermudagrass stubble (Eichhorn and Venuto, 1999). In some environments, ryegrass has produced as much if not more ADG and LWG when planted alone on prepared seedbed as in mixtures of ryegrass and small grains (Bransby, 1996). Edwards et al. (1978) evaluated various rates of N fertilization (165 to 371 kg N ha -1 ) of Gulf ryegrass and wheat mixtures. Average daily gain was consistent across treatments at 0.82 kg d -1 Total liveweight gain ranged from 622 kg ha -1 on pastures that received 165 kg N ha -1 to 1030 kg ha -1 on pastures that received 371 kg N ha -1 Bransby (1993) compared rye-ryegrass pastures under continuous and rotational stocking at 2.5, 3.7, and 5.0 (270 kg) steer ha -1 stocking rates. Average daily gain for calves on the rotationally-stocked pastures was 0.82, 0.65, and 0.63 kg d -1 respectively, for the three stocking rates compared to 1.05, 0.72, and 0.43 kg d -1 for the continuously stocked pastures, respectively. Thus, an advantage for rotational stocking was obtained only at high stocking rates. In Brazil, Restle et al. (2000) studied beef heifers grazing oat

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18 ryegrass pastures under continuous stocking and fertilized with 200 kg N ha -1 Average daily gain and gain ha -1 were 0.58 kg d -1 and 429 kg ha -1 respectively. Results of a 5-yr study were presented by Rouquette et al. (1992) who compared a range of stocking rates on pure stands of ryegrass plus N and ryegrass-arrowleaf clover (Trifolium vesiculosum Savi.), sod seeded on Coastal bermudagrass pastures. Cow-calf pairs grazed the pastures. Average daily gains of suckling calves were similar among treatments and ranged from 1.0 to 1.4 kg d -1 for ryegrass and arrowleaf pastures at stocking rates of 5.98 to 2.37 animal units (AU) (680-kg liveweight) ha -1 The most significant responses to the use of cool-season forages are not only calf ADG but also weaning weight of fall-born calves. Using a fall calving season, the stocker phase is essentially eliminated because calves are heavy enough to enter the feedlot immediately upon weaning (Rouquette et al., 1997). After an extensive research program with cool-season grasses, Rouquette et al. (1997) concluded that achieving economic optimum grazing management and use of cool-season grasses is not an easy task. A knowledge base of forage growth expectations and the art of managing proper defoliation regimes will allow for the greatest opportunity for positive economic returns and an acceptable transition from cool-season to warm-season pastures. Supplementation Flexible management of the grazing system is required to maximize animal performance since forage production and nutritive value are not uniform throughout the year. The use of supplementary feed can overcome the shortfalls of the forage component. When energy and protein requirements increase due to lactation, pregnancy, and growth, part of the roughage component of the diet may need to be replaced by

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19 concentrates (Fontaneli, 1999). Concentrates generally are more digestible than forages and have much higher fermentation rates. According to Stockdale et al. (1987), several factors may affect the response to supplement, including quality of the pasture and supplement, amount of pasture and supplement fed, and degree to which supplemental feeds replace pasture intake. Protein Crude protein concentration of high nutritive value herbage may exceed 250 g kg -1 however, 650 to 850 g kg -1 of total protein is degradable in the rumen, with 150 to 350 g kg -1 escaping the rumen. Microbial protein is a major protein source for ruminants; therefore, optimizing microbial protein synthesis is essential. In a review of protein supplementation of grazing livestock, Petersen (1987) stated that enhanced forage utilization occurs in several ways. Protein supplements provide amino acids, carbon skeletons, and minerals that help satisfy microbial requirements, thereby increasing microbial growth and/or fermentation. These factors combine to increase intake through increased microbial activity and rate of passage (Allison, 1985). In addition to increasing available nutrients in the rumen, protein supplements may also increase the quantity of protein reaching the small intestine through undegraded or bypass protein. Non-structural carbohydrates are needed to utilize the forage N in the rumen, to reduce energy cost of excreting N, and to supply nutrients to the small intestine. Microbial protein synthesis can become a performance-limiting factor for ruminants consuming highly digestible ryegrass diets. Hill (1991) observed that in young calves grazing ryegrass, synthesis of microbial protein was 6 to 10 g 100 g -1 of OM digested in the rumen, and energy/protein supplements increased the flow of microbial and undegraded dietary protein to the intestines, and increased forage intake proportionally. In a study with beef steers grazing

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20 rye-ryegrass pasture and receiving free-choice mineral, Grigsby et al. (1991) compared corn meal and fish meal concentrates. Average daily gain was 1.5 and 1.0 kg d -1 for corn meal and fish meal, respectively. Supplemental energy may have provided a source of rapidly fermentable carbohydrate, which was well synchronized with ammonia and peptide production from forage protein degradation, which in turn may have resulted in a greater synthesis of microbial protein. Moore et al. (1999) suggested that the ratio of digestible organic matter (DOM) to CP should be no more than seven to one based on the observation that animals consuming forages with ratios greater than seven were likely to respond to CP supplement. Hill et al. (1990) studied beef heifers grazing ryegrass pastures (243 g CP kg -1 ) and supplemented with maize, cottonseed meal, blood meal, feather meal, fishmeal, or condensed molasses with cottonseed meal. Meal-based supplements were given at 3.5 g kg -1 of body weight (DM basis). Forage organic matter intake (OMI) (650 g kg -1 digestible) was increased by blood and fishmeal supplements and decreased by cottonseed and feather meal. It was concluded that blood and fish meal seemed to be more effective in supplying required protein and essential amino acids that stimulate intake of forage. In C4 grasses, protein may be excessive or seriously deficient, depending on the pasture species, season, and maturity of growth (Poppi and McLennan, 1995). Minson (1990) compiled a series of studies in which CP was below 62 g kg -1 in the base forage. There was an average 40% increase in intake due to protein supplement and a 34% increase due to supplementary urea. Probably, the greatest and most uniform response to supplemental N occurs when the CP concentration of the base forage is below 70 g kg -1

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21 (Coleman et al., 2004). Bodine et al. (1999) reported the effect of four sources of ruminal degradable protein and two sources of energy supplementation on steers consuming low-quality hay (< 70 g CP kg -1 OM). Forage OMI and total OMI increased quadratically as rumen degradable protein increased for both energy supplements. When DOM:CP ratios of limpograss [Hemarthria altissima (Poir.) Stapf and Hubb.] were between 8 and 10, cattle responses to CP supplementation were significant (Lima et al., 1999b). Hammond et al. (1994) suggested assessing blood urea N (BUN) concentrations as a method to monitor the need for supplements by grazing animals when forage nutritive value data are not available. Cattle BUN concentrations from 9 to 12 mg dL -1 are considered to be a transition range below which gain response to protein supplementation has been positive. Hoffman et al. (1993a) reported a summary of BUN values from several studies with dairy cows. There was an apparent excess of N in the rumen in relation to carbohydrates and wastage of highly degradable N in pasture forage when BUN values ranged from 18.5 to 28.6 mg dL -1 Newman et al. (2002b) observed higher BUN concentrations (18 vs. 15 mg dL -1 ) on heifers grazing limpograss either with or without CP supplementation, respectively. Energy Energy supplements for forages generally fall into three categories: starch, sugars, and fiber (Poppi and McLennan, 1995). Sources of fibrous highly-digestible energy, such as soybean hulls or wheat middlings, have shown the most consistent response, presumably because of the good synchrony with NH 3 released (Poppi and McLennan, 1995). Johnson et al. (2001a) supplemented mature cows on bermudagrass pastures with soyhulls or corn at 0.17% BW. There was no difference in dry matter intake (DMI) or ADG between the treatments. These results suggest that low levels of corn

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22 supplementation did not affect forage intake and animal performance. Supplementing forage with a starch source, especially at levels > 250 g kg -1 of the total DM tends to reduce fermentation of the basal forage [Goetsch et al. (1991), cited by Coleman et al. (2004)]. According to Fieser and Vanzant (2004), when fed at similar levels of OM, soybean hull supplementation provided an average of 60 g kg -1 greater DOM intake than corn supplementation. Ordonez-Tercero et al. (2003) used two energy sources [sugar-cane (Sacharum officinarum L.), molasses and maize] to test the effect of energy supplementation on intake and digestibility of the basal diet (elephantgrass) [Pennisetum purpureum Schum.] and a foliage mix of Brosimum alicastrum Sw.and leucena [Leucaena leucocephala (Lam). Dewit] There were no differences in basal diet intake (5.54 kg DM d -1 ) among treatments. The potentially degradable DM (337 g kg -1 ) and OM (364 g kg -1 ) fractions were not affected by energy supplementation. Frizzo et al. (2003) studied the effect of levels of energy supplementation on the productive and reproductive performance of Charolais heifers maintained in a cultivated pasture of black oat (Avena strigosa Chreb.) [Avena nuda L.]) plus annual ryegrass. It was shown that supplementation increased ADG, stocking rate, and LWG. Heifers kept only on pasture had lower body condition and showed lower estrus percentage than heifers supplemented with 0.7 and 1.4% of LWG d -1 The estimated intake of DM was lower for the 1.4% BW d -1 supplementation level. In South Brazil, Rocha et al. (2003) evaluated performance of beef heifers grazing oat-annual ryegrass mixtures with and without ground maize supplement. The utilization of oat and ryegrass pasture with supplementation resulted in higher ADG, stocking rate, and LWG than the mixture with no supplement. Bertrand and Dunavan (1977) studied the

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23 effects of increasing energy supplementation from 0 to 15 g kg -1 BW on performance of beef steers (180 kg liveweight) grazing ryegrass-rye-white clover (Trifolium repens L.). The increase in supplementation levels increased ADG (0.89-1.10 kg d -1 ), stocking rate (4.09-5.32 head ha -1 ), and LWG (582-936 kg). Interactions between Supplements and Forages When forages are offered free choice and supplemental concentrates are offered in restricted amounts, forage DMI may either increase, decrease, or remain the same (Moore, 1992). In many cases, animal responses to supplements are either greater or less than expected. The deviations between expected and observed performance are usually explained by associative effects of supplements upon voluntary intake and digestibility of the total diet (Moore et al., 1999). In general, concentrates will decrease forage intake when forage quality is high, other nutrients are in balance with energy, and concentrate is fed in large amounts. Large differences in substitution rates have been reported and the effects have greater relation to differences among forages rather than to differences among concentrates (Waldo, 1986). On the other hand, small amounts of concentrate may increase voluntary intake when forages nutritive value is low, especially when the forage has a high ratio of TDN to CP (Moore, 1994). Sarker and Holmes (1974) fed supplement in increments of 2, 4, 6, or 8 kg OM d -1 to non-lactating cows grazing ryegrass. Though total OMI increased with increasing amounts of concentrate, the average increase in forage intake was 0.46 kg OM kg -1 of concentrate OM fed. Meijs (1986) fed high-starch supplements or high fiber supplements to cows grazing predominantly perennial ryegrass (Lolium perenne L.) swards. Supplement intake was 5.5 and 5.3 kg OM d -1 with forage intakes of 11.5 and 12.6 kg OM d -1 for high and low starch treatments, respectively. Average forage substitution rate

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24 for animals receiving supplements was 0.45 vs. 0.21 kg herbage kg -1 concentrate for animals receiving starch and fibrous supplements, respectively. Royes et al. (2001) evaluated different sources of energy (maize, sugar cane molasses, or soybean hulls) and feeding rates (0, 1.4, or 2.8 kg DM steer -1 d -1 in growth trials; 0, 150, or 300g kg -1 of the ration DM in digestion trials) of supplements fed to cattle receiving ammoniated stargrass hay. Increasing the level of supplementation decreased hay intake but increased total dietary intake for all diets. Daily gain and feed efficiency of steers improved with supplementation. Steers supplemented with corn or soybean hulls at 2.8 kg DM d -1 had a higher ADG (0.92 kg) and gain:feed (0.103) than steers supplemented with molasses (0.78 kg) at the same level. According to Galloway et al. (1992), moderate dietary levels of supplement (200-300 g kg -1 ) can improve nutrient intake and performance by cattle consuming bermudagrass. At greater amounts, nutrient digestion, intake, or both, of the forage portion of the diet can be affected negatively. Cattle grazing forages with DOM:CP ratios greater than seven to eight are likely to have inadequate ruminally degradable protein and should respond to N supplemention by increasing intake and animal performance (Moore and Kunkle, 1999). Wheeler et al. (2002) tested the effects of increasing supplement protein concentration on performance and forage intake of beef steers consuming bermudagrass forage. Treatments were no supplement or daily equivalents of 0.2, 0.4, and 0.6 g of supplemental protein kg -1 of BW. Forage intake increased 16% and total OMI increased 30% in supplemented compared to unsupplemented steers. Diet OM digestibility increased 145 g kg -1 in supplemented compared to unsupplemented steers.

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25 A special case of substitution and associative effects is the use of total mixed rations (TMR) for lactating dairy cows. Such diets may include several sources of forage and combinations of concentrates. Because TMRs often are based on high quality forage and include high percentages of concentrates, intake and digestibility of the forage component is very likely less than expected when forage is fed alone (Moore, 1994). Fike et al. (2003) studied the effects of two levels of supplementation on forage and total intake of lactating dairy cows grazing bermudagrass and rhizoma peanut (Arachis glabrata Benth.) pastures. The substitution of forage OM by supplement OM (kg kg -1 ) was 0.48 for rhizoma peanut and 0.06 for bermudagrass. Feeding additional supplement increased total OMI by 230 and 9 g kg -1 for cows grazing bermudagrass and rhizoma peanuts pastures, respectively. Behavior According to Stobbs (1975), the energy required to prehend forage from the standing biomass is an insignificant part of the maintenance requirements, but energetic costs of activities associated with grazing have been estimated to account for 25 to 50% of an animals daily energy requirements. Numerous studies have shown that supplying supplemental protein and (or) energy to cattle consuming lowto moderate-quality forages can increase animal performance and forage OMI, however, data are limited showing the effect of supplementation on grazing behavior (Krysl and Hess, 1993). Adams (1985) postulated that disruption of normal grazing activity resulting from supplementation regimens could adversely affect forage intake and animal performance. In research with yearling beef steers grazing ryegrass, the same authors noticed that although there was an 11.3% decrease in forage intake with supplementation, total daily grazing time was not altered. Macoon (1999) observed that dairy cows supplemented

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26 with high levels of concentrate spent less time grazing during daylight (135 min) than cows that received low levels of concentrate (180 min). Supplementation can cause shifts in daylight grazing behavior patterns; however, there is no indication that supplemented animals alter their percentage of daytime vs. nighttime grazing compared to unsupplemented animals (Adams, 1985). Providing protein supplements to cattle grazing low-quality forages increased harvest efficiency (forage OMI kg -1 BW min -1 spent grazing) from 8 to 60% compared with unsupplemented cattle (Barton et al., 1992). In contrast to protein meal-based supplements, high-starch supplements either failed to alter or decreased harvest efficiency (Adams, 1985). Allden and Whittaker (1970) indicated an upper limit of grazing time of ~ 600 to 720 min d -1 for sheep. Similarly, Stobbs (1975) observed that fatigue limits the time that can be spent grazing to ~ 720 min d -1 in cattle. Numerous studies have reported daily grazing time to be in the range from 359 to 771 min d -1 for cattle over a wide range of environmental temperatures, supplementation regimens, grazing management practices, and forage types (Krysl and Hess, 1993). Intake Intake is the most important determinant of performance by grazing ruminants (Lippke, 2002). As much as 60 to 90% of the variation in digestible energy intake may be due to animal variability, with 10 to 40% due to diet digestibility (Crampton et al., 1960). The most important mechanisms controlling intake in ruminants are metabolic, distention, and behavioral. Conrad et al. (1964) examined results from 114 trials with lactating cows and reported the situations where physical and physiological factors were regulating feed intake. Intake of diets having digestibility between 500 and 670 g kg -1 was thought to be limited by physical factors such as digestibility of the feed and its rate

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27 of passage through the digestive tract. Intake of diets having a digestibility > 670 g kg -1 was limited primarily by physiological control mechanisms. Intake of perennial warm-season grasses is considered to be limited by physical (distention) mechanisms due to their high fiber concentration and low digestibility. Van Soest (1965) reported negative correlations between cell wall concentration and voluntary intake. Factors controlling ruminant intake should be assumed to function through multiple interactions, and the simplistic view should be abandoned that only physical factors limit the intake of roughage diets and that only chemical or physiological factors limit intake of concentrate diets (Fisher, 2001; Moore, 1994). Hodgson (1982) stated that under grazing conditions, ingestive behavior (i.e., bite mass, bite rate, and grazing time) might be a more important determinant of forage intake than physical and metabolic controls. Low forage availability and accessibility are situations where ingestive behavior may be an important controlling factor on forage intake. The DMI on pastures is a result of a complex interaction among pasture canopy characteristics, ingestive behavior, and grazing time (Burns and Sollenberger, 2002). Forage intake is forage species-specific and altered by plant maturity and morphology. The eating behavior of ruminants has been clearly characterized as selective, with strong preference for green leaf and against dead and stem tissues (Minson, 1990). Greater selectivity by the animal for leafy tissue from within the bounds of forage on offer will generally result in a higher quality diet than that on offer and greater DMI (Burns et al., 1991). Measurements of Forage Intake of Grazing Animals The ability to measure or predict DMI and nutritive value of forages on pastures has been the impetus for studies of the plant-animal interface (Burns and Sollenberger,

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28 2002). A number of techniques have been adopted to estimate intake because it is not practical to measure intake on pastures directly (Lippke, 2002). There are two general approaches to the indirect estimation of pasture intake. According to Moore and Sollenberger (1997) the techniques may be classified as follows: 1) estimates for individual animals by fecal output and forage digestibility or ingestive behavior, and 2) estimates for groups of animals by disappearance of herbage mass or calculation from animal performance. Each method employs different assumptions which must be met if the estimates are to be valid. To avoid errors associated with the estimation of forage intake on pastures, researchers have cut and carried green pasture herbage to confined animals kept in confinement. Though this approach affords a great degree of precision, it may be highly inaccurate because it reduces the opportunity for diet selection. Experimental results are likely most affected when swards are highly heterogeneous, and when environmental factors, sward density, and forage accessibility have large effects on grazing behavior. Intake calculations based on fecal output and forage digestibility have been used extensively. Fecal output is the measure of interest because it can be used to calculate intake using the following equation: Intake = fecal output / (100 diet digestibility). Early attempts to estimate intake from total feces collection in individual animals included use of fecal collection bags. In addition to the potential for loss or urine contamination due to poor design or lack of fit, the bags also have the potential to stress the animal and to alter intake by changing grazing behavior. The alternative is to estimate fecal output using marker technology. Functionally, markers are classified as being internal to the feedstuff or external, i.e., added to the

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29 feedstuff or dosed separately to the animal (Lippke, 2002). Markers are reference compounds used to investigate both chemical and physical digestive processes (Owens and Hanson, 1992). Characteristics of an ideal external marker were outlined by Owens and Hanson (1992) and include the following traits: 1) it should be unabsorbable, 2) it should not affect or be affected by animal or microbial digestive processes, 3) its flow should closely mimic that of the material it marks, and 4) it must be analyzable with a specific and sensitive methodology. No single marker currently meets all of these criteria. Chromium oxide has been the most widely used external marker for estimation of intake during the last 50 yr (Huston et al., 1999). The primary problem associated with this marker is that it moves through the digestive tract independently of the undigested particles of the diet, and fecal concentrations of chromium may exhibit strong diurnal variation. Measuring daily DM intake with Cr requires a continuous period of Cr intake and frequent spot feces collection during different hours on the days following dosing. This additional handling of the animals is undesirable, particularly when it has potential to disturb established patterns of grazing behavior. An alternative method of calculating fecal output from a single dose of Cr has been presented by (France et al., 1988). Animals are dosed once with labeled feed fractions, and numerous fecal samples are collected over a period of time long enough for the label source to clear the animal. A nonlinear equation relating time after dosing to fecal Cr concentration is used to generate parameters for the estimation of fecal output (Pond et al., 1989). According to Buntinx et al. (1992), controlled-release Cr capsules offer the potential of reduced labor, coupled with the possibility of preventing adverse effects on

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30 animal behavior from repeated daily dosing, or from the presence of fecal bags during total fecal collection. Controlled-release devices greatly reduce the within-day coefficient of variation for fecal concentration of Cr in sheep and cattle when compared with daily dosing (Adams et al., 1991). However, total fecal collection must be used on a subset of animals to correct for possible payout differences from manufacturer specifications and/or subsampling influences on chromium recovery. Dosed and plant alkanes are options as external and internal markers, respectively. Mayes et al. (1984) showed that dosed even-chain length n-alkanes and naturally occurring odd-chain plant n-alkanes could be used as fecal markers to determine forage intake. Forage species contain variable quantities of alkanes, and the concentration of odd-chain alkanes may be too low in some tropical forages for them to be used as an internal marker (Laredo et al., 1991). The problems associated with species-specific alkane concentrations and analytical determination of the n-alkanes make the technique less likely to be used in a wide range of environments. Associated with the use of inert markers in estimating DM intake of grazing animals is the requirement of knowing the digestibility of the animals diet. Obtaining samples that represent the diet of the dosed animals can frequently be the major factor limiting the accuracy of the method. Surgically altered animals and "hand plucked" samples are the two most-used methods to obtain samples of the diet in grazing studies. After a representative sample is collected, the accuracy of the marker technique resides with the method used to estimate the absolute digestibility of the diet (Burns et al., 1991). In vitro dry matter digestibility has been the most-used method because it is cheap, precise, and in most of the cases, well correlated with in vivo digestibility. Although in

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31 vitro dry matter digestibility (IVDMD) has perhaps been most widely used to predict digestible DM, a variety of internal markers also have been used. Ingestive behavior has been suggested as a method of estimating daily DM intake (Coleman et al., 1989). The parameters to be measured are bite mass, bite rate, and grazing time. Mechanical methods for short-interval measurements have been reviewed, but recent advances have occurred through automation to aid quantification. Bite rate may be recorded by observing, listening, or by electronic devices, however, it is necessary to discriminate between biting and non-biting jaw movements. Artificial swards can be used in measurement of bite weight, or esophageal extrusa can be collected over a short period of time. Finally, grazing time can be recorded visually or by using electronic devices. Errors associated with these devices and especially with extrapolation of intake during short measurement periods to daily intake are the most common. If the pastures are uniform and under intensive management, forage disappearance can provide a reasonable estimation of forage intake. Forage mass is measured before and after grazing, and the forage that disappears is considered to have been consumed. The technique is very labor intensive because a large number of sites have to be measured to provide an acceptable estimation of herbage mass. The difficulty of achieving accurate estimations of herbage mass, as well as trampling, defoliation by insects, and senescence, make the herbage disappearance technique not reliable enough to be recommended for routine estimates of pasture intake (Moore and Sollenberger, 1997). The animal performance technique, also called "reverse energy calculations", is the calculation of intake by using, in reverse, accepted energy requirements for maintenance, growth, and lactation (McMeniman, 1997). For a given rate of performance, the daily

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32 amount of energy that must be consumed in order to achieve that level of performance can be obtained from tables and equations. Accurate determinations of changes in body weight are required when using this technique. Digestibility is the main determinant of forage energy concentration and efficiency of energy utilization (Moore and Sollenberger, 1997). Digestibility of the forage can be obtained by the methods reported before, and concentration of digestible energy (DE) is calculated by assuming 4.4 Mcal DE kg -1 DOM. Concentration of metabolisable energy (ME) is calculated by multiplying DE concentration by 0.82 and net energy for maintenance and for gain are calculated from ME concentration by (NRC, 1996). The main constraint of this technique is that factors other than forage energy concentration, such as nutrient imbalances and environmental effects may influence the efficiency of conversion of net energy for gain to animal product. N Fractioning and in situ Methodology Diet formulation systems have been developed that attempt to balance diets for both ruminal protein degradability and carbohydrate availability with the goals of maximizing microbial protein synthesis and minimizing protein wastage as ammonia (Nocek and Russell, 1988). Rumen protein degradability is one of the most important qualitative factors determining the protein value of a feed. Protein degradability determines the supply to the rumen microbes of both ammonia and branched-chain amino acids, as well as the supply to the small intestine of the rumen-undegraded feed protein as a potential source for amino acid absorption. The protein concentration in feeds may be expressed on a CP basis. The CP is composed of two fractions: rumen degradable protein (RDP) and rumen undegradable protein (RUP). The RDP consists of non-protein N and true protein while the RUP is

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33 formed only by true protein. The protein degradation rate will vary depending on the proportions of NPN, true protein that degrades at variables rates, and unavailable protein (Pichard and Van Soest, 1977). In situ Procedure Numerous methods currently are available for estimating ruminal DM and N disappearance of feedstuffs. The in situ technique involves the placement of indigestible bags containing a feedstuff into the rumen of a fistulated animal for various time intervals and the measurement of the amount of DM and N that disappears from the bags over time (Stern and Satter, 1984). According to Nocek (1988) there is no better way to simulate the rumen environment within a given feeding regime, although in the ruminal environment, the feed sample in the bag is not subject to the total ruminal experience: i.e.; mastication, rumination, and passage. The in situ procedure can be used to quantify the protein fractions, the instantly degradable Fraction A; the undegradable fraction after an extended rumen incubation, Fraction C; and Fraction B, the slowly degradable fraction, obtained by difference (%Fraction B= 100 % Fraction A % Fraction C). Although the technique allows the quantification of the different N fractions of the forage as well as the rate of digestion of B, it cannot be used to quantify the rate at which A is degraded. Nocek (1985) demonstrated that a certain portion of the test feed escapes the bag prior to ruminal degradation. This fraction is generally assumed to be readily available to rumen microbes and digested at a rapid rate. Potentially degradable nutrient fractions have been described by first-order kinetic rate constants (Mertens, 1973). Primary assumptions are that the pools in question are homogeneous and that the substrate remaining will be degraded as a linear function of time in the rumen. Orskov and McDonald (1979) and other researchers

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34 have developed mathematical models to fit the estimated ruminal degradability of feedstuffs. The most common approach included the use of non-linear regression. Mertens and Loften (1980) proposed the inclusion of lag time for protein degradation in the models used for fitting N disappearance. If a delay appears at the beginning of the disappearance of Fraction B, a lag component should be included in the model (McDonald, 1981). Various aspects of the in situ technique interact and can influence the interpretation of the results. Bag porosity is a compromise between limiting the influx of rumen contents not associated with the test feed and allowing influx of microbes to degrade the test feed, while at the same time limiting the efflux of undegradable feed particles (Nocek, 1988). Uden and Van Soest (1984) showed cell wall digestion of timothy (Phleum pratense L.) increased with increasing bag pore size (20, 37, and 53 m). Digestion of DM was similar for pore size ranging from 40 to 102 m, and estimated rumen N availability was greater for bags with pore size of 6 or 20 m (Uden and Van Soest, 1984). Feed particle size is another important aspect in the in situ disappearance determination. Generally, longer and coarser material is associated with slower rates of digestion and greater variation. However, finely ground materials are subject to greater mechanical losses from the bags (Nocek, 1988). The same author states different feedstuffs can require different particle sizes, and the particle size for cereal grains, fibrous products, forage, and silages ranges from 2 to 5 mm. Sample weight is also an important consideration. The optimum sample weight is that which provides enough residue at the end of the extended rumen incubation for

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35 chemical analysis without over-filling the bag so as delay bacterial attachment, increase lag time, and underestimate digestion rates (Nocek, 1988). For forages, Van Hellen and Ellis (1977) recommends 10 mg cm -2 of bag surface area, whereas Uden and Van Soest (1984) recommend 6 to 7 mg cm -2 The host animal diet is the major factor determining quantity and types of microbes and therefore the rate and extent of digestion of dietary nutrients. de Faria and Huber (1984) evaluated in situ DM digestion of corn silage, alfalfa (Medicago sativa L.) silage, and grass hay in diets containing three different protein (81, 113, 133 g kg -1 ) and three different acid detergent fiber (ADF) concentrations (390, 299, and 210 g kg -1 ). Neither protein nor ADF of the host animals diet had a significant effect on DM digestibility for any forage tested, however, specific sampling time by forage interaction was noted. The animal species most commonly used for in situ experiments are cattle and sheep. Siddons and Paradine (1983) compared sheep and steers fed similar diets at maintenance. Sheep had higher ruminal ammonia N than steers, lower VFA, and similar ruminal pH and rumen dilution rate. In situ Disappearance of Warmand Cool-season Grasses High quality cool-season pastures and many tropical legumes would be expected to have high ruminal ammonia losses when their CP concentrations exceeded 150 g kg -1 because of the high concentration of ruminal degradable protein (Poppi and McLennan, 1995). According to Beever (1984), 800 to 900 g kg -1 of ryegrass CP can be degradable in the rumen. Van Vuuren et al. (1991) stated that OM and CP in situ total degradability decreased with increasing ryegrass sward maturity and with decreasing rate of N application. Salaun et al. (1999) studied the relation between N fertilization and rumen CP degradability of perennial ryegrass. They concluded that reducing N fertilizer

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36 application decreased the theoretical degradability in the rumen due to decrease in the NPN. The total N disappearance was 600 and 710 g kg -1 for plots fertilized with 250 and 550 kg N ha -1 Assis et al. (1999) did not find significant difference of CP Fraction A in Tifton 85 bermudagrass with levels of N fertilization from 0 to 400 kg ha -1 Yan and Agnew (2004) tested the effective degradability of perennial ryegrass N assuming a ruminal outflow rate of 0.02, 0.05, or 0.08 % h -1 The objective was to use these data to develop prediction equations for N degradability in C3 grass silages. At 0.02 % h -1 ruminal outflow, total N degradability was 934 g kg -1 The N degradability was negatively related to ADF, NDF, and lignin concentrations but positively related to CP concentration. Warm-season grass CP tends to be more slowly degraded in the rumen than that of cool-season grasses (Minson, 1990). Akin and Burdick (1975) concluded that digestibility differences between C3 and C4 grasses were associated with the parenchyma bundle sheath (PBS) of C4 grasses. The C4 plants have a lower soluble protein concentration (260 to 300 g kg -1 ) than C3 plants (330 to 480 g kg -1 ), which is thought to be mainly due to differences in Rubisco concentration and location. Further, PBS tissue from C4 plants, the primary location for Rubisco, is slowly or incompletely digested in the rumen (Akin and Burdick, 1975). Redfearn et al. (1995) studied the rates of ruminal protein disappearance for switchgrass (Panicum virgatum L.), big bluestem (Andropogon gerardii Vitman), and smooth bromegrass (Bromus inermis Leyss.) and suggested that generalizations regarding ruminal protein degradability should not be made among forage species. The total protein disappearance was 847, 795, and 974 mg g -1 for switchgrass, big bluestem, and smooth

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37 bromegrass, respectively. Mandebvu et al. (1999) compared in situ disappearance of Tifton 85 and coastal bermudagrass. There was no difference in the potentially digestible fractions (A+B) among the cultivars with approximate values of 600 g kg -1 Assis et al. (1999) evaluated the in situ N disappearance of Cynodon spp, Tifton 44 and Tifton 85 bermudagrasses, and Puerto Rico stargrass. Tifton 85 had the smallest concentrations of the potentially digestible CP fractions (A+B), but it had the largest B fraction concentration. Johnson et al. (2001b) reported that with no fertilization, the Fraction A concentration was 212 (bahiagrass), 310 (bermudagrass), and 279 g kg -1 (stargrass) of the total N, but upon application of 160 kg N ha -1 per cutting, Fraction A represented 285 (bahiagrass), 400 (bermudagrass), and 421 g kg -1 (stargrass) of the total N. Newman et al. (2002a) fractionated the N content of limpograss at different grazing heights. Pastures grazed to 20 cm had the greatest proportion of soluble N and the least portion of undegradable CP. They concluded that low total CP in the DM, resulting in low concentrations of CP Fractions A and B in the rumen, and the long lag phase for degradation of the B fraction, may contribute to reported protein deficiencies of cattle grazing limpograss. Other Methods to Estimate Ruminal Protein Degradation Although in vivo and in situ procedures for estimating degradability of protein are not without drawbacks, they are typically considered to most accurately reflect reality. However, labor requirements, cost, time, variability, and the need for access to fistulated ruminants limit the feasibility of widespread use of these procedures for routine analysis (Mathis et al., 2001). In vitro estimation of rumen-degradable protein is appealing because it addresses many of the limitations of in situ and in vivo procedures and would be relatively easy to standardize (Stern et al., 1997). While numerous in vitro methods are

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38 currently available for estimating ruminal protein degradation of feedstuffs, assays based on Streptomyces griseus are probably the most commonly used (Roe et al., 1991). In the Streptomyces griseus method, feeds are incubated in a borate-phosphate buffer at pH 7.8 to 8.0 to facilitate optimum enzyme activity, and enzyme is added at a level of 6.6 units of enzyme g -1 sample DM (Krishnamoorthy et al., 1983). Streptomyces griseus is a broad spectrum, commercially available protease that has both exopeptidase and endopeptidase activity, as in the rumen. Ruminal protein degradation also has been estimated by the ficin method, which breaks down protein by proteolytic ficin (Ficus glabrata) used at 8.24 units g -1 of sample DM in a phosphate buffer at pH 6.5 (Poos-Floyd et al., 1985). Finally, the neutral protease with amylase method makes use of an endopeptidase, Bacillus subtilus, at 9.6 units g -1 sample DM, in a sodium citrate buffer, in addition to an enzyme that contains endo--glucanase and -amylase activities (Assounami et al., 1990). Roe et al. (1991) compared the three in vitro methods of ruminal protein degradability with the in situ methodology. None of them resulted in feed protein degradation curves that had a consistent relationship with those generated by the in situ technique. However, relationships were found between protein degradability estimates obtained by the neutral proteases with the amylase method at specific time points and those measured in situ.

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CHAPTER 3 SUPPLEMENTATION EFFECTS ON FORAGE CHARACTERISTICS AND PERFORMANCE OF EARLY WEANED CALVES GRAZING RYE-RYEGRASS PASTURES Introduction Grassland covers ~ 4.5 million ha in Florida and most of this area is utilized by beef cattle. Floridas beef cattle industry is based on cow-calf enterprises with a total of 950 000 beef cows. Sales of cows and calves accounted for $348 million of income to Florida farmers and ranchers in 2003 (Florida Agriculture Statistics, 2004). The profitability of the Florida cow-calf enterprise is largely dependent on the cows productivity. The combination of low quality warm-season forages and forage quantity limitations during winter reduce cow reproductive performance. Early weaning the calves of young cows and first-calf heifers is a management strategy that has potential to address this problem. At early weaning, cow requirements for total digestible nutrients (TDN) decreases by as much as 49%. As a result they are able to restore body condition rapidly (Arthington and Kalmbacher, 2003). Heifers whose calves were weaned early had higher pregnancy rates (94%) than those whose calves were weaned at the normal time (65%) during a 2-yr study in South Florida (Arthington and Kalmbacher, 2003). Although the benefits of early weaning, i.e., improving reproduction and reducing nutrient requirements of the cow, have been recognized for many years, the factor limiting practical application of early weaning has been management of the early weaned calves. Mild winters in the southern USA offer an opportunity to raise calves on pasture systems that include high nutritive value winter-annual forages (Arthington and 39

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40 Kalmbacher, 2002). Annual ryegrass (Lolium multiflorum Lam.), a high-yielding, high quality grass, is the most commonly grown cool-season pasture forage in the southern and southeastern USA (Evers et al., 1997). The use of small grains such as oat (Avena sativa L.), wheat (Triticum aestivum L.), and rye (Secale cereale L.) in mixtures with ryegrass generally extends the grazing season (Rouquette et al., 1997). Small grains grow better from late December to mid-February, and ryegrass is most productive during March to late May. The practice of raising early weaned calves on winter pastures is a feasible alternative, especially if pasture quality and quantity are superior to hay available for winter feeding (Barnes et al., 1996). Weder (1997) reported average daily gain of 0.8 kg d -1 for early weaned calves grazing annual ryegrass, however, the limited ruminal capacity of early weaned calves at young ages may limit forage intake and result in low animal performance. Early weaned calves grazing wheat pastures consumed 27% less forage dry matter (DM) during the first 20 d exposed to pastures than during the subsequent 50 d (Paisley et al., 1998). The authors suggested that initial performance of early weaned calves was limited by low forage intake. Moreover, calves grazing high quality cool-season pasture may have high ruminal ammonia losses, if forage crude protein (CP) concentrations exceed 150 g kg -1 (Poppi and McLennan, 1995). Loss of N from the rumen is costly due to significant energetic expenditure associated with urea synthesis and excretion. Non-structural carbohydrates are needed to utilize the forage N, to reduce energy cost of excreting N, and to supply nutrients to the small intestine. Microbial protein synthesis can become a performance-limiting factor for ruminants consuming ryegrass diets. Rocha et al. (2003) evaluated beef heifers grazing oat-annual

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41 ryegrass with and without ground maize (Zea mays L.) supplement. Supplementation resulted in higher average daily gain, stocking rate, and production per unit of land area. An effective supplementation program is necessary to overcome potentially low forage intake and maximize the efficiency of utilization of cool-season grasses in early weaned calf systems. The objective of this study was to evaluate the effect of different levels of concentrate on performance, stocking rate, gain per hectare, and forage intake of early weaned calves grazing rye-ryegrass pastures in North-Central Florida. Material and Methods The study was located at the Beef Research Unit, 18 km northeast of Gainesville, FL (30 o N). Data were collected during the 2003 and 2004 winter-spring seasons. The soils at the research site are Adamsville fine sand (Aquic Quartzipsamments, hyperthermic, uncoated) and Pomona sand (sandy, siliceous, hyperthermic Ultic Alaquod). These soils have from poor to somewhat poor drainage with rapid permeability. Prior to initiation of winter grazing, mean soil pH (1:2 soil:deionized H 2 O ratio) was 6.5, and Mehlich-I (0.05 M HCl + 0.0125 M H 2 SO 4 ) extractable P, K, Mg, and Ca in the Ap1 horizon (0to 15-cm depth) were 52, 19, 117, 761 mg kg -1 respectively. Starting dates for the trial were 28 Jan. 2003 and 15 Jan. 2004. Grazing ended on 14 May 2003 and 29 Apr. 2004. Monthly rainfall totals and temperature data during the trial are presented in Table 1 (Appendix A). In early October 2003 and 2004, the area was sprayed with 3.0 L ha -1 of glyphosate (Monsanto Co., St. Louis, MO; isopropylamine salt [10 g kg -1 ] of N-phosphonometyl glycine) to kill bahiagrass (Paspalum notatum Flgge). Three weeks later, the cool-season forages were planted using a no-till drill. The seeding rates were 20 kg ha -1 of Jumbo ryegrass and 80 kg ha -1 of a mixed-blend rye (Grazemaster). All pastures

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42 received an initial application of 40 kg N ha -1 17 kg P ha -1 and 66 kg K ha -1 3 wk after planting. Additional applications of 40 kg N ha -1 were made in late December, early February, and early March in both years. Calves were weaned on 2 Jan. 2003 and 5 Jan. 2004 at ~90 d of age and 100 kg of body weight (BW). They were held in dry lot for 7 d with access to bermudagrass [Cynodon dactylon L. (Pers.)] hay (ad libitum), 1 kg of pre-conditioning medicated concentrate d -1 and water. Early weaned calves were Angus-sired (crossbred-cows sired by Angus bulls) products from primiparous and multiparous cows in 2003 and 2004, respectively. Calves were dewormed with Ivermectin 10 g kg -1 concentration (Ivomec, Merck & Company, Rahway, New Jersey, USA) on 10 Apr. 2003, and 12 Feb. and 29 Apr. 2004. Pasture size was 0.2 ha, and each pasture was subdivided into four paddocks. Pastures were rotationally stocked with a 7-d grazing and 21-d resting period. Two early weaned calves (1 steer and 1 heifer) were assigned as testers to each experimental unit such that BW was equal ( 5 kg) across all experimental units. Put and take early weaned calves of comparable age and weight to the testers were used to maintain similar herbage allowance across experimental units. The literature does not define specific herbage allowance targets for the species in this study, thus our primary objectives in managing pasture quantity were to avoid i) low herbage allowances that would limit calf ADG and ii) variation among experimental units in herbage allowance within a year that could be confounded with the responses to supplement treatment. It was also recognized that variable growth conditions, especially due to rainfall, could result in year differences in herbage allowance.

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43 Treatments were three levels of a commercial pelleted concentrate, 10, 15, and 20 g kg -1 of calf BW, offered daily. Each treatment was replicated three times in a completely randomized design, so there were a total of nine experimental units in the study. In 2004, for observation purposes, two experimental units were added to the study in which the EWC did not receive concentrate supplementation. The concentrate composition is described in Table 3.1. A salt-based trace mineral mix was supplied free choice throughout the grazing season. Pasture Sampling In each 28-d grazing cycle, herbage mass was determined twice using a double-sample technique (on Paddocks 1 and 3). Indirect estimates of herbage mass were done using a 0.25-m 2 aluminum disk plate meter, and direct measures were done by clipping herbage in the same area to a 3-cm stubble. Twenty disk settling height measures were taken the day before the paddock was grazed and twenty were taken at the end of the grazing period. Twenty, 0.25-m 2 sites, representing the range of herbage mass on the nine pastures, were selected for disk calibration once a month. Disk height was measured and the herbage at these sites was then clipped to a stubble height of 3 cm, dried at 60 o C in a forced-air drier to constant weight, and weighed. A calibration equation was developed using these double sampling data by regressing actual herbage mass on disk height. To predict herbage mass for any given preor post-graze sampling date, the average of the 20 disk heights measurements was calculated and entered into the calibration equation. The regression equations and r 2 for each double sampling date are shown in Table 2 (Appendix A).

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44 Table 3-1. Ingredient and chemical composition of supplement fed to early weaned calves on pasture (DM basis). Ingredients Concentration (g kg -1 ) Corn meal 25.0 Wheat middling 400.0 Soybean meal (48% CP) 25.0 Cottonseed meal (41% CP) 50.0 Soybean hulls 392.5 Cottonseed hulls 32.5 Calcium carbonate 12.5 Salt 6.3 Quadra-plex 1.3 Molasses 50.0 Chemical composition (g kg -1 DM) Dry matter 900.0 Ash 62.3 Crude protein 146.9 TDN 700.0 Calcium 7.6 Phosphorus 5.7 Salt 6.3 Sodium 3.7 Magnesium 2.5 Potassium 12.6 Sulfur 1.4 Cobalt (mg kg -1 ) 3.36 Cooper (mg kg -1 ) 30.48 Iodine (mg kg -1 ) 0.12 Iron (mg kg -1 ) 200.12 Manganese (mg kg -1 ) 111.84 Selenium (mg kg -1 ) 0.15 Zinc (mg kg -1 ) 103.13 Provided by the manufacturer (Lakeland Animal Nutrition) Pregraze herbage mass during a grazing cycle was calculated as the average of Paddocks 1 and 3. Herbage accumulation for a given paddock was calculated by subtracting postgraze herbage mass of cycle n-1 from pregraze herbage mass of cycle n. Average herbage allowance was computed as the ratio of average herbage mass [(pregraze + postgraze) / 2] to early weaned calf liveweight exposed to the experimental

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45 unit during the grazing period (Sollenberger et al., 2005). Stocking rate (SR) was a response variable (variable SR study) and was expressed as animal units (AU; 1 AU = 500 kg of liveweight 0.75 ) ha -1 Hand-plucked samples were used to estimate nutritive value of the grazed portion of the canopy. Samples were taken at 20 locations per paddock on the day prior to grazing and herbage was removed to the stubble height at which the previous paddock was grazed. These samples were dried at 60 o C in a forced-air drier to constant weight, ground in a Wiley mill to pass a 1-mm stainless steel screen, and taken to the laboratory for analyses. Nitrogen concentration was measured using a modification of the aluminum block digestion technique (Gallaher et al., 1975). Concentration of CP in herbage DM was calculated as N x 6.25. In vitro digestible organic matter (IVDOM) concentration was determined by the two-stage procedure of Tilley and Terry (1963) modified by Moore and Mott (1974). Rumen degradable protein and rumen undegradable protein were estimated by the in vitro method proposed by Roe et al. (1991). Hand-plucked samples from 15 g kg -1 BW supplement treatment pastures were analyzed. Samples were incubated in a buffer/protease solution for 48 h, the residue recovered through filtering, and it was analyzed for CP concentration. Rumen degradable protein is that which was digested during 48 h, and rumen undegradable protein is the difference between the original CP content and the rumen degradable protein. Animal Response Variables Total organic matter intake (OMI) was computed based on fecal output and diet digestibility, viz., total OMI= OM output of feces / (1[OM digestibility/ 100]). Fecal output was estimated using a controlled release Cr marker (Captec New Zealand Limited,

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46 Auckland, New Zealand). The devices were administered orally to tester calves on 25 Mar. 2003 and 26 Feb. 2004. Four fresh fecal samples were obtained during the morning on Days 11, 13, 15, and 17 after administration. In addition, four early weaned calves fitted with the controlled release chromium marker were used for total fecal collection from Days 11 to 17 after dosing. These four calves were confined in metabolic crates and fed freshly harvested rye-ryegrass ad libitum and were supplemented with 15 g kg -1 BW concentrate. The total daily fecal output from each calf was collected, weighed, mixed and a subsample analyzed for DM and Cr concentration. Total Cr output was calculated by multiplying the fecal Cr concentration by total daily fecal weight. The total daily Cr output was used to estimate total fecal output of the tester calves by relating total daily Cr output and Cr concentration in the feces OM. Individual daily total Cr output is described in Tables 3 and 4 (Appendix A). Fecal samples were dried at 60 o C in a forced-air oven and ground in a Wiley mill to pass a 4-mm stainless steel screen. Chromium concentration in the feces was assayed by atomic absorption spectrophotometry following the procedure described by Williams et al. (1962). The samples were analyzed by d of collection in duplicate and analyses were repeated for samples where the difference of Cr concentration between duplicates exceeded 100 g kg -1 Total OMI was calculated based on fecal output estimations and IVDOM of the respective pasture herbage. Forage OMI was the difference between total OMI and the known amount of concentrate OM fed. The fecal output estimated with the marker did not equal the fecal output predicted based on estimated forage and supplement

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47 digestibilities. For this reason, an interactive SAS Institute Inc. (1991) program was used to adjust the total OMI based on the equations developed by Moore et al. (1999). This adjustment accounted for the differences in total diet digestibility due to associative effects from concentrate and forage (Appendix A). To calculate the forage intake, the following assumptions were made: 1) Digestibility of forage and supplement were determined by IVDOM 2) Digestibility of forage was affected by the level of supplement intake, as determined by the equations described by Moore et al. (1999) Diurnal grazing time of calves was evaluated on a 28-d interval. Calves were observed and actual grazing time recorded from 0700 to 1800 h. These observations were performed on the first day of the grazing period in Paddock 2 of each grazing cycle. The weight of the animals was recorded on a 28-d interval at 0900 h. The change in unshrunk weight of the tester animals was used to calculate average daily gain (ADG). Liveweight (LWG) per ha in each 28-d period was determined based on the ADG of the testers multiplied by the number of calves within the pasture during that period and adjusted to a hectare basis. In 2004 only, blood was collected from the jugular vein on a 28-d interval. Samples were collected into 9-mL, Na-heparinized syringes (Luer Monovette, LH, Sarstedt, Inc., Newton, NC) and placed on ice. Blood was centrifuged (2000 xg rcf for 30 min) and plasma was separated and frozen at -20 o C on the same day. Blood urea N (BUN) was determined using a kit (Kit B-7551-120, Pointe Scientific, Inc., Detroit, MI) and read on a plate reader at 620 nm. Economic Analysis The data and assumptions used in the economic analysis were:

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48 Concentrate cost = $0.22 kg -1 Calf price = $2.2 kg -1 Days of grazing = 100 d Where LW = calf liveweight per hectare, formulas used to calculate the results were: Concentrate cost ha -1 = LW*Concentrate level*Days of grazing*Concentrate cost Income ha -1 = gain ha -1 *calf price The return was the difference between concentrate cost ha -1 and income ha -1 The objective of the economic analysis was to compare the specific treatments used in this study; it should not be extrapolated to a wide range of production systems. Statistical Analysis All responses were analyzed by fitting mixed effects models using the PROC MIXED procedure of SAS (SAS Institute Inc., 1996). Replicate and its interactions were considered random effects. Single degree of freedom orthogonal polynomial contrasts were used to test concentrate effects. Treatments were considered different when P <0.05. Interactions not discussed in the Results and Discussion section were not significant (P>0.05). The means reported in this text are least squares means. For pasture and pasture-animal interface variables, the model used was: Y ijk = + A i + T j + P k + (AT) ij + (AP) ik + (TP) jk + (ATP) ijk + e ijk Where Y ijk is the dependent variable is the overall mean T j is the concentrate effect (main plot) A i is the year effect (sub-plot)

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49 P k is the period effect (sub-sub-plot) (AT) ij is the year*concentrate interaction (AP) ik is the year*period interaction (TP) jk is the concentrate*period interaction (ATP) ijk is the year*concentrate*period interaction e ijk is the error Main plot (concentrate) error was replicate*concentrate; sub-plot (year) error was replicate*year; sub-sub-plot (period) error was the residual. For the animal variables, the model used was: Y ijkl = + A i + T j + P k + S l + (AT) ij + (AP) ik +(PS) kl +(AS) il + (TP) jk +(TS) jl +(ATP) ijk +(ATS) ijl +(TPS) jkl +(ATPS) ijkl + e ijkl Where Y ijkl is the dependent variable is the overall mean T j is the concentrate effect (main plot) A i is the year effect (sub-plot) S l is the sex effect (sub-sub-plot) P k is the period effect (sub-sub-sub-plot) (AT) ij is the year*concentrate interaction (AP) ik is the year*period interaction (AS) il is the year*sex interaction (TP) jk is the concentrate*period interaction (TS) jl is the concentrate*sex interaction (PS) kl is the concentrate*sex interaction

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50 (ATP) ijk is the year*concentrate* period interaction (ATS) ijl is the year*concentrate*sex interaction (TPS) jkl is the concentrate*period*sex interaction (ATPS) ijkl is the year*concentrate*period*sex interaction e ijkl is the error Main plot (concentrate) error was replicate*concentrate; sub-plot (year) error was replicate*year; sub-sub-plot (sex) error was replicate*sex; sub-sub-sub-plot (period) error was the residual. Results and Discussion Pasture Responses There were no differences in pregraze herbage mass among the different levels of concentrate (Table 3-2). There was a significant year*period interaction that occurred because herbage mass was greater during January to March 2004 than in 2003, but there was no year effect for April (Fig. 3-1). The greater rainfall from November 2002 to March 2003 (Fig. 3-2) resulted in pastures with excessive soil moisture and likely reduced the efficiency of utilization of N fertilizer causing lesser herbage mass during the first 3 mo of the 2003 experimental period. The soils at the research site are characterized by poor to somewhat poor drainage and are susceptible to flooding under conditions of excessive rainfall. In contrast, limited rainfall in March and April 2004 likely resulted in lesser mass during April than in January through March. Macoon (1999) reported average herbage mass on rye-ryegrass pastures of 1.4 Mg ha -1 for dairy cows at a SR of five cows ha -1 In that study, there was a significant effect of month of sampling. Herbage mass was greater (1.3 Mg ha -1 ) in March than in January (0.8 Mg ha -1 ). According to Rouquette et al. (1997), small grains grow better from late

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51 December to mid-February and ryegrass has rapid forage growth during March to late May, and as a result stocking rate often needs to be increased to efficiently use ryegrass pastures. In the current study, this type of response was observed only in 2003. Table 3-2. Pregraze herbage mass and herbage accumulation of rye-ryegrass pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight) Concentrate Herbage mass Herbage accumulation -----------g kg -1 BW----------------------Mg ha -1 ----------------------kg ha -1 d -1 --------10 1.6 31 15 1.5 27 20 1.7 35 Polynomial contrast NS NS P value 0.18 0.44 SEM 0.1 3.0 NS = not significant Herbage accumulation was similar among treatments, with an average of 31 kg DM ha -1 d -1 (Table 3-2). These rates are comparable to those reported by Assmann et al. (2004). In their study in southern Brazil, ryegrass pastures were grazed by beef steers (3 AU ha -1 ), fertilized with 100 kg N ha -1 and herbage accumulation was 37 kg DM ha -1 d -1 For heavily-stocked (5 cows ha -1 ) rye-ryegrass pastures grazed by lactating dairy cows, Macoon (1999) observed 22 kg DM ha -1 d -1 There was a significant year*period interaction for herbage accumulation (Fig. 3-3). This interaction occurred because herbage accumulation in April 2003 was much greater than in 2004, while in other months there were no differences. Excessive rainfall during November and December 2002 (220 mm vs. 30-yr average of 133 mm) resulted in standing water in some pastures and decreased herbage accumulation during the first 2 mo after planting. In April and May 2003, soils retained sufficient moisture despite fewer

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52 rainfall events to provide a better growth environment for the plants. In contrast, the decreased herbage accumulation in April 2004 reflects lower than normal rainfall in March (51 vs. 30-yr average of 93 mm) and April (26 vs. 30-yr average of 75 mm) of that year (Fig. 3-2). Table 3-3. Herbage crude protein (CP) and in vitro digestible organic matter (IVDOM) concentrations of rye-ryegrass pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight) Concentrate CP IVDOM -----------g kg -1 BW----------------------g kg -1 DM--------------------g kg -1 OM----------10 273 841 15 282 837 20 283 838 Polynomial contrast NS NS P value 0.13 0.51 SEM 4.0 3.0 NS = not significant There was no concentrate level effect on herbage CP and IVDOM (Table 3-3). The CP concentrations were greater in 2004 (302 g kg -1 ) than 2003 (257 g kg -1 ) in all periods likely because the very high rainfall during parts of the 2003 season. An anaerobic root environment caused by excess rain can decrease absorption of N, resulting in lesser N concentration in the plants (Salisbury and Ross, 1991). Also, heavy rain likely leached some N out of the root zone.

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53 0.00.51.01.52.02.5Jan.Feb.Mar.Apr.MonthsHerbage mass (Mg DM ha-1) Year 2003 Year 2004******NSbababaab Fig. 3-1. Year*period interaction effect on pregraze herbage mass of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate. Year effect within period; **, P 0.01; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05). 050100150200250OctNov.Dec.Jan.Feb.Mar.Apr.MayMonthsRainfall (mm) 2002/03 2003/04 Fig. 3-2. Rainfall distribution from October to May 2002/03 and 2003/04.

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54 0102030405060708090Jan.Feb.Mar.Apr.MonthsHerbage accumulation (kg DM ha-1d-1) Year 2003 Year 2004NSNSNS** b bbabbab Fig. 3-3. Year*period interaction effect on average herbage accumulation of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate. Year effect within period; **, P 0.01; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05). There was a year*period interaction for CP concentration (Fig. 3-4). The herbage CP concentration was least in January compared to other months because of maturity. There was greater length of growth period of this forage herbage (~80 d from planting) relative to regrowth grazing (21 d). Additionally, the lowest CP observed in this study was in January 2003 following very heavy November and December rains. The greater CP concentrations in February and March are associated with the dates of N fertilization on the pastures (early February and early March). There also was year*period interaction for IVDOM (Fig. 3-5). There was a significant year effect on IVDOM concentration in January only. The IVDOM was not affected by climatic conditions to the same degree as herbage mass, accumulation, and CP concentrations were. The IVDOM decreased in both years, from January to April, primarily because of the presence of reproductive tillers after February. Highest IVDOM,

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55 unlike CP, occurred in January. Other studies (Chapter 5) have shown cool-season grass IVDOM to be relatively insensitive to length of regrowth period during winter. The CP and IVDOM concentrations were similar to the average CP (285 g kg -1 ) and IVDOM (756 g kg -1 ) values reported by Arthington and Kalmbacher (2002) for ryegrass pastures grazed by early weaned calves. Redfearn et al. (2002) showed CP and IVDMD concentrations to be 232 and 846 g kg -1 respectively, for ryegrass pastures grazed by beef steers. Fractionation of CP showed ruminal-degradable protein concentration to be 730 g kg -1 CP and ruminal-undegradable protein concentration to be 270 g kg -1 CP (data not shown). These very high concentrations of ruminal degradable protein have the potential to result in significant loss of N in ruminants. Excessive ruminal-degradable protein consumed by the animal is absorbed by the rumen epithelium and excreted as urea in the urine. The NRC (1996) reported lower values for the ruminal degradable protein of ryegrass hay (650 g kg -1 ). Using the in situ methodology, Michalet-Doureu and Ould-Bah (1992) observed similar ruminal degradable protein concentration of 740 g kg -1 on ryegrass hay. Herbage allowance was similar among the treatments (Table 3-4). This was achieved by increasing SR on high concentrate treatment pastures to account for substitution of concentrate for forage. The average herbage allowance through the experimental period was greater in 2004 (0.8) than in 2003 (0.4) due to superior fall growth preceding the initiation of grazing in the second year.

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56 Table 3-4. Herbage allowance and SR of rye-ryegrass pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight) Concentrate Herbage allowance SR -----------g kg -1 BW----------------kg DM kg -1 BW-----------------AU ha -1 ------------10 0.62 5.5 15 0.59 5.9 20 0.66 6.5 Polynomial contrast NS L P value 0.38 < 0.01 SEM 0.02 0.13 AU = 500 kg BW 0.75 NS = not significant, L = linear Phillips (1989) reported that to prevent a decline in individual dairy cow performance, herbage allowance should allow for dry matter intake of at least 40 g OM kg -1 BW d -1 McCartor and Rouquette (1977) studied the effect of different levels of herbage allowance on performance of beef calves grazing pearl millet [Pennisetum glaucum (L.) R. Br.] and concluded that animal performance declines as herbage allowance levels fall below 1 kg DM kg -1 BW. According to Le Du et al. (1979), the maximum intake of cool-season grasses occurs when herbage allowance is at least twice the herbage intake. The average herbage allowance observed in this study was always more than twice the early weaned calves forage intake. There was no correlation (r 2 = 0.03) between herbage allowance and ADG (Table 3-5) of the calves implying that factors other than herbage allowance were influencing animal performance.

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57 050100150200250300350400Jan.Feb.Mar.Apr.MonthsCP (g kg-1DM) Year 2003 Year 2004*******dabccabc Fig. 3-4. Year*period interaction effect on crude protein (CP) concentration of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate. Year effect within period; **, P 0.01; *, P 0.05; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05). 700750800850900Jan.Feb.Mar.Apr.MonthsIVDOM (g kg-1OM) Year 2003 Year 2004**NSNSNSaabbccdd Fig. 3-5. Year*period interaction effect on in vitro digestible organic matter (IVDOM) concentration of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate. Year effect within period; **, P 0.01; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05).

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58 There was a significant year*period interaction on herbage allowance (Fig. 3-6). There was no difference in herbage allowance during January and February 2003. Herbage allowance was less in March than January, but there was no difference between February and March. During April, herbage allowance was greater than in the other months. This was a result of the greater herbage accumulation (Fig. 3-1) during that month. In 2004, herbage allowance was greatest in January and decreased through April. High herbage allowance in January 2004 was due to excellent forage growth during fall 2003 and conservative stocking decisions to start the experiment in 2004. 00.20.40.60.811.21.4Jan.Feb.Mar.Apr.MonthsHA (kg DM kg-1BW) Year 2003 Year 2004********babcbcbac Fig. 3-6. Year*period interaction effect on herbage allowance of rye-ryegrass pastures grazed by early weaned calves supplemented with different levels of concentrate. Year effect within period; **, P 0.01; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05). Horn et al. (1995) conducted a 3-yr experiment to evaluate the influence of energy supplements on performance of stocker cattle grazing wheat pasture. They concluded that energy supplements, such as maize and soybean [Glycine max (L.) Merrill] hulls, when fed at 7.5 g kg -1 BW allowed a 33% increase in stocking rate compared to

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59 unsupplemented calves. In this study, there was 18% increase in SR when the supplement level was increased from 10 to 20 g kg -1 BW (Table 3-5). Animal Response Variables There was a linear increase in ADG and LWG as supplementation level increased (Table 3-5). The average daily gain reported for calves supplemented at the 10 g kg -1 BW concentrate level in this study was the same as that reported by Arthington and Kalmbacher (2003) for calves grazing ryegrass in south Florida. There were no effects due to sex or year on ADG, however, there was a year*period interaction (Fig. 3-7). The decrease in ADG during February and March 2003 was associated with an infestation of endoparasites that was not detected by visual examination. After the depression in ADG was observed in March, feces samples were collected, examined, and a high incidence of parasites detected. Calves were individually treated with 6 ml of Ivermectin (10 g kg -1 ) and a large increase in ADG was observed during April. The ADG and LWG of calves that did not receive concentrate supplementation were 0.3 kg d -1 and 650 kg ha -1 respectively. These values were well below the ADG and gain per ha of calves supplemented with 10 g kg -1 BW in concentrate, 0.74 kg d -1 and 950 kg ha -1 respectively. Although the forage had nutritive value similar to the concentrate, the low DM concentration (150-180 g kg -1 ) of the forage may be limited the total OMI of the animals resulting in decreased performance. The linear increase in LWG was a function of the linear increase in both SR and ADG as concentrate level increased. Horn et al. (1995) observed greater LWG on steers grazing wheat pastures supplemented with concentrate at 7.5 g kg -1 BW (153 kg ha -1 ) than for the unsupplemented calves (103 kg ha -1 ). Restle et al. (2000) studied beef heifers grazing oat-ryegrass pastures fertilized with 200 kg N ha -1 under continuous stocking.

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60 Average daily gain and LWG were 0.58 kg d -1 and 429 kg ha -1 respectively, less than the values in our study. The results of the current study agree with the literature on the effects of supplementation on beef calves grazing cool-season grass pastures, i.e., energy supplementation leads to substitution of forage by concentrate, allowing higher stocking rates and gains per unit land area. There was no difference in total OMI among treatments; however, there was a linear decrease in forage OMI with increasing levels of supplement (Table 3-6). Total OMI was similar to that reported by Paisley et al. (1998) for early weaned calves grazing wheat pastures (28 g kg -1 BW). In general, forage intake is decreased by feeding concentrate when forage quality is high, other nutrients are in balance with energy, and concentrate is fed in large amounts (Moore, 1994). Carey et al. (1993) supplemented beef steers grazing tall fescue (Festuca arundinacea Schreb.) pastures with different sources of energy. Forage intake was less for steers supplemented with energy than for control steers, but total intake (forage + supplement) did not differ among treatments. Cravey (1993) reported substitution ratios of 1:0.93 (forage:concentrate) when steers grazing wheat pastures were fed energy concentrate at 10 g kg -1 BW. The substitution rate of concentrate OM (0.8, 1.2, and 1.6 for 10, 15, and 20 g kg -1 treatment respectively) for forage OM in the current study was 1:1 and consistent across treatments, confirming observations in the literature of substitution of forage by concentrate intake at similar proportions by calves receiving high amounts of energy supplement and grazing high nutritive value cool-season pastures.

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61 Table 3-5. Average daily gain (ADG) and liveweight gain (LWG) of early weaned calves grazing rye-ryegrass pastures and supplemented with three levels of concentrate. (BW=body weight) Concentrate ADG LWG -----------g kg -1 BW-----------------------kg d -1 ------------------------kg ha -1 -----------10 0.74 950 15 0.81 1080 20 0.89 1320 Polynomial contrast L L P value 0.01 0.01 SEM 0.03 42 Gain per ha during a 106-d period. L = linear 00.20.40.60.811.2Jan.Feb.Mar.Apr.MonthsADG (kg d-1) Year 2003 Year 2004**NS****acbbcaaab Fig. 3-7. Year*period interaction effect on average daily gain of early weaned calves grazing rye-ryegrass pastures and supplemented with different levels of concentrate. Year effect within period; **, P 0.01; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05). The greater ADG at higher concentrate levels despite similar total intake may be a function of the different sources of nutrients provided by the concentrate that resulted in

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62 better N and energy synchronization in the rumen and increased microbial protein production. In addition concentrate had more rumen-undegradable protein than the forage, potentially leading to more amino acids being absorbed post rumen and decreasing the levels of ammonia absorbed by the rumen epithelium. Absorption and excretion of excess ammonia is a process that spends energy that otherwise could be used to improve animal performance. Calves that received greater concentrate rates spent less time grazing during daylight (Table 3-7) likely reducing their energy requirement for maintenance. This could also be a factor contributing to greater ADG for calves receiving greater levels of concentrate, despite total OMI being the same across treatments.. Table 3-6. Total organic matter intake (OMI) and forage OMI of rye-ryegrass pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight) Concentrate Total OMI Forage OMI ---------g kg -1 BW----------------------(% BW)--------------10 2.6 1.8 15 2.4 1.3 20 2.7 1.1 Polynomial contrast NS L P value 0.68 < 0.01 SEM 0.01 0.01 Total OMI intake data adjusted according to Moore et al. (1999). NS = not significant, L = linear Grazing time differed among the calves receiving different levels of concentrate. Calves supplemented with 10 g kg -1 BW concentrate spent more time grazing during daylight hours than calves receiving 15 and 20 g kg -1 BW (Table 3-7). There was no sex effect on grazing time.

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63 In research with yearling beef steers grazing ryegrass, Adams (1985) reported that although there was an 11% decrease in forage intake by supplemented calves compared with control calves, total daily grazing time was not altered. Macoon (1999) observed that dairy cows supplemented with high levels of concentrate spent less time grazing (135 min) than cows that received low rates of concentrate (180 min). Cowan et al. (1977) observed decreased grazing time of 23 min d -1 for each additional kg of concentrate fed to cows grazing grass-legume pastures. Increased grazing time in the current study suggests that calves supplemented with lower rates of concentrate grazed longer during the day in an attempt to meet their nutritional requirements. According to Macoon (1999), lactating cows receiving low levels of concentrate tended to increase the night-time grazing as well. Table 3-7. Grazing time during daylight hours of early weaned calves grazing rye-ryegrass pastures and supplemented with different levels of concentrate. Concentrate Grazing time -----------g kg -1 BW------------------------------------------min------------------------------10 284 15 230 20 234 Polynomial contrast Q P value < 0.08 SEM 17 Q = quadratic. There was no effect of three levels of concentrate supplementation on calf BUN concentrations (Table 3-8). Blood urea N levels between 11 and 15 mg dL -1 were associated with maximum rates of gain for growing steers (Byers and Moxon, 1980), thus it is likely that the calves had an adequate protein supply during the entire experimental

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64 period. Increased solubility and degradability of dietary protein can lead to increased ruminal ammonia concentrations resulting in increased BUN concentrations (Hammond et al., 1994). Protein in intensively fertilized cool-season grasses is highly degradable and thus easily fermented to volatile fatty acids and ammonia in the reticulo-rumen. Ammonia that is not captured is absorbed and excreted in the urine (Van Vuuren et al., 1991). It was expected that calves receiving greater amounts of concentrate would have a lesser BUN concentration because the concentrate had lower CP and ruminal degradable protein concentrations than the rye-ryegrass pastures. There was a numerical suggestion of lesser blood urea N for calves receiving the 20 g kg -1 BW concentrate compared to the 15 g kg -1 level, however, no statistical difference was observed. Table 3-8. Blood urea nitrogen (BUN) concentrations of early weaned calves grazing rye-ryegrass pastures supplemented with three levels of concentrate. (BW=body weight) Concentrate BUN ---------g kg -1 BW---------------------------------------mg dL -1 ------------------------------10 14.1 15 15.1 20 13.0 Polynomial contrast NS P value 0.26 SEM 1.0 NS = not significant According to the NRC (1996), the ruminal-degradable protein requirement for 120 kg calves is 300 g d -1 Based on the estimations of forage and concentrate intake for calves receiving the 10 g kg -1 BW supplement and the data for CP fractionation, ryegrass provided ~ 380 and concentrate 100 g of ruminal degradable protein d -1 Therefore, the estimated ruminal degradable protein consumed was 60% greater than the requirement.

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65 The excessive rumen degradable protein consumed by the calves was not expressed in increased BUN concentrations that were in the range considered adequate for growing calves. Economic Analysis There was a linear increase in supplement cost and income with increasing supplement levels; however, there was no difference in return among the treatments (Table 3-9). Provided that there are no economic incentives to producing heavier calves, these data do not justify use of supplement levels above 10 g kg -1 BW. Table 3-9. Economic analysis of three concentrate levels on early weaned calves grazing rye-ryegrass pastures. Concentrate Concentrate cost Income Return --------g kg -1 BW-----------------------------------$ ha -1 ------------------------------10 600 2100 1500 15 970 2370 1400 20 1430 2900 1470 Polynomial contrast L L NS P value < 0.01 < 0.01 0.86 SEM 25 91 92 L = linear; NS = not significant Summary and Conclusions Herbage mass, accumulation, allowance, and nutritive value of rye-ryegrass pastures did not differ among levels of concentrate. Seasonal variation in forage responses were the result of the climatic conditions during the trial, especially the very high rainfall that occurred during the 2002/03 season. It resulted in lower herbage mass and CP during that year.

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66 Total OMI did not vary among the three supplementation levels, but forage OMI decreased linearly as supplement level increased. Concentrate substituted for forage intake in ~ a 1:1 ratio. To enable greater forage intake, calves receiving the 10 g kg -1 BW concentrate level spent more time grazing during daylight hours than calves receiving the 15 and 20 g kg -1 BW treatments. Average daily gain, SR, and LWG increased linearly with increasing levels of concentrate. Increased ADG at the same level of intake implies better synchronization of energy and N in the rumen, and possibly an effect associated with the greater amounts of ruminal undegradable protein provided by the concentrate. Stocking rate increased because of the substitution of concentrate for forage in the 15 and 20 g kg -1 BW treatments. There was a trend for BUN concentration to be lower for the 20 than 15 g kg -1 BW treatment, likely the result of the lower CP and ruminal degradable protein concentration in the concentrate compared to the forage. The BUN concentrations of all treatments were in the range considered adequate for growing cattle. The economic analysis implies that there is no economic benefit of increasing the supplementation levels above 10 g kg -1 BW, unless a greater price is paid for heavier weight calves. In practice, the optimum supplementation level will be dependent upon forage availability, cost of the concentrate, and calf price. Given the pasture conditions, and the concentrate and calf pricing in this study, the 10 g kg -1 BW supplement level is recommended. Future studies should evaluate lesser levels, e.g., 5 g kg -1 to determine if more economical gains can be achieved with less supplement.

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CHAPTER 4 SUPPLEMENTATION EFFECTS ON FORAGE CHARACTERISTICS AND PERFORMANCE OF EARLY WEANED CALVES GRAZING TIFTON 85 BERMUDAGRASS PASTURES Introduction There are 39 million ha of perennial pastures grown in the eastern USA and almost 30 million ha are located in the South. These pastures are mainly used for beef cows and calves and stockering of weaned calves (Hoveland, 1992). Among several species found in the southeastern USA, bermudagrass [Cynodon dactylon (L.) Pers.] is one of the most important with ~ 10 to 12 million ha of planted for livestock grazing and hay. Johnson et al. (2001b) compared different species of warm-season grasses in Florida and showed that bermudagrass produced more forage DM (1540 43 kg ha -1 cutting -1 ) than stargrass (Cynodon nlemfuensis Vanderyst) (1400 43 kg ha -1 cutting -1 ) or bahiagrass (Paspalum notatum Flgge) (1300 43 kg ha -1 cutting -1 ). Moreover, bermudagrass was more digestible (575 4 g kg -1 ) than stargrass (546 4 g kg -1 ) and bahiagrass (519 4 g kg -1 ). In 1993, Tifton 85, a hybrid between a South African bermudagrass and Tifton 68 stargrass was released. Burton et al. (1993) described it as taller, with larger culms, broader leaves, and darker color than other bermudagrass hybrids. Compared with Coastal, Tifton 85 yielded 26% more dry matter, was 110 g kg -1 more digestible, and more succulent at harvest. Warm-season grasses have relatively larger proportions of cell-wall material. Cell wall is potentially digestible, but because of the chemical composition and anatomical structure of warm-season grasses the rate of degradation and fermentation is relatively 67

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68 slow (Coleman et al., 2004). Although forage nutritive value of C4 grasses is generally inferior to that of C3 grasses, animal production can be high on C4 grasses because of their high DM production potential. Pedreira et al. (1998) studied Tifton 85 bermudagrass in a 3-yr grazing trial. The pastures were continuously stocked with a fertilization program of 210 kg N ha -1 yr -1 Tifton 85 pastures supported six heifers ha -1 resulting in live weight gain (LWG) 648 kg ha -1 during spring-summer in North-Central Florida. The use of supplementary feed is a management practice to overcome the nutritional shortfalls of the forage component of the diet. When energy and protein requirements are high, e.g., for lactation, pregnancy, and growth, part of the roughage component may need to be replaced by feeding of concentrates (Fontaneli, 1999). Rouquette et al. (2003) reported that stocker steers gained 1.03 kg d -1 on Tifton 85 compared to 0.75 kg d -1 on Coastal bermudagrass without supplementation. With supplementation of 1.4 kg d -1 of a 3:1 maize (Zea mays L.):soybean [Glycine max (L.) Merr.] meal concentrate, steers grazing Tifton 85 gained 1.25 kg d -1 According to Galloway et al. (1992), moderate dietary levels of supplement (200-300 g kg -1 of the diet DM) can improve nutrient intake and performance by cattle consuming bermudagrass. At greater amounts, nutrient digestion, intake, or both of the forage portion of the diet can be affected negatively. Wheeler et al. (2002) tested the effects of increasing supplement protein concentration on performance and forage intake of beef steers consuming bermudagrass forage. Treatments were no supplement or daily equivalents of 0.2, 0.4, and 0.6 g of supplemental protein kg -1 of body weight (BW). Forage intake increased 16% and organic matter (OM) intake increased 30% in supplemented compared to unsupplemented steers. Diet OM digestibility increased 145 g

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69 kg -1 and total digestible OM intake increased 490 g kg -1 in supplemented compared to unsupplemented steers. The success of the Florida cow-calf enterprise depends upon maintaining high calving percentage. First-calf heifers and other young cows may experience low conception rates due to stress of lactation. Early weaning of calves of young cows is one approach to reduce cow nutrient requirement, increase body condition, and increase the likelihood of rebreeding. Management practices for raising the early weaned (EW) calf have not been widely studied. In particular there is little information to guide supplementation programs for EW calves grazing C4 grass pastures. In a study done in south Brazil, Muehlmann et al. (1997) reported average daily gains (ADG) of 0.2 kg d -1 for EW calves grazing a bermudagrass-native warm-season grasses mixture. Vendramini et al. (2003) observed that supplementing EW calves with concentrate at 10 g kg -1 BWof body weight resulted in gains of 0.59 and 0.44 kg d -1 on stargrass and atrapaspalum (Paspalum atratum Swallen), respectively. Additional information is needed regarding the amount of supplement required by EW calves grazing C4 grass pastures. The objective of this study was to evaluate the effect of different levels of concentrate on ADG, stocking rate (SR), LWG, and forage intake of EW calves grazing Tifton 85 bermudagrass pastures in North-Central Florida. Material and Methods The research site was the Florida Beef Research Unit, 18 km northeast of Gainesville, FL (30 o N). Dates for the trial during the 2 yr were 14 May through 13 Aug. 2003 (86 d) and 18 May through 10 Aug. 2004 (86 d). Monthly rainfall and temperature data during the trial are presented in Table 1 (Appendix A).

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70 The soils at the research site are Adamsville fine sand (Aquic Quartzipamments, hyperthermic, uncoated) and Sparr fine sand (Grossarenic Paleudult, hyperthermic, loamy siliceous). These soils are moderately well drained with rapid permeability. Prior to initiation of summer grazing at the site, mean soil pH (1:2 soil:deionized H 2 O ratio) was 5.5, and Mehlich-I (0.05 M HCl + 0.0125 M H 2 SO 4 ) extractable P, K, Mg, and Ca in the Ap1 horizon (0to 15-cm depth) were 44, 17, 40, 328 mg kg -1 respectively. Bermudagrass pastures were established in August 2002 and fertilized with 40 kg ha -1 of N, 17 kg ha -1 of P, and 66 kg ha -1 of K 1 mo after establishment. In April 2003 and 2004, the pastures received 40 kg N ha -1 17 kg P ha -1 and 66 kg K ha -1 An additional 80 kg N ha -1 was split in two applications of 40 kg ha -1 in early June and early July of both years. Treatments were three levels of a commercial pelleted concentrate, 10, 15, and 20 g kg -1 of the calves body weight (BW), offered daily. The concentrate composition was described in Table 3.2 (Chapter 3). Each treatment was replicated three times in a completely randomized design so there were a total of nine experimental units in the study. In 2004, for observation purposes, two experimental units were added to the study in which the EWC did not receive concentrate supplementation. The EWC were described in Chapter 3. At the start of this trial, they were ~ 200-d old with an average liveweight of 190 kg. Calves grazed rye (Secale cereale L.)-annual ryegrass (Lolium multiflorum Lam.) pastures from January to April. Calves received the same supplement treatment in summer as they did during the winter-spring period. At the initiation of grazing of the Tifton 85 pastures, calves within a supplement level were randomly allocated to the three replicates of that treatment. They were vaccinated with

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71 Ultrabac 8 and Bovashield 4 (Pfizer Animal Health New York, NY) and dewormed with ivermectin (Ivomec, Merck & Company, Rahway, New Jersey, USA) 10 g kg -1 concentration on 23 July 2003 and 14 July 2004. Pasture size was 0.15 ha subdivided in three paddocks for rotational stocking. The grazing period was 7 d and the rest period was 14 d. Two EW calves (1 steer and 1 heifer) were assigned as testers to each pasture and put and take EW calves were used to maintain herbage allowance at ~1 kg forage DM kg -1 of calf BW. Pasture Sampling In each grazing cycle (21 d), herbage mass was determined on Paddock 2 using a double sampling technique. The procedure of herbage sampling was similar to the one described in Chapter 3. A total of 40, double-samples were taken every 3 wk to calibrate the disk meter. This greater number than used during the winter was necessary because C4 grasses usually have a greater heterogeneity of herbage bulk density through the different heights of the canopy (Sollenberger and Burns, 2001). Thus, separated preand post-graze calibration equations resulted in more precision of the height/weight relationship than combining them into one. The regression equations and r 2 of the preand post-graze double sampling technique are shown in Table 1 (Appendix B). The methodology used to determine herbage allowance, stocking rate, and forage nutritive value were the same as those described in Chapter 3. Animal Response Variables Total organic matter intake (OMI) and forage OMI were estimated only in 2003. The controlled-release devices used in the trial were designed for sheep or small calves. When used by 200-kg calves, most of the calves regurgitated the device before the fecal collection period. The data are presented only for 2003 and reflect results from only 8 out

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72 of 18 calves dosed with the controlled-release device. The total Cr output calculated from total fecal collection from three calves is presented in Table 2 (Appendix B). The Cr analyses procedure and total and forage OMI determinations were described in Chapter 3. Grazing time of EW calves during daylight hours was evaluated every 21 d. Calves were observed and time spent grazing was recorded from 0700 to 1900 h. These observations were performed on the first day of the grazing period in Paddock 2 in each grazing cycle. The weight of the animals was recorded on a 21-d interval at 0900 h. The change in unshrunk weight of the tester animals was used to calculate ADG. The LWG in each 21-d period was determined based on the ADG of the testers multiplied by the number of calves within the pastures during that period and adjusted to a hectare basis. Blood urea N (BUN) sampling and determination were conducted as described in Chapter 3. Economic Analysis The assumptions used in the economic analysis were: Concentrate cost = $0.22 kg -1 Calf price = $2.20 kg -1 Live weight (LW) = SR*500 kg Days of grazing = 90 d The formulas used to calculate the results were described in Chapter 3. Statistical Analysis The models, procedure, and software used for statistical analyses were described in Chapter 3.

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73 Results and Discussion Pasture Responses There was no difference in pregraze herbage mass and herbage accumulation among the treatments (Table 4-1). Table 4-1. Pregraze herbage mass and herbage accumulation of Tifton 85 pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight) Concentrate Herbage mass Herbage accumulation -----------g kg -1 BW--------------------Mg ha -1 -----------------------kg ha -1 d -1 --------10 4.9 101 15 4.8 95 20 4.7 84 Polynomial contrast NS NS P value 0.20 0.11 SEM 0.1 15 NS = not significant Using continuous stocking, Pedreira et al. (1998) reported greater yields of Tifton 85 than Florakirk bermudagrass early in the grazing season. Herbage accumulation was 73 kg DM ha -1 d -1 for Tifton 85 vs. 44 kg DM ha -1 d -1 for Florakirk. Under continuous stocking, average herbage mass during a 3-yr study with Tifton 85 grazed by beef steers in Georgia was 2.8 Mg ha -1 and ADG decreased when herbage mass was below 2.5 Mg ha -1 (Hill et al., 1993). Greater average herbage mass was maintained in our study to provide opportunity for the calves to select forage that was high in nutritive value and maximize animal performance.

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74 0.01.02.03.04.05.06.07.0MayJun.Jul.Aug.MonthsHerbage mass (Mg DM ha-1) Year 2003 Year 2004****NS**aabcbabcd Fig. 4-1. Year*period interaction effect on pregraze herbage mass of Tifton 85 pastures grazed by early weaned calves supplemented with different levels of concentrate. Year effect within period; **, P 0.01; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05). There was a significant year period interaction effect on herbage mass (Fig. 4-1). The lesser herbage mass present in the first 2 mo of 2004 can be attributed to very low rainfall that occurred during May (15 mm vs. the 30-yr average of 106 mm). Herbage accumulation also varied throughout the grazing season ranging from 45 to 121 kg ha -1 d -1 in 2003 and from 51 to 133 kg ha -1 d -1 in 2004 (Fig. 4-2). According to Dubeux et al. (2003), rotationally stocked pastures of bahiagrass had greater herbage accumulation than continuously stocked pastures. The use of a rotational stocking may be one of the factors responsible for greater herbage accumulation in this study compared to those of Hill et al. (1993) and Pedreira et al. (1998) who stocked their pastures continuously.

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75 020406080100120140160Jun.Jul.AugMonthsHerbage accumulation (kg DM ha-1d-1) Year 2003 Year 2004******cbbaac Fig. 4-2. Year*period interaction effect on average herbage accumulation of Tifton 85 pastures grazed by early weaned calves supplemented with different levels of concentrate. Year effect within period; **, P 0.01; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05). Herbage CP and IVDOM concentrations were not affected by treatment and averaged 184 g kg -1 and 624 g kg -1 respectively (Table 4-2). These concentrations are greater than those reported by Pedreira et al. (1998) (120 g kg -1 ) and Hill et al. (1993) (140 g kg -1 ) for Tifton 85 pastures stocked continuously. Fike et al. (2002) used rotational stocking with similar resting period (21 d) and N fertilization program (45 kg N ha -1 on 21 May, 8 June, and 7 August) on Tifton 85 pastures. They reported CP and IVDOM values (140 and 600 g kg -1 respectively) lower than those observed in this study. Masticate samples from Tifton 85 pastures had CP and IVDMD concentrations of 180 and 650 g kg -1 (Hill et al., 1998). With a 35-d regrowth interval, Mislevy and Martin (1998) reported CP of 170 g kg -1 DM for hand-plucked samples of stargrass pastures grazed by beef steers.

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76 Table 4-2. Crude protein (CP) and in vitro digestible organic matter (IVDOM) concentration of Tifton 85 pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight) Concentrate CP IVDOM -----------g kg -1 BW---------------------g kg -1 DM---------------------g kg -1 OM---------10 182 626 15 181 610 20 189 636 Polynomial contrast NS NS P value 0.49 0.44 SEM 6 8 NS = not significant There was a year*period interaction for CP concentration (Fig. 4-3). In both years, the lower CP during the first month of grazing was attributed to a longer growth period preceding the first grazing cycle than subsequent cycles. At initiation of grazing in May, the herbage present reflected growth that had occurred during the spring and not from the most recent 14 d. Thus, hand-plucked samples included older herbage with more stem material resulting in lesser CP and IVDOM concentrations. Pedreira (1995) reported similar results from bermudagrasses pastures grazed by beef heifers. Herbage CP was 70 g kg -1 in June while the average for June through August was 125 g kg -1 Similar results were presented by Hill et al. (1993), with CP concentrations of 114 (May) and 156 g kg -1 (September) in the first and last month of the grazing study with Tifton 85.

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77 050100150200250300MayJun.Jul.Aug.MonthsCP (g kg-1DM) Year 2003 Year 2004NS****NSaabcaabc Fig. 4-3. Year*period interaction effect on crude protein (CP) concentration of Tifton 85 pastures grazed by early weaned calves supplemented with different levels of concentrate. Year effect within period; **, P 0.01; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05). There was no year*period interaction effect on herbage IVDOM, but there were year and period (Fig. 4-4) effects. Mean herbage IVDOM was greater in 2004 (640 g kg -1 ) than in 2003 (580 g kg -1 ). The herbage IVDOM period response was similar to that of CP concentration, with the lowest values occurring during the first month of grazing because of the longer regrowth period of the forage. Herbage IVDOM was greater in June and decreased in July and August. The estimated rumen-degradable and rumen-undegradable protein concentrations were 550 and 450 g kg -1 CP, respectively (data not shown). Similar values for bermudagrass rumen-degradable protein (570 g kg -1 ) were found by Basurto et al. (2001). Mathis et al. (2001) measured bermudagrass ruminal degradable protein from seven different locations using the enzymatic methodology proposed by Roe et al. (1991). The average across locations was 610 g kg -1 CP.

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78 050100150200250300350400450500550600650700750MayJun.Jul.Aug.MonthsIVDOM (g kg-1DM)abcd Fig. 4-4. Period effect on IVDOM concentration of Tifton 85 herbage grazed by early weaned calves supplemented with different levels of concentrate. Means are different if followed by different letters (P 0.05). There was a linear decrease in average herbage allowance with increased supplementation levels, although the magnitude of the change was small (Table 4-3). Put and take animals were used to maintain similar herbage allowance among treatments. Even though herbage mass was not different among treatments, the greater SR for the 20 g kg -1 BW treatment caused herbage allowance to be slightly reduced. There was no period or period year interaction on herbage allowance, however, a year effect was observed with averages across treatments of 1.0 and 0.8 kg DM kg -1 BW for 2003 and 2004, respectively. Herbage allowance of at least 1.0 kg DM kg -1 BW was associated with highest gains of unsupplemented yearling beef heifers on continuously-stocked Tifton 85 bermudagrass (Pedreira, 1995). Although the herbage allowance in this study was below 1 kg DM kg -1 BW in 2004, it is important to consider that calves were supplemented with at least 10 g kg -1 BW concentrate. This amount of concentrate

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79 accounts for 33% of the daily energy requirement of these animals, likely resulting in lower forage allowance requirements. Table 4-3. Herbage allowance and stocking rate of Tifton 85 pastures grazed by early weaned calves supplemented with three levels of concentrate. (BW=body weight) Concentrate Herbage allowance Stocking rate -----------g kg -1 BW----------------kg DM kg BW -1 -------------------AU ha -1 ---------10 0.97 11.1 15 0.95 11.2 20 0.81 13.7 Polynomial contrast L L P value <0.01 <0.01 SEM 0.02 0.4 AU = 500 kg LW 0.75 L=linear Hernandez Garay et al. (2004) studied the effect of N fertilization and stocking rate on weanling bulls grazing stargrass. Animal ADG decreased quadratically with increasing stocking rate from 1.3 to 3.8 AU (500 kg LW) ha -1 The LWG was maximized at stocking rates of ~ 2.5 AU ha -1 for N fertilization rates of 112 and 224 kg ha -1 over a 280-d grazing season. Pedreira (1995) verified a mean seasonal stocking rate of 5.9 AU ha -1 on Tifton 85 pastures with maximum stocking rate of 9 AU ha -1 in July 1994. The greater stocking rates in this study are the result of supplementation of the calves. The substitution of forage by concentrate resulted in more forage available for grazing, resulting in a linear increase in stocking rates with increased level of concentrate.

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80 Animal Response Variables There was linear increase in ADG and LWG as level of supplement fed increased (Table 4-4). The performance of early weaned calves grazing Tifton 85 and receiving 10 g kg -1 BW in concentrate was similar to the results presented by Vendramini et al. (2003) for early weaned calves grazing stargrass (0.59 kg d -1 ) and receiving the same amount of concentrate. When Harvey and Burns (1989) evaluated performance of EW calves grazing pearl millet [Pennistum glaucum (L.) R. Br.] with ground ear maize fed ad libitum, the ADG of the calves was 0.92 kg d -1 In the current study, average daily gain increased only 25% with an increase of 100% in amount of supplement fed. Because the level of gain observed is well below the genetic potential for these calves, it is likely that energy and/or protein limited growth of the calves. According to the NRC (1996) simulations calves grazing Tifton 85 with 10 g kg -1 BW supplementation would gain 0.9 kg of liveweight d -1 The lesser ADG observed in this study may be related to greater than expected negative associative effect of concentrate on forage intake. In addition, high temperature and humidity during the grazing season may have contributed to the relatively poor performance of the calves. The ADG and LWG of calves that did not receive concentrate supplementation were 0.33 kg d -1 and 950 kg ha -1 respectively. These values were inferior to the ADG and gain per ha of calves supplemented with 10 g kg -1 BW in concentrate, 0.52 kg d -1 and 1080 kg ha -1 respectively. There was a year*period interaction on calf ADG (Fig. 4-5). The greater ADG observed during May 2003 may have resulted from differences in gut fill during the transition of the calves from cool-season to warm-season pastures. In 2003, calves were moved from rye-ryegrass directly to Tifton 85, and a greater amount of fill associated

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81 with consumption of Tifton 85 relative to rye-ryegrass could explain the greater observed gain in May. In 2004, the calves spent 2 wk grazing warm-season grasses before the initiation of the warm-season study, decreasing the variation in fill effect between the first two weight measurements. With the exception of May 2003, there was a tendency of increasing ADG through the experimental period in both years. This may be related to generally increasing CP of pastures throughout the experimental period. Table 4-4. Average daily gain (ADG) and liveweight gain (LWG) ha -1 of early weaned calves grazing Tifton 85 pastures supplemented with different levels of concentrate. (BW=body weight) Concentrate ADG LWG -----------g kg -1 BW---------------------kg d -1 -------------------------kg ha -1 -----------10 0.52 1080 15 0.65 1450 20 0.65 1550 Polynomial contrast L L P value 0.04 <0.01 SEM 0.02 85 LWG during a period of 86 d. L = linear Using a fixed stocking rate, Rouquette et al. (2003) reported that supplementation with 0.9 kg of a mixture of maize and soybean meal increased LWG of yearling heifers on Tifton 85 pastures from 521 (unsupplemented control) to 615 kg ha -1 In the current study, gain ha -1 increased 43% when supplementation level increased from 10 to 20 g kg -1 BW. There was no difference in total and forage OMI (Table 4-5). Trends were observed, especially for forage OMI, but missing data due to bolus regurgitation weakened the statistical tests.

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82 00.20.40.60.81MayJun.Jul.Aug.MonthsADG (kg d-1) Year 2003 Year 2004**NS**NSacbbaabb Fig. 4-5. Year*period interaction effect on average daily gain of early weaned calves grazing Tifton 85 pastures and supplemented with different levels of concentrate. Year effect within period; **, P 0.01; NS= not significant. Period effect within year; means are different if followed by different letters (P 0.05). Table 4-5. Total organic matter intake (OMI) and forage OMI of Tifton 85 pastures grazed by early weaned calves supplemented with three levels of concentrate. Concentrate Total OMI Forage OMI ---------g kg -1 BW---------------------(% BW)--------------10 3.2 2.3 15 2.8 1.6 20 2.5 1.2 Polynomial contrast NS NS P value 0.45 0.21 SEM 0.4 0.4 Total organic matter intake data adjusted according to Moore et al. (1999). NS = not significant Nelson et al. (1980) reported that intake of Coastal bermudagrass hay was 23.9 g kg -1 BW for forage harvested at 4-wk regrowth. Hill et al. (1993) studied DM intake of

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83 freshly harvested Tifton 85 forage at different maturities. Intake varied from 18 to 20 g kg -1 BW for 3to 7-wk regrowth. When supplemented with maize at 10 g kg -1 BW, beef steers reduced the bermudagrass hay intake 0.46 kg for each kg of maize fed. Burns et al. (1991) compared intake of different species of warm-season grasses in North Carolina. Bermudagrass (22.3 g kg -1 BW) was the lowest compared with switchgrass (Panicum virgatum L.) and flaccidgrass (Pennisetum flaccidum Griseb.) (30.9 g kg -1 BW). The authors attributed the lower bermudagrass intake to low forage digestibility. Wheeler et al. (2002) verified increased DM intake in beef steers grazing stockpiled bermudagrass pastures when supplemented with concentrate of different CP concentrations. The intake was 21 and 23 g kg -1 BW for steers supplemented with 16 and 40% CP supplement. It is expected that calves grazing bermudagrass and consuming over 30 g kg -1 of the diet as an energy supplement would have decreased forage OMI. The data collected in this study support this trend. However, the problems experienced with the methodology limit the conclusions that can be drawn. Grazing time during daylight hours decreased linearly with increasing supplementation levels (Table 4-6). These date corroborate the results of Macoon (1999) who found that dairy cows supplemented with high levels of concentrate spent less time grazing (135 min) than cows that received low rates of concentrate (180 min). There was a significant year effect on grazing time with calves grazing 240 and 106 min for Years 2003 and 2004, respectively. The large difference in grazing time between years can be explained in part by the greater herbage mass present in 2003.

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84 Table 4-6. Grazing time during daylight hours of early weaned calves grazing Tifton 85 pastures supplemented with three levels of concentrate. (BW=body weight) Concentrate Grazing time -----------g kg -1 BW---------------------------------------min------------------------------10 193 15 181 20 147 Polynomial contrast L P value <0.01 SEM 8 L=linear There was no effect of increasing supplementation rates on calf BUN (Table 4-7). Based on the high CP concentration of the forage (180 g kg -1 ) and concentrate (160 g kg -1 ), it was expected that calves consuming more forage would have a higher BUN concentration; however, the forage rumen-degradable protein (550 g kg -1 ) was less than concentrate rumen degradable protein (700 g kg -1 ) (NRC, 1996). As a result, calves consumed similar total rumen degradable protein, resulting in similar BUN among the treatments. According to Hammond et al. (1993), cattle BUN concentrations from 9 to 12 mg dL -1 are considered to be in a transition range below which daily response to protein supplementation has been positive. The calf BUN concentrations in this study suggest that protein supplementation is unlikely to improve performance of the calves. Further studies are necessary to clarify the relation of BUN concentrations and performance of early weaned calves grazing warm-season pastures.

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85 Table 4-7. Blood urea nitrogen (BUN) concentrations of early weaned calves grazing Tifton 85 pastures supplemented with three levels of concentrate. (BW=body weight) Concentration BUN -----------g kg -1 BW-----------------------------------------mg dL -1 ----------------------------10 14.8 15 15.4 20 14.0 Polynomial contrast NS P value 0.64 SEM 0.9 NS = not significant Economic Analysis Cost and income increased linearly with increasing supplement rate (Table 4-8). The increase in income was more than compensated for by greater concentrate cost at the highest supplementation rate. There was a quadratic effect on economic return indicating that the 15 g kg -1 BW treatment provided greater return than the average of the 10 and 20 g kg -1 treatments. Table 4-8. Economic analysis of three concentrate levels for early weaned calves grazing Tifton 85 pastures. Concentrate Concentrate cost Income Return -------g kg -1 BW--------------------------------$ ha -1 --------------------------10 1100 2380 1280 15 1670 3200 1530 20 2710 3400 700 Polynomial contrast L L Q P value <0.01 <0.01 0.04 SEM 74 187 284 L = linear, Q = quadratic

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86 Summary and Conclusions The different levels of supplement fed to early weaned calves grazing Tifton 85 pastures did not alter herbage mass, accumulation, and nutritive value of the forage. Herbage allowance was slightly lower for 20 g kg -1 BW supplementation, the result of the greater number of put and take calves assigned to that treatment. Nevertheless, there was no evidence that the herbage allowance in that treatment negatively affected calf performance. Average daily gain, SR, and LWG increased with greater rates of concentrate. Greater SR was associated with a tendency for calves consuming more concentrate to consume less forage. However, a problem with calves regurgitating the marker boluses limited the number of observations and made it impossible to draw firm conclusions about intake. Calves receiving lower concentrate rates had greater grazing time during daylight, supporting the observation of a trend toward lower forage intake with higher rates of supplement. Blood urea N concentrations did not differ among treatments. This can be explained by similar rumen-degradable protein consumed by the calves among the different treatments. Although greater supplementation levels resulted in greater ADG and gain per unit of land area, economic analysis of the data did not support use of the 20 g kg -1 BW treatment. Economic return was greater for the 15 g kg -1 BW concentrate treatment than the average of the 10 and 20 g kg -1 treatments, thus the 15 g kg -1 level is recommended for EW calves grazing Tifton 85 bermudagrass.

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CHAPTER 5 BOTANICAL COMPOSITION, HERBAGE ACCUMULATION, NUTRITIVE VALUE, AND CRUDE PROTEIN FRACTIONATION OF RYE-RYEGRASS HERBAGE AT DIFFERENT REGROWTH INTERVALS AND N FERTILIZATION LEVELS Introduction Warm-season grasses dominate most of the pastures in the southeastern USA, but climatic conditions allow for use of cool-season annual grasses, either sod-seeded or in prepared seedbeds. Reasons for use of cool-season annual grasses include extending the grazing period, high nutritive value, compatibility and ease of establishment in warm-season perennial grass pastures, and tolerance to different defoliation regimens and stocking rates (Rouquette et al., 1997). Annual ryegrass (Lolium multiflorum Lam.), a high-yielding, nutritious grass, is the most commonly grown cool-season pasture forage in the southern and southeastern USA from November to May (Evers et al., 1997). Small grains are mixed with annual ryegrass to improve early season forage production. Annual ryegrass-small grain mixtures provide the longest grazing season and the most forage production if planted early on a well-prepared seedbed. However, they also are relatively expensive and are most economically used by lactating dairy cows, stocker calves, replacement heifers, or for limit grazing by beef cows nursing fall calves (Evers et al., 1997). Small grains grow better from late December to mid-February, and ryegrass results in rapid forage growth during March to late May, often requiring frequent increases in stocking rate to efficiently use the herbage produced. 87

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88 Many studies have shown the effect of N fertilization on herbage accumulation and nutritive value of cool-season forages. Allen et al. (1974) tested different levels of N in oat (Avena sativa L.)-ryegrass mixed swards. Forage yield was 4.3 Mg ha -1 and 9.0 Mg ha -1 with 0 and 280 kg N ha -1 Morris et al. (1994) reported ryegrass herbage accumulation of 10.5 Mg ha -1 when 280 kg N ha -1 was applied. Ryegrass forage in vegetative stages of growth usually has in vitro digestible organic matter (IVDOM) > 700 g kg -1 approaching and sometimes exceeding 800 g kg -1 in the first weeks of the grazing season (Ulyatt, 1981). In vitro digestibility of annual ryegrass grown at different N rates has been evaluated in Louisiana by Allen et al. (1974). The authors reported a 60 g kg -1 increase in digestibility as N fertilization levels increased from 224 to 448 kg N ha -1 The crude protein (CP) concentration of cool-season grasses is strongly influenced by the available soil N level. Application of N fertilizer to grasses usually increases CP concentration as well as crop growth. The majority of the increase in CP is non-protein N (NPN) in the form of nitrates and free amino acids (Van Soest, 1982). Crude protein concentrations in annual ryegrass tend to be high (Haby and Robinson, 1997), commonly averaging 150 to 200 g kg -1 with no N applied and increasing to 280 g kg -1 at N rates of 448 kg ha -1 Redfearn et al. (2002) analyzed nutritive value of different cultivars of ryegrass. The CP and in vitro digestible dry matter concentration (IVDDM) in January were 234, 229, and 232 g CP kg -1 and 849, 849, and 846 g IVDDM kg -1 for Gulf, Jackson, and Marshall, respectively. In general, CP concentrations of cool-season grasses range from 200 to 300 g kg -1 when well fertilized and in vegetative growth stage.

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89 High quality cool-season pastures and many tropical legumes would be expected to have high ruminal ammonia losses when their CP concentrations exceed 150 g kg -1 because of the high concentration of rumen-degradable protein (Poppi and McLennan, 1995). According to Beever (1984), 800 to 900 g kg -1 of ryegrass CP can be degraded in the rumen. Van Vuuren et al. (1991) stated that organic matter and CP in situ total degradability decreased with increasing ryegrass sward maturity and with decreasing rate of N application. Salaun et al. (1999) studied the relation between N fertilization and CP rumen degradability of perennial ryegrass (Lolium perenne L.). They concluded that reducing N fertilizer application decreased the theoretical degradability in the rumen due to decreases in NPN. The total N disappearance was 600 and 710 g kg -1 for plots fertilized with 250 and 550 kg N ha -1 In Florida beef cow-calf systems, early weaning of fall-born calves increases rebreeding of first-calf heifers and young cows (Arthington and Kalmbacher, 2003). More information is needed to develop pasture-based programs for feeding these calves during the cool season. Rye (Secale cereale L.) ryegrass mixtures are important cool-season pastures in Florida, but to achieve the desired levels of calf weight gain it is thought that some supplement must be fed. In development of supplementation programs, seasonality of forage production, botanical composition, and nutritive value responses to management are key considerations. In addition, the patterns of forage dry matter (DM) and CP degradation in the rumen can provide useful information to guide the choice of supplement ingredient and chemical composition. Therefore, as part of a research program evaluating strategies for feeding early weaned calves, this study was conducted with the objectives of evaluating the effect of N fertilization levels and regrowth interval

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90 on rye-ryegrass botanical composition, herbage accumulation, nutritive value, and in situ N fractionation and disappearance. Material and Methods The study was located at the University of Florida Beef Research Unit, 18 km northeast of Gainesville, FL (30 o N). Data were collected during the 2003 and 2004 winter-spring seasons. The soil at the research site is Plummer sand (Grossarenic Paleaquults, loamy, siliceous, thermic). Drainage of these soils ranges are from poor to moderately rapid in A and moderate in B horizons. Prior to initiation of the study, mean soil pH (1:2 soil:deionized H 2 O ratio) was 6.0, and Mehlich-I (0.05 M HCl + 0.0125 M H 2 SO 4 ) extractable P, K, Mg, and Ca in the Ap1 horizon (0to 15-cm depth) were 4, 15, 109, and 661 mg kg -1 respectively. Temperature and rainfall during the study period are described in Table 1 (Appendix A). In early October, the area was sprayed with 3.0 L ha -1 of glyphosate (Monsanto Co., St. Louis, MO; isopropylamine salt [10 g kg -1 ] of N-phosphonometyl glycine) to kill bahiagrass. Three weeks later, the pastures were overseeded with a mixture of rye-ryegrass using a John Deere 1590 no-till drill. The seeding rates were 20 kg ha -1 of Jumbo ryegrass and 80 kg ha -1 of a mixed blend rye (Grazemaster). All pastures received an initial application of 40 kg N ha -1 17 kg P ha -1 and 66 kg K ha -1 3-wk after planting. Treatments were the factorial combinations of two regrowth intervals (3 and 6 wk) and three N fertilization levels (0, 40, and 80 kg N ha -1 per season) evaluated in two seasons, winter (January-February) and spring (March-April). There were six plots per block and treatments were arranged in three randomized complete blocks. The criteria used for blocking was primarily soil drainage. All plots were staged at the beginning of

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91 the winter season (10 Jan. 2003 and 16 Jan. 2004), and the same plots were harvested during both winter and spring seasons. The second 3-wk harvest and the first 6-wk harvest of the winter season also served as the staging cut for the spring season. Plot size was 2 x 4 m with 1-m distance between plots. In this study, winter was the 6-wk period starting on 10 Jan. 2003 and 16 Jan. 2004, and spring was the 6-wk period starting on 21 Feb. 2003 and 27 Feb. 2004. The 6-wk cutting-interval plots received the entire N application for that season at the beginning of the 6-wk growth period, while the 3-wk plots received half of seasonal rate at the beginning of each of the two, 3-wk growth periods per season. Herbage was harvested to a 5-cm stubble from two representative 0.25-m 2 quadrats per plot and was dried at 60 o C for 48 h to measure herbage accumulation. Herbage accumulation data are reported by season (winter and spring), and data from a given season is the total of one harvest of the 6-wk treatment and two harvests of the 3-wk treatment. One additional 0.25-m 2 quadrat per plot was clipped, hand separated into rye and ryegrass, and used to calculate botanical composition of the swards. Laboratory Analyses For laboratory analyses, herbage from the two quadrats used to determine herbage accumulation (i.e., not separated by species) was ground in an Udy cyclone mill (Udy Corporation, Fort Collins, CO.). to pass a 4-mm screen, and stored for chemical analyses. Samples were analyzed for IVDOM using the two-stage technique described by Tilley and Terry (1974) modified by Moore and Mott (1974). Nitrogen concentration was determined using a micro-Kjeldahl method, a modification of the aluminum block digestion technique described by (Gallaher et al., 1975).

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92 In situ Disappearance Procedure Rye-ryegrass samples from a given experimental unit were composited across seasons. Four grams of dried and ground sample were weighed and placed into N-free polyester bags (pore size, 50-60 m). Bags were 20 x 10 cm in and heat sealed using an impulse sealer (model MP-8; Midwest Pacific, Co., Baltimore MD). The ratio of weight to surface area was 20 mg cm -2 Duplicate samples were incubated for 0, 3, 6, 9, 12, 24, 48, and 72 h. The bags were soaked in water, attached to a rope, and placed into a ruminally-fistulated Holstein cow. The cow was housed in a free-air circulating barn in an individual stall. The cow was fed a diet of bermudagrass [Cynodon dactylon (L.) Pers.] hay ad libitum supplemented with 0.4 kg of soybean (Glycine max L.) meal d -1 For a given incubation time, 72 bags representing all experimental units (three N levels, two regrowth periods, 2 yr, and three replications; all in duplicate) were incubated and withdrawn at the same time. This procedure guaranteed identical ruminal conditions among treatments at each incubation time. Samples were composited across seasons (within plot) to ensure sufficient forage for analyses. The 0-h bags were not incubated in the rumen but were subjected to the same rinsing procedure used for the ruminally incubated bags. After withdrawal of bags from the rumen, bags were placed in a plastic bucket with water and rinsed repeatedly until the rinse water was colorless. Bags were frozen (-20 o C), and at the end of the experiment all bags were washed at the same time in a clothes washing machine set at low water levels for one cycle. Afterwards, bags were dried at 60 o C in a forced-air oven for 48 h and weighed. No corrections were made for microbial N contamination. According to Vanzant et al. (1998), techniques to predict microbial contamination have proven unsatisfactory for universal application. The N

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93 concentration in the samples post-incubation was determined using the micro-Kjeldahl procedure described earlier. Dry Matter and Crude Protein Degradation Kinetics Crude protein fractions were estimated using an in situ ruminal degradation method where N is partitioned into fractions A, B, and C (Krishnamoorthy et al., 1983). Fraction A represents the soluble portion of the N and was assumed to be degraded rapidly and completely. It was measured as the N that washed out of the bag when rinsed with water. Fraction C was considered to be ruminally unavailable N and was the portion of N remaining in the bags following incubation for 72 h. Fraction B was calculated by difference (B = A C) and represents the insoluble, potentially degradable N. Kinetic parameters of DM disappearance were estimated using the non-linear model proposed by (McDonald, 1981). P = A + B(1-e -c(t-L) ) Where P = DM degraded at time t (g kg -1 ), A = wash loss (g kg -1 ), B = potentially degradable fraction (g kg -1 ), c = the rate at which b is degraded (g kg -1 h -1 ), t = time (h) incubated in the rumen, and L = lag time (h). The constants A, B, c, and L were estimated using the nonlinear regression procedures (SAS Institute Inc., 1996). For CP disappearance kinetics, the non-linear model proposed by Orskov and McDonald (1979) was used. This model used the same principles as the model described above, however, it does not include lag time. High solubility of the N present in coolseason grasses results in no lag time after incubation in the rumen. The estimation of potential DM and CP degradability was calculated by fixing the particle turnover at 0.06 h -1 (Cerneau and Michalet-Doureu, 1991; cited by Michalet

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94 Doureu and Ould-Bah, 1992). The model used was proposed by Orskov and McDonald (1979) [PD= a + (b x c)/(c+0.06)]. Statistical Analyses Data were analyzed using mixed model methodology through MIXED procedure (SAS Institute Inc., 1996). Nitrogen fertilization, regrowth interval, year, and season were considered fixed effects and block and its interactions were considered random effects. The means reported in this text are least squares means. For botanical composition, herbage accumulation, CP, and IVDOM, the model used was: Y ijkl = + A i + N j + R k + S l + B m + (AN) ij + (AR) ik + (AS) il + (NR) jk + (NS) jl +(RS) kl + (ANR) ijk + (ANS) ijl + (NRS) jkl + (ANRS) ijkl + e ijkl Where Y ijkl is the dependent variable is the overall mean N j is the N effect (main plot) R k is the regrowth interval effect (main plot) A i is the year effect (sub-plot) S l is the season effect (sub-sub-plot) B m is the block effect (NR) jk is the N*regrowth intervasl interaction (AN) ij is the year*N interaction (AR) ik is the year*regrowth interval interaction (AS) il is the year*season interaction (NS) ijl is the N*season interaction (RS) kl is the regrowth interval*season interaction

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95 (ANR) ijk is the year*N*regrowth interval interaction (ANS) ijl is the year*N*season (ARS) ikl is the year*regrowth interval*season interaction (NRS) jkl is the N*regrowth interval*season interaction (ANRS) ijkl is the year*N*regrowth interval*season interaction e ijkm is the error Main plot (N and Regrowth interval) error was block*N*regrowth interval; sub-plot (year) error was block*year; sub-sub-plot (season) error was the residual. For the DM and N in situ fractionation response variables the model used was: Y ijkl = + A i + N j + R k + B m + (AN) ij + (AR) ik + (NR) jk + (ANR) ijk + e ijk Where Y ijkl is the dependent variable is the overall mean N j is the N effect (main plot) R k is the regrowth interval effect (main plot) A i is the year effect (sub-plot) B m is the block effect (AN) ij is the year*N interaction (AR) ik is the year*regrowth interval interaction (NR) jk is the N*regrowth interval interaction (ANR) ijk is the year*nitrogen*regrowth interval interaction e ijkml is the error Main plot (N, regrowth interval) error was block*N*regrowth interval; sub-plot (year) error was the residual.

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96 Results and Discussion Botanical Composition, Herbage Accumulation, and Nutritive Value Botanical composition was not affected by N fertilization level or regrowth interval; however, there was a season*year interaction (Table 5-1). In 2003, rye was predominant in the canopy during the winter and ryegrass was predominant during the spring. However, the low rainfall from March through May 2004 (92 mm vs. the 30-yr average of 274 mm) reduced ryegrass contribution during spring 2004, and there was no seasonal difference in ryegrass percentage. Fontaneli et al. (2000) observed ~ 80% rye and 15% ryegrass in the canopy in early winter (December February) 1996. In early spring (March May), there was a greater proportion of ryegrass (60%) than rye (30%). According to Rouquette et al. (1997), small grains grow better from late December to mid-February and ryegrass growth is rapid during March to late May. There were effects of N level, season*regrowth interval interaction, and year season interaction on herbage accumulation. Herbage accumulation increased linearly from 0.9 to 1.8 Mg ha -1 at the 6-wk regrowth interval as N increased from 0 to 80 kg N ha -1 (data not shown). Allen et al. (1974) tested different levels of N fertilization on oat-ryegrass mixed swards. Forage yield was 4.3 Mg ha -1 and 9.0 Mg ha -1 with 0 and 280 kg N ha -1 yr -1 In studies where the maximum N rate was below 450 kg ha -1 the greatest yields have commonly occurred at the highest N rate applied (Morris et al., 1994). There was a season*regrowth interval interaction for herbage accumulation (table 5-2). Interaction occurred because during the winter period there was no effect of regrowth interval on accumulation, but in spring accumulation was greater for the 6-wk interval. At both intervals, spring herbage accumulation was greater than winter. Craigmiles and Weihing (1971) harvested annual ryegrass every 1, 2, 3, or 4 wk

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97 beginning in January. Highest yields were obtained by harvesting every 4 wk, with average forage yields of 1 Mg ha -1 per 4-wk period. Altom et al. (1996) compared the herbage accumulation of rye-wheat-ryegrass at different N rates during fall-winter and spring. Herbage accumulation was 1 and 1.2 Mg ha -1 (0 kg N ha -1 ) and 1.1 and 3.2 Mg ha -1 (110 kg N ha -1 ) for fall-winter and spring accumulation, respectively. In the current study, lesser herbage accumulation in January and February likely was caused by the lower temperatures (Table 1, Appendix A). In contrast, during March-April, warmer temperatures resulted in greater herbage accumulation. There also was a year*season interaction for herbage accumulation (Table 5-1). Herbage accumulation was greater during the spring than winter both years; however, interaction occurred because the magnitude of the increase from winter to spring was greater in 2004 than in 2003. Extremely high rainfall in 2003 (209 mm above average for February and March) likely reduced the efficiency of N fertilizer utilization and resulted in lesser herbage accumulation that year. Although herbage accumulation was greater in 2004, the season effect was similar to that in 2003. In general, herbage accumulated during winter was ~ 50% of that during the spring. Cuomo et al. (1999) studied herbage accumulation of ryegrass established in different warm-season grass residues. There was a significant effect of season for all treatments. Average forage accumulation across treatments and years ranged from 0.2 Mg ha -1 on January to 3.0 Mg ha -1 on May. When annual ryegrass is planted using no-till systems, forage production has been inconsistent, particularly early in the growing season (Cuomo et al., 1999). However, greater consistency can be gained in no-till annual ryegrass production when warm-season annual

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98 grass swards are treated with glyphosate 30 d before planting or where the residue was burned before planting (Cuomo et al. 1999). There were effects of N*season interaction, regrowth interval*season interaction, and year*season interaction on herbage CP and IVDOM (Table 5-3). There was a linear increase in CP concentrations in both seasons as N fertilization levels increased. Nitrogen*season interaction occurred because the range in the response to N was greater in the winter than spring. In addition, except for 0 N level, CP concentrations were always greater in winter than in spring. This can be attributed to the dilution effect of greater herbage mass produced in the spring that resulted in lesser CP concentrations. There was a linear increase in IVDOM during the winter as N level increased, however, there was only a trend (P = 0.11) toward fertilization level effect on IVDOM during the spring. Appearance of ryegrass reproductive tillers likely diminished the effects of N fertilization level on IVDOM concentration. There was a significant regrowth interval*season interaction effect on CP and IVDOM (Table 5-2). During the winter, there was a small decrease in CP as regrowth interval increased, however, during the spring, because of the appearance of reproductive tillers, the rate of decline with increasing regrowth interval was more rapid. Crude protein concentrations of the herbage harvested at 6 wk in the winter and at 3 wk in the spring were similar. There was no difference in IVDOM concentrations for 3-wk and 6-wk regrowth intervals during the winter, but there was a significant decrease in IVDOM with increasing regrowth interval observed during the spring. There was a trend (P = 0.06) for herbage harvested at the 3-wk regrowth interval to have lesser IVDOM (30 g kg -1 ) in the spring than in the winter, but for the 6-wk regrowth interval, IVDOM concentration was

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99 much less (150 g kg -1 ) in the spring than in the winter. Data from Buxton and Russell (1988) for forages sampled following the beginning of reproductive development indicate a decline in IVDOM from 761 to 552 g kg -1 in orchardgrass ( Dactylis glomerata L.) leaves from May to July. Buxton and Merten s (1989) reported that herbage IVDOM of four cool-season grass species decreased linearly with time during the spring. Plants with moderate number of reproductiv e tillers had an average decline in degradability of 3.9 g kg -1 d -1 Year*season interaction effects were obs erved in CP and IVDOM concentrations (Table 5-1). The extremely high rainfall duri ng the winter of 2003 resulted in herbage with lesser CP concentrations than the he rbage harvested during the winter of 2004. The CP concentration was greater during the spring of 2003 than the spring of 2004. In addition, CP concentrations were always greater in the winter than in the spring. Greater herbage accumulation was always associated with lesser CP concentrations because of the dilution effect mentioned before. The same pattern of response was observed for IVDOM concentrations as already described for CP. Protein Fractionation of the Forage The CP fractions in rye-ryegrass herbage are presented as a proportion of the total CP. There was a significant effect of N fer tilization level on concentrations of the CP fractions in rye-ryegrass herbage (Fig. 5-1) Fraction A increased linearly and Fraction B decreased linearly as level of N fertil ization increased from 0 to 80 kg N ha -1 In addition, there was a linear decrease in Fraction C from 73 g kg -1 to 47 g kg -1 as N fertilization level increased from 0 to 80 kg ha -1

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100 Table 5-1. Year*season interaction forage effects on botanical composition, crude protein (CP), and in vitro digestible organic matter (IVDOM) of rye-ryegrass mixtures. Year Ryegrass SE P Herbage accumulation SE P CP SE P IVODM SE P Winter Spring Winter Spring Winter Spring Winter Spring --------%----------Mg ha -1 -----------------------g kg -1 ------------------2003 47 95 3.5 <0.01 0.6 1.2 0.1 <0.01 200 170 6 <0.01 740 700 9 <0.01 2004 30 35 3.5 0.22 1.2 2.2 0.1 <0.01 220 150 6 <0.01 770 640 9 <0.01 P value <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 SE 4 4 0.1 0.1 6 6 9 9 As a proportional of total herbage accumulation on a dry matter basis P value for effect of season within year P value for effect of year within season

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101 Table 5-2. Season*regrowth interval interaction effects for herbage accumulation and crude protein (CP) and in vitro digestible organic matter (IVDOM) concentrations of rye-ryegrass forage mixtures during 2003 and 2004. Regrowth interval Herbage accumulation SE P CP SE P IVDOM SE P Winter Spring Winter Spring Winter Spring ----wk-------Mg ha -1 ----------------------g kg -1 -----------------3 0.9 1.6 0.1 <0.01 220 200 7 0.02 760 730 12 0.06 6 1.0 2.2 0.1 <0.01 200 130 7 <0.01 750 600 12 <0.01 P 0.70 <0.01 <0.01 <0.01 0.73 <0.01 SE 0.1 0.1 7 7 12 12 P value for effect of season within cutting interval P value for effect of cutting interval within season

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102 Table 5-3. N*season interaction effects on crude protein (CP) and in vitro digestible organic matter (IVDOM) concentrations of rye-ryegrass forage. Season CP SE OPC IVDOM SE OPC N fertilization (kg ha -1 ) N fertilization (kg ha -1 ) 0 40 80 0 40 80 -----------g kg -1 ------------------g kg -1 --------Winter 150 210 260 7 L 720 770 770 24 L Spring 130 160 200 7 L 660 660 690 16 0.11 P 0.06 <0.01 <0.01 <0.01 <0.01 <0.01 SE 9 9 9 16 16 16 OPC = orthogonal polynomial contrast for effect of N fertilization levels within season L = linear (P<0.01) P for linear contrast for effect of N fertilization on IVDOM in spring P for effect of season within N fertilization levels. The concentrations of Fraction A in rye-ryegrass swards in this study are in the range reported by Hoffman et al. (1993b) for five species of cool-season grasses (from 370 to 600 g kg -1 ). Similar values of Fractions A, B, and C (430, 490, and 80 g kg -1 respectively) were observed by Michalet-Doureu and Ould-Bah (1992) with perennial ryegrass. Salaun et al. (1999) studied the relationship between N fertilization and CP rumen degradability of perennial ryegrass. They concluded that reducing N fertilizer application decreased the theoretical degradability in the rumen due to the decrease in NPN. The total N disappearance was 600 and 710 g kg -1 for plots fertilized with 250 and 550 kg N ha -1 Van Vuuren et al. (1991) observed increased levels of instantly degradable protein of perennial ryegrass as N fertilization levels increased from 0 to 700 kg ha -1

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103 4105045935154393597456480200400600800100004080N fertilization (kg ha-1)Fraction concentration (g kg-1 CP) A B C Fig. 5-1. Nitrogen fertilization effects on crude protein (CP) fraction concentrations in total CP of rye-ryegrass forage. There was a linear increase in Fraction A (P<0.01, SE = 28), and a linear decrease in Fraction B (P<0.01, SE = 23) and Fraction C (P<0.05, SE =12). There was significant regrowth interval effect on Fraction C concentration. Forage harvested at 3 wk (48 g kg -1 ) had less Fraction C concentration than that harvested at 6 wk (71 g kg -1 ). According to Van Vuuren and Meijs (1987), CP of highly-fertilized young grass is characterized by a high rate of rumen degradation that will result in substantially greater ruminal ammonia concentrations and potentially greater losses of N via urine. The greater Fraction A associated with greater N fertilization levels in this study suggest that this may occur.

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104 Degradation Parameters of DM and CP There was no effect of N fertilization level or regrowth interval on in situ DM degradation rates, lag time, and potential DM degradability (Table 5-4). Cool-season grasses generally have less DM disappearance lag time than warm-season grasses. For highly-digestible warm-season forages like elephantgrass (Pennisetum purpureum Schum.), Vieira et al. (1997) found lag times values of 4 h. With cool-season grasses, Waghorn and Burke (2001) reported values ranging from 0 to 4.5 h. The DM disappearance lag time in this study was 3 h. Although there was a lag for DM disappearance, there was none for CP disappearance (Table 5-4). A difference in the most soluble fraction of the CP was observed among fertilization levels from 0 to 20 h of incubation, but CP disappearance was similar among treatments thereafter (Fig. 5-2). Newman et al. (2002a) observed a linear increase in lag time from 16 to 21 h as canopy height of continuously stocked limpograss increased. It is expected that warm-season grasses have longer lag time and slower N disappearance because of the lower soluble protein concentration than in cool-season grasses, which is thought to be mainly due to differences in Rubisco concentration (Akin and Burdick, 1975). Furthermore, much of the protein in warm-season grasses may be protected structurally because of its association with bundle-sheath cells (Mullahey et al., 1992). There was a linear increase in potential CP degradability of the forage as N fertilization level increased, however, no effect of N levels were verified on CP lag time and rate of disappearance (Table 5-5). Potentially degradable CP was greater in 2004 (780 g kg -1 ) than in 2003 (690 g kg -1 ). The average CP rate of disappearance in this study is similar to rates described by Krishnamoorthy et al. (1983) for cool-season grasses and

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105 legumes, 0.05 h -1 Other authors have found greater rates of disappearance for legumes. Hoffman et al. (1993b) reported protein disappearance rates of 0.40 h -1 for birdsfoot trefoil (Lotus corniculatus L.) in the late-bud stage of growth, and Brown and Pitman (1991) found an average of 0.43 h -1 for different species of tropical legumes. There was no significant effect of the N fertilization levels in this study on DM and CP rate of disappearance. Van Vuuren et al. (1991) found significant increases in OM (0.03-0.06 h -1 ) and CP (0.05-0.09 h -1 ) disappearance rates of perennial ryegrass as N fertilization level increased from 0 to 700 kg ha -1 Michalet-Doureu and Ould-Bah (1992) observed rate of CP degradation and potential N degradability of perennial ryegrass herbage of 0.1 h -1 and 740 g kg -1 respectively. Table 5-4. Effect of N fertilization levels on DM and CP degradation parameters of rye-ryegrass forage. N fertilization levels (kg ha -1 ) SE OPC P 0 40 80 DM Lag time (h) 2.7 3.1 3.4 1.2 0.54 Rate of disappearance (h -1 ) 0.08 0.09 0.16 0.06 0.21 Potential degradability (g kg -1 ) 670 670 690 23 0.50 CP Lag time (h) 0 0 0 Rate of disappearance (h -1 ) 0.08 0.06 0.06 0.01 0.14 Potential degradability (g kg -1 ) 700 720 770 9 L OPC = orthogonal polynomial contrast L=linear (P<0.01)

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106 200300400500600700800900100001020304050607080HoursCP disappearance (g kg-1CP) N 0 N 40 N 80 Fig. 5-2. N fertilization rate on crude protein (CP) disappearance of rye-ryegrass forage. Summary and Conclusions Botanical composition of rye-ryegrass swards was affected by season with the proportion of rye being greater during the winter and proportion of ryegrass greater during the spring of a high rainfall year. During a drier year there was no difference in seasonal botanical composition due to reduced contribution from ryegrass in spring. Nitrogen fertilization and regrowth interval did not affect botanical composition. Herbage accumulation increased linearly with increased levels of N fertilization. More herbage accumulated with the 6-wk than the 3-wk regrowth interval in spring, but during winter, regrowth interval did not affect total accumulation. Accumulation was greater in spring than winter regardless of regrowth interval. The effect of regrowth interval on herbage CP and IVDOM concentrations varied with season. There was no difference in IVDOM during the winter for the two regrowth intervals, however, lesser IVDOM concentrations were observed for the 6-wk than the 3-wk regrowth interval during the spring. Herbage CP concentrations were greater for the

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107 3-wk regrowth interval during the winter and spring, but the difference was much greater in spring. Thus, regrowth interval is of much greater importance in determining forage nutritive value and herbage accumulation during spring than winter and during the spring season, greater N fertilization levels and shorter regrowth periods than in winter may be necessary to maintain the nutritive value of the forage. There was a significant effect of N fertilization level on the CP fractions of rye-ryegrass. Increased N fertilization increased the concentration of Fraction A and decreased Fraction B and C. More Fraction A at greater fertilization levels has the potential to result in less efficient use of N by ruminants that can lead to greater N losses through the urine. No lag time was observed for in situ CP disappearance indicating that the CP present in rye-ryegrass herbage is highly soluble. Greater N fertilization increased the potential CP disappearance linearly. When considering utilization of rye-ryegrass herbage for early weaned calves, the greater total CP and Fraction A concentrations in rye-ryegrass with increased levels of N fertilization implies that there may be an excess of ruminal degradable protein under these circumstances. Reduced N fertilization levels lower herbage accumulation and may not be a practical strategy to address this problem especially during a time of the year when quantity of forage is often the primary factor limiting animal performance. Utilization of high energy, low protein supplements may be a useful management practice to improve better N and energy synchronization in the rumen and increase the efficiency of rumen degradable protein utilization by early weaned calves grazing cool-season grass pastures.

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CHAPTER 6 HERBAGE ACCUMULATION, NUTRITIVE VALUE, AND CRUDE PROTEIN FRACTIONS OF TIFTON 85 SWARDS AT DIFFERENT CUTTING INTERVALS AND N FERTILIZATION LEVELS Introduction Bermudagrass is an important forage for beef and dairy cattle in the southern USA (Hill et al., 1998). Tifton 85 bermudagrass (Cynodon spp.) has been widely grown in the USA, central and South America, and southern Africa (Mandebvu et al., 1999). When compared with Coastal and Tifton 78bermudagrass, Tifton 85 is higher yielding and more digestible (Hill et al., 1993). Warm-season grasses have larger proportions of cell-wall material compared with C3 grasses. This fraction is potentially digestible, but because of its chemical composition and anatomical structure, the rate of degradation and fermentation in warm-season grasses is relatively slow (Coleman et al., 2004). On the other hand, warm-season grasses have more efficient carbon fixation, resulting in greater total forage production when compared with cool-season grasses. Bermudagrass herbage yield and nutritive value are dependent upon management practices, including N fertilization and regrowth interval. According to Wilkinson and Langdale (1974), Coastal and other bermudagrass cultivars produced maximum yields of about 27 Mg ha -1 yr -1 at N rates of 1200 kg ha -1 with near linear yield response to N up to about 600 to 700 kg N ha -1 Prine and Burton (1956) evaluated the effects of different N rates and harvest frequencies on Coastal bermudagrass forage production and nutritive value. There was a curvilinear increase in herbage accumulation as N rate increased from 108

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109 0 to 1008 kg N ha -1 yr -1 with maximum dry matter (DM) accumulation of 18.4 Mg ha -1 yr -1 with a 6to 8-wk harvest interval. The forage crude protein (CP) concentration ranged from 97 to 190 g kg -1 across the range of N fertilization rates. Burton et al. (1963) studied different defoliation intervals of Coastal fertilized with 660 kg N ha -1 yr -1 The maximum herbage accumulation was 22 Mg ha -1 using a 6-wk defoliation interval. Warm-season grass CP tends to be more slowly degraded in the rumen than that of cool-season grasses (Minson, 1990), and CP concentration often does not meet the nutrient requirement of growing ruminants (Moore, 1992). The C4 grasses have a lower soluble protein concentration (260 to 300 g kg -1 ) than C3 grasses (330 to 480 g kg -1 ). This is thought to be mainly due to lower Rubisco concentration and to the relative abundance of Rubisco in the slowly or incompletely-digested bundle sheath cells of C4 plants vs. the prevalence of Rubisco in the rapidly degraded mesophyll of C3s (Akin and Burdick, 1975). Redfearn et al. (1995) studied the rates of ruminal protein disappearance for switchgrass (Panicum virgatum L.), big bluestem (Andropogon gerardii Vitman), and smooth bromegrass (Bromus inermis Leyss.) and suggested that generalizations regarding ruminal protein degradability should not be made among forage species. The total protein disappearance was 847, 795, and 974 mg g -1 for switchgrass, big bluestem, and smooth bromegrass, respectively. Newman et al. (2002a) fractionated the N content of limpograss (Hermathria altissima Stapf. And Hubb.). They concluded that low total CP in the DM, low concentrations of CP Fractions A and B in the rumen, and the long lag phase for degradation of the B fraction, may contribute to reported protein deficiencies of cattle grazing limpograss. Johnson et al. (2001b) reported that with no N fertilization, Fraction A concentration was 212 (bahiagrass[Paspalum notatum Flgge]), 310 (bermudagrass),

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110 and 279 g kg -1 (stargrass [Cynodon lnemfuensis Vanderyst]) of the total N, but upon application of 160 kg N ha -1 per cutting, Fraction A represented 285 (bahiagrass), 400 (bermudagrass), and 421 g kg -1 (stargrass) of the total N. Assis et al. (1999) studied the effects of N fertilization levels on CP fractions of Tifton 85. Fraction A ranged from 270 to 300 g kg -1 with no significant effect of fertilization levels from 0 to 400 kg N ha -1 In Florida cow-calf systems, early weaning has been an effective management practice to increase the rebreeding rates of first-calf heifers. Tifton 85 bermudagrass is an important warm-season grass in southeast USA that could be used in pasture-based systems for early weaned calves. However, warm-season grasses usually have low nutritive value and are incompatible with the high nutritional requirements of early weaned calves. In order to maximize the performance of these calves on pastures, an effective supplementation program must be developed to overcome the limitations of the forage. For development of a supplementation program it is important to know seasonal patterns of herbage accumulation and nutritive value. In addition, measurements of DM and CP degradation in the rumen can provide useful information to guide the choice of supplement composition. Therefore, as part of a research program evaluating strategies for feeding early weaned calves, this study was conducted to evaluate the effect of N fertilization level and regrowth interval on herbage accumulation, nutritive value, and CP in situ fractions of Tifton 85 bermudagrass swards. Material and Methods The plot study was conducted at the University of Florida Beef Research Unit, 18 km northeast of Gainesville, FL (30 o N). Data were collected during 2003 and 2004. The soils at the research site are Adamsville fine sand (Aquic Quartzipamments, hyperthermic, uncoated) and Sparr fine sand (Grossarenic Paleudult, hyperthermic, loamy

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111 siliceous). These soils are moderately well-drained with rapid permeability. Prior to initiation of summer grazing at the site, mean soil pH (1:2 soil:deionized H 2 O ratio) was 5.5, and Mehlich-I (0.05 M HCL + 0.0125 M H 2 SO 4 ) extractable P, K, Mg, and Ca concentrations in the Ap1 horizon (0to 15-cm depth) were 44, 17, 40, and 328 mg kg -1 respectively. Average annual temperature and rainfall are described in Table 1 (Appendix A). Treatments were the factorial combinations of two regrowth intervals (2 and 4 wk) and three N fertilization levels (0, 40, and 80 kg ha -1 in each 4-wk period) evaluated in two periods of 4 wks in each of 2 yr. Treatments were replicated three times in a completely randomized design. Plot size was 2 x 4 m with a 1 m border between plots. In early April of 2003 and 2004, plots received an initial application of 40 kg N ha -1 17 kg P ha -1 and 66 kg K ha -1 to stimulate growth and provide maintenance P and K. Plots were staged to a 15-cm stubble on 20 June 2003 and 21 May 2004 to begin the first 4-wk period. The first experimental period was from 20 June to 18 July 2003 and 21 May to 18 June 2004. The second 4-wk period followed immediately after the harvests on 18 July 2003 and 18 June 2004 and ended on 15 Aug. 2003 and 16 July 2004. The 2-wk treatment plots were harvested twice in each 4-wk period and the 4-wk treatment plots once. Forage samples were harvested at a 15-cm stubble height using a sickle bar mower. The central 1-m 2 area of the plot was harvested, the wet weight recorded, and a subsample dried at 60 o C for 48 h. After drying, the subsample was weighed and ground in an Udy mill (Udy Corporation, Fort Collins, CO.) to pass a 4-mm screen. Herbage accumulation, laboratory analyses, and in situ determinations were the same as described in Chapter 5.

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112 Dry Matter and Crude Protein Degradation Kinetics The DM and CP degradation kinetics measurements were similar to those reported in Chapter 5. The exception was that DM and CP disappearance were estimated using the non-linear model proposed by (McDonald, 1981). Unlike the cool-season C3 grasses, warm-season C4 grasses usually present a lag time for in situ CP disappearance. P= A + B(1-e -c(t-L) ) In this model, P= DM degraded at time t (g kg -1 ), A = wash loss (g kg -1 ), B = potentially degradable fraction (g kg -1 ), c = the rate at which b is degraded (g kg -1 h -1 ), t = time (h) incubated in the rumen, and L = lag time (h). The constants A, B, c, and L were estimated using the nonlinear regression procedures of SAS, version 8 (SAS Institute Inc., 1996). Potential DM and CP disappearance were calculated by fixing the particle turnover at 0.04 h -1 The model used to estimated the potential CP disappearance was proposed by Orskov and McDonald (1979) [PD= a + (b x c)/(c+0.04)]. Statistical Analyses Data were analyzed using mixed models methodology through PROC MIXED (SAS Institute Inc., 1996). Nitrogen fertilization, regrowth interval, year, and period were considered fixed affects, and replicates and their interactions were considered random effects. The nature of the response to N was determined using orthogonal polynomial contrasts. The means reported are least squares means. For herbage accumulation, CP, and IVOMD, the model used was: Y ijkl = + A i + N j + R k + P l + (AN) ij + (AR) ik + (AP) il + (NR) jk + (NP) jl + (RP) kl + (ANR) ijk + (ANP) ijl +(NRP) jkl +(ANRP) ijkl + e ijkl Where Y ijkl is the dependent variable

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113 is the overall mean N j is the N effect (main plot) R k is the regrowth interval effect (main plot) A i is the year effect (sub-plot) P l is the period effect (sub-sub-plot) (AN) ij is the year* N interaction (AR) ik is the year*regrowth interval interaction (AP) il is the year*period interaction (NP) ijl is the N*period interaction (RP) kl is the regrowth interval*period interaction (ANR) ijk is the year*N*regrowth interval interaction (ANP) ijl is the year*N*period interaction (ARP) ikl is the year*regrowth interval*period interaction (NRP) jkl is the N*regrowth interval*period interaction (ANRP) ijkl is the year*N*regrowth interval*period interaction e ijkl is the error Main plot (N, regrowth interval, and N*regrowth interval) error was replicate*N*regrowth interval; sub-plot (year) error was replicate*year; sub-sub-plot (season) error was the residual. For the DM and N in situ fractionation response variables, season was not included in the model because samples from a given experimental unit were composited across seasons. The model used was: Y ijkl = + A i + N j + R k + (AN) ij + (NR) jk + (AR) ik + (ANR) ijk + e ijk

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114 Where Y ijkl is the dependent variable is the overall mean N j is the N effect (main plot) R k is the regrowth interval effect (main plot) A i is the year effect (sub-plot) (AN) ij is the year*N interaction (AR) ik is the year*regrowth interval interaction (NR) jk is the N*regrowth interval interaction (ANR) ijk is the year*N*regrowth interval interaction e ijk is the error Main plot (N, regrowth interval, and N*regrowth interval) error was replicate*N*regrowth interval; sub-plot (year) error was the residue. Results and Discussion Herbage Accumulation and Nutritive Value There were N-rate main effect on herbage accumulation. That resulted from a linear increase (1.6-2.6 Mg ha -1 per 4-wk period) in herbage accumulation as levels of N fertilization increased from 0 to 80 kg ha -1 (Table 6-1). Prine and Burton (1956) reported a curvilinear increase in herbage accumulation as N rate increased from 0 to 1008 kg N ha -1 The highest efficiency in production (kg DM kg -1 N ha -1 ) occurred at 336 kg N ha -1 which approximately corresponds to the highest N rate in the current study if one extends the current 8-wk trial to four cuttings at 4-wk intervals over an entire growing season. Wilkinson and Langdale (1974) reported linear yield responses of several bermudagrass cultivars to N rates of 600 to 700 kg ha -1

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115 The year*regrowth interval interaction occurred because there was no difference in herbage accumulation between 2-wk (1.3 Mg ha -1 ) and 4-wk (1.4 Mg ha -1 ) intervals in 2003, but there was a difference in 2004 (2.3 vs. 3.5 Mg ha -1 for 2and 4-wk intervals, respectively (data not shown). The reason for lack of a regrowth interval effect in 2003 is not clear, and results of most previous studies are more like 2004 results in the current study. Prine and Burton (1956) observed greater herbage accumulation when Coastal bermudagrass was harvested from 6to 8-wk of regrowth compared to shorter intervals. Mandebvu et al. (1999) reported bermudagrass herbage accumulation of 5.8 Mg ha -1 at a 7-wk cutting intervals compared with 2.2 Mg ha -1 at 3 wk. Burton et al. (1963) studied different defoliation intervals of Coastal fertilized with 660 kg N ha -1 Maximum herbage accumulation occurred when using a 6-wk defoliation regimen. There was a year*period interaction for herbage accumulation (data not shown). There was no difference in herbage accumulation between the periods in 2003 (1.4 vs. 1.2 Mg ha -1 ), however, in May to June 2004 Tifton 85 accumulated more DM than during June to July (3.7 vs. 2.0 Mg ha -1 respectively). The pattern of Tifton 85 herbage accumulation has not been consistent across periods of the growing season in the literature, and this is likely due to changes in year-to-year and location-to-location weather conditions. Mandebvu et al. (1999) did not find differences in herbage accumulation of Tifton 85 from July to September. The authors suggested that weather conditions are important factors affecting variation in herbage accumulation through the season. There was no indication that weather conditions affected herbage accumulation during the two periods per year in the current study. It is likely that the greater herbage

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116 accumulation during May to June 2004 was due in part to this periods proximity in time to the initial fertilization made in April. Table 6-1. N fertilization effects on herbage accumulation and in vitro digestible organic matter (IVDOM) of Tifton 85 forage during 42-d periods. Variable N fertilization (kg ha -1 period -1 ) SE OPC 0 40 80 ---------------------Mg ha -1 --------------------Herbage accumulation 1.6 2.2 2.6 0.1 L IVDOM 480 500 530 12 L OPC = orthogonal polynomial contrast L = linear (P<0.01) There was a linear increase in IVDOM with increasing N fertilization level (Table 6-1). In addition, there was a N fertilization* regrowth interval interaction in CP concentration of Tifton 85 (Table 6-2). At both regrowth intervals, there was a linear increase in CP concentration with increasing N fertilization, but interaction occurred because the rate of increase was greater for the 2-wk than the 4-wk interval. The CP concentrations were greater at 2than 4-wk regrowth intervals at all N fertilization levels tested. Johnson et al. (2001b) reported a linear increase in bermudagrass CP concentration with increaseing levels of N fertilization. Herbage CP concentrations were 98, 146, and 180 g kg -1 for fertilization levels of 0, 78, and 156 kg N ha -1 Similar results were found by Lima et al. (1999b) with limpograss. Increasing the N fertilization levels from 50 to 150 kg ha -1 increased CP concentrations from 97 to 115 g kg -1 Stallcup et al. (1986) reported that CP concentrations in bermudagrass pastures fertilized with 0 and 50 kg N ha -1 increased from 114 to 143 g kg -1

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117 Table 6-2. Regrowth interval*N fertilization level interaction effects on CP concentrations of Tifton 85 forage. Regrowth interval N fertilization SE OPC (wk) (kg ha -1 period -1 ) 0 40 80 ------------------g kg -1 ---------------2 140 155 190 6 L 4 115 120 135 6 L P <0.01 <0.01 <0.01 SE 6 6 6 OPC = orthogonal polynomial contrast L = linear (P<0.01) P regrowth interval within N fertilization level Mandebvu et al. (1999) tested the effects of regrowth interval on in vitro digestible dry matter (IVDMD) and CP concentrations of Tifton 85 swards and found a linear decrease in IVDMD of Tifton 85 from a 3-wk (620 g kg -1 ) to a 7-wk (510 g kg -1 ) regrowth period. In addition, a linear decrease in CP concentration was observed from 210 to 136 g kg -1 for 3and 7-wk regrowth herbage. Protein Fractionation of the Forage Nitrogen fertilization levels did not affect Fraction A in Tifton 85 herbage. However, there was a linear increase in fraction B and a linear decrease in fraction C as level of N fertilization increased (Fig. 6-1 A). Johnson et al. (2001b) observed a linear increase in Fraction A (310-400 g kg -1 ) as N fertilization of bermudagrass increased from 0 to 157 kg N ha -1 According to Follett and Wilkinson (1995), increased level of N

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118 fertilization increases nitrate accumulation in the plant, and nitrate is a component of Fraction A. In contrast, Assis et al. (1999) tested two N fertilization levels (0 and 400 kg ha -1 ) on Tifton 44 and Tifton 85 and did not find differences in Fraction A concentration. Fraction A was 304 and 302 g kg -1 for Tifton 44, and 270 and 300 g kg -1 for Tifton 85 at N rates of 0 and 400 kg ha -1 respectively. In this study, there was a linear decrease in Fraction C of Tifton 85 as fertilization increased from 0 to 80 kg N ha -1 Similar results were found by Johnson et al. (2001b) who reported a linear decrease (80 to 60 g kg -1 ) in bermudagrass Fraction C as N fertilization level increased from 0 to 157 kg ha -1 Regrowth interval did not affect concentrations of Fractions A and B, but a smaller Fraction C was observed in Tifton 85 at the 2(207 g kg -1 ) than at the 4-wk (282 g kg -1 ) interval (Fig. 6-1 B). Lima et al. (1999a) reported a greater proportion of the CP linked to the acid detergent fiber (ADF) and unavailable for ruminal digestion for Tifton 85 at 8than 4-wk regrowth. The values for Fraction C at 4-wk regrowth found in this study were similar to those reported by Malafaia et al. (1997) for palisadegrass (Brachiaria brizantha Stapf.) at 4 wk of regrowth (280 g kg -1 ) but greater than the values found for Tifton 85 (170 g kg -1 ).

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119 A3813453433304064552832432080100200300400500600700800900100004080N fertilization (kg ha-1)Fraction concentration (g kg-1CP) C B A B3843294023922072820100200300400500600700800900100024Regrowth interval (wk)Fraction concentration (g kg-1CP) C B A Fig. 6-1. N fertilization (Fig. A) and regrowth interval (Fig. B) effects on crude protein (CP) fraction concentrations in total CP of Tifton 85 forage. There was no effect of N fertilization on Fraction A (P>0.10 SEM=23), a linear increase (P<0.05, SEM=30) in Fraction B, and a linear decrease (P<0.01, SEM=11) in Fraction C. There was no effect of regrowth interval on Fractions A and B. Fraction C was different (P<0.05) between intervals. Disappearance Parameters of DM and CP There was no effect of N fertilization on rate of DM disappearance and lag time, however, a linear increase in potentially degradable DM was observed with increasing N

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120 fertilization levels (Table 6-3). There was a significant effect of regrowth interval on rates of DM disappearance and potentially degradable DM (Table 6-4). In contrast to these results, Mandebvu et al. (1999) did not find differences in rate of DM disappearance (0.03 h -1 ) or potential degradability (650 g kg -1 ) of Tifton 85 at regrowth intervals from 3 to 7 wk. Newman et al. (2002a) did not find differences in lag time (9 h) or rate of disappearance (0.06) for limpograss herbage from continuously stocked swards grazed at different canopy heights, however, a linear decrease in potentially degradable DM was observed as canopy height increased from 20 to 60 cm. Assis et al. (1999) tested the effect of N fertilization levels of 0 and 400 kg ha -1 on DM digestibility parameters of Tifton 85. They found no significant effect of N fertilization level on lag time (3.5 h) or potentially degradable DM (400 g kg -1 ). The DM disappearance lag time found in this study (5.9 h) was greater than those reported by Vieira et al. (1997) for elephantgrass (Pennisetum purpureum Schum.) (3 h) and shorter than the values reported by Brown and Pitman (1991) for bahiagrass (14.1 h). The rate of CP disappearance and lag time were not affected by the N fertilization level with average values of 0.06 h -1 and 4.2 h, respectively (Table 6-3). The CP lag time observed in this study was similar to the lag times reported by Assis et al. (1999) (3.5 6.0 h) for Tifton 85 and shorter than the average lag time found by Newman et al. (2002a) for limpograss (18 h). Assis et al. (1999) did not find significant effects of N fertilization on rate of degradation, corroborating the data presented in this study.

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121 Table 6-3. Effect of N fertilization rates on dry matter (DM) and crude protein (CP) degradation of Tifton 85 forage. N fertilization (kg ha -1 ) SE OPC P value 0 40 80 DM Lag time (h) 5.7 5.6 6.3 1.0 0.55 Rate of disappearance (h -1 ) 0.04 0.04 0.08 0.01 0.77 Potential degradability (g kg -1 ) 476 512 512 13 L CP Lag time (h) 4.4 4.0 4.2 2.2 0.94 Rate of disappearance (h -1 ) 0.02 0.05 0.11 0.07 0.27 Potential degradability (g kg -1 ) 567 597 611 17 L OPC = orthogonal polynomial contrast L=linear, (P<0.05) The potential degradability of the CP increased linearly with increasing N fertilization levels (Table 6-3). Assis et al. (1999) also found greater potential CP degradability of Tifton 85 fertilized with 400 kg N ha -1 (490 g kg -1 ) than non-fertilized (470 g kg -1 ). Newman et al. (2002a) observed a linear decrease in potential CP degradability when canopy heights of continuously-stocked limpograss pastures increased from 20 to 60 cm. Although there was no regrowth interval effect on DM disappearance lag time, rate of DM disappearance and potentially degradable DM were less at the 4-wk than the 2-wk regrowth interval (Table 6-4).

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122 Table 6-4. Effect of regrowth interval on dry matter (DM) and crude protein (CP) degradation of Tifton 85 forages. Regrowth interval SE P DM 2 wk 4 wk Lag time (h) 6.5 5.0 0.6 0.06 Rate of disappearance (h -1 ) 0.05 0.03 0.01 < 0.01 Potential degradability (g kg -1 ) 530 470 8 < 0.01 CP Lag time (h) 1.1 7.3 1.3 < 0.01 Rate of disappearance (h -1 ) 0.05 0.07 0.04 0.70 Potential degradability (g kg -1 ) 590 581 12 0.14 There was no effect of regrowth interval on rate of CP disappearance and potentially degradable CP (Table 6-4). However, lag time was shorter for Tifton 85 at a 2(1.1 h) than a 4-wk (7.3 h) regrowth. The lag time observed in this study for 2-wk-old Tifton 85 is comparable to the lag time in cool-season grasses (0 to 4.5 h) reported by Waghorn and Burke (2001), possibly because of the greater concentration of soluble protein in younger plants (Minson, 1990). There was an increase in CP disappearance from 0 to 72 h of incubation at all N fertilization levels and regrowth intervals (Fig. 6-2A). The CP disappearance of Tifton 85 herbage from the three N fertilization levels was similar during the first 3 h of incubation. A difference between the 0 N treatment and the others was observed after 6 h when the 0 N treatment started to show a greater proportion of undegradable CP. This continued

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123 through 72 h. A similar pattern of CP disappearance was observed for the two regrowth intervals (Fig. 6-2B). After 6 h of incubation, the forage harvested at a 4-wk regrowth was less degraded and that difference was maintained through 72 h. A0200400600800100001020304050607080HoursCP disappearance (g kg-1CP) 0 40 80 B0200400600800100001020304050607080HoursCP disappearance (g kg-1CP) 2 wks 4 wks Fig. 6-2. Tifton 85 crude protein (CP) disappearance at three N fertilization levels (A) and two regrowth intervals (B). According to Minson (1990), the CP in warm-season grasses is more slowly degraded in the rumen than that of cool-season grasses. The C4 plants have a lower soluble protein concentration, which is thought to be mainly due to lower Rubisco

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124 concentration than for C3s. Most of the Rubisco in C4 plants is located in the parenchyma bundle sheath, which is slowly or incompletely digested in the rumen (Akin and Burdick, 1975) and further contributes to the less rapid release of CP by C4 grass species. Summary and Conclusions Herbage accumulation of Tifton 85 bermudagrass increased linearly as the N fertilization level increased and was greater for 4than the 2-wk regrowth period in 1 of the 2 yr. Increasing N fertilization rates from 0 to 80 kg ha -1 resulted in a linear increase in CP and IVDOM concentrations, and herbage harvested at 4 wk contained less CP and IVDOM when compared with 2-wk regrowth. Crude protein Fraction A was not affected by N fertilization levels or maturity, however, there was a linear increase in Fraction B and linear decrease in Fraction C as N fertilization increased. Forage harvested at 4 wk had a greater proportion of Fraction C than forage harvested at 2 wk. In general, the proportion of the more degradable fractions (A and B) was greater for less mature and more heavily fertilized herbage. Level of N fertilization did not affect in situ lag time in DM disappearance, rate of disappearance, or potential disappearance. However, DM rate and potential disappearance were less with a longer regrowth period. Potential CP disappearance increased linearly as N fertilization level increased. The in situ lag time for CP was shorter for forage harvested at 2 than at 4 wk of regrowth. The in situ CP lag time for Tifton 85 harvested at 2 wk was comparable to lag times reported in the literature for cool-season grasses. In situ CP disappearance increased from 270 to 380 g kg -1 at 0 h to 720 to 790 g kg -1 at 72 h of incubation for the N fertilization levels and regrowth intervals tested in this study.

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125 In order to supply better nutritive value forage to ruminants, N fertilization and shorter regrowth intervals must be considered. In Tifton 85 bermudagrass longer regrowth intervals and/or no N fertilization decreased rumen degradable CP concentration in the forage. It is likely that if the grass was fed to ruminants there would be a deficiency of rumen-degradable protein. Diets that are low in rumen-degradable protein will have a greater negative impact on performance of ruminants with high CP requirements, e.g., early weaned calves and dairy cows. Use of N fertilization and a shorter regrowth interval are easily implemented management practices to improve nutritive value and potential rumen-degradable protein in Tifton 85 swards. Their use in production systems will depend upon the economics of fertilization and the potential yield reductions and lower stocking rates associated with shorter regrowth intervals. From a supplementation perspective, the issues are clearly different for Tifton 85 than for the rye (Secale cereale L.)-ryegrass (Lolium multiflorum Lam.) mixture. For rye-ryegrass, rapid release of N in the rumen suggests the need for high energy supplements with slowly ruminally degraded CP sources, while for Tifton 85, especially if more mature or less well fertilized, the supplement must assure that N levels in the rumen are sufficiently high to allow utilization of available energy.

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CHAPTER 7 SUMMARY AND CONCLUSIONS The profitability of the Florida cow-calf enterprise is largely dependent on the reproductive performance of the cow herd. Low body condition is responsible for lower conception rates and overall cow herd productivity. Early weaning of the calves of young cows and first-calf heifers is a management strategy that has potential to increase their reproductive rates by allowing more rapid restoration of body condition after parturition. Although the benefits of early weaning, i.e., improving reproduction and reducing nutrient requirements of the cow, have been recognized for many years, one factor limiting practical application of early weaning has been management of the early weaned (EW) calves. Producers that early wean their calves could possibly send these animals directly to the feedlot; however, this has not been the most profitable. The necessity of keeping the calves during the winter to get a higher price has resulted in recent studies evaluating feed management of EW calves. Mild winters in the southern USA offer an opportunity to raise calves on forage-based grazing systems using high nutritive value cool season annual forages in winter and warm-season perennial grasses during summer, however, some concentrate supplementation may be needed because the calves have a limited rumen capacity and high nutrients requirements. The major expense in raising these calves is feed cost, therefore low-cost alternatives to high concentrate diets, e.g. pasture-based systems, are needed. Two grazing studies (Chapters 3 and 4) were conducted to test the effects of supplementation level on herbage characteristics, forage intake, and performance of early 126

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127 weaned calves grazing rye (Secale cereale L.)-ryegrass (Lolium multiflorum Lam.) and Tifton 85 bermudagrass (Cynodon spp.) pastures during the winter-spring and summer seasons, respectively. In addition, two plot studies were conducted to assess the effects of N fertilization levels and regrowth interval on herbage accumulation, nutritive value, and in situ crude protein (CP) fractions of rye-ryegrass and Tifton 85 (Chapters 5 and 6). The overall goal of the research effort was to assess a range of strategies for raising early weaned calves and to explain the biological basis for the success or failure of these approaches. All studies were conducted at the University of Florida Beef Research Unit, 18 km northeast of Gainesville, FL. Rye-ryegrass Grazing Study Treatments were three levels of a commercial pelleted concentrate (147 g CP kg -1 and total digestible nutrient concentration of 700 g kg -1 ), 10, 15, and 20 g kg -1 of calf BW, offered daily. Each treatment was replicated three times in a completely randomized design, so there were a total of nine experimental units in the study. Data were collected during the 2003 and 2004 winter-spring seasons. Starting dates were 28 Jan. 2003 and 15 Jan. 2004, and grazing ended on 14 May 2003 and 29 Apr. 2004. Calves were weaned on 2 Jan. 2003 and 5 Jan. 2004 at approximately 90 d of age and 100 kg of body weight (BW). Pasture size was 0.2 ha, and each pasture was subdivided into four paddocks. Pastures were rotationally stocked with a 7-d grazing and 21-d resting period. Two early weaned calves (1 steer and 1 heifer) were assigned as testers to each experimental unit such that BW was equal across all experimental units. Put and take early weaned calves of comparable age and weight to the testers were used to maintain similar herbage allowance across experimental units.

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128 Rye-ryegrass pastures grazed by EW calves receiving different levels of supplement had similar herbage mass, accumulation, allowance, and nutritive value. Seasonal variation in forage responses were the result of the wheather conditions during the trial, especially the very high rainfall that occurred during the 2002/03 season. It resulted in lower herbage mass and CP during that year. Total organic matter intake (OMI) did not vary among the different supplementation levels, but forage OMI decreased linearly as supplement level increased. Average daily gain, SR, and LWG increased linearly with increasing levels of supplement, but grazing time decreased as greater amounts of supplement was fed. Greater ADG at the same level of intake suggests that there may have been better synchronization of energy and N in the rumen of the higher supplement treatment animals, and possibly an effect associated with the greater amounts of ruminal undegradable protein provided by the concentrate. Stocking rate increased because of the substitution of concentrate for forage in the 15 and 20 g kg -1 BW treatments. The blood urea nitrogen (BUN) concentrations of all treatments were in the range considered adequate for growing cattle. Economic analysis showed that there was no economic benefit of increasing the supplementation levels above 10 g kg -1 BW. Given the pasture conditions, and the concentrate and calf pricing in this study, the 10 g kg -1 BW supplement level is recommended for EW calves grazing cool-season grass pastures. Tifton 85 Bermudagrass Grazing Study Treatments were three levels, 10, 15, and 20 g kg -1 of calf BW offered daily, of the same commercial pelleted concentrate used in the winter study. Each treatment was replicated three times in a completely randomized design so there were a total of nine

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129 experimental units in the study. Dates for the trial during the 2 yr were 14 May through 13 Aug. 2003 (86 d) and 18 May through 10 Aug. 2004 (86 d). Calves had grazed rye-annual ryegrass pastures from January to April. In the summer study, they received the same supplement treatment as they did during the winter-spring period. At the start of the summer trial, they were approximately 200-d old with an average liveweight of 190 kg. Pasture size was 0.15 ha subdivided in three paddocks for rotational stocking. The grazing period was 7 d and the rest period was 14 d. Two early weaned calves (1 steer and 1 heifer) were assigned as testers to each pasture and put and take early weaned calves were used to maintain herbage allowance at approximately 1 kg forage DM kg -1 of calf BW. The different levels of supplement fed to early weaned calves grazing Tifton 85 pastures did not affect herbage mass, accumulation, or nutritive value of the pasture. Herbage allowance was slightly lower for the 20 g kg -1 BW supplementation, the result of the greater number of put and take calves assigned to that treatment. Nevertheless, there was no evidence that the herbage allowance in that treatment negatively affected calf performance. Average daily gain, SR, and LWG increased with greater levels of supplementation. Greater SR was associated with a tendency for calves consuming more concentrate to consume less forage; however, a problem with calves regurgitating the marker boluses limited the number of observations and made it impossible to draw firm conclusions about intake. Calves receiving lower concentrate rates had greater grazing time during daylight, supporting the observation of a trend toward lower forage intake with higher rates of supplement.

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130 Although greater supplementation levels resulted in greater average daily gain and gain per unit of land area, economic analysis of the data did not support use of the 20 g kg -1 BW treatment. Economic return per hectare was greater for the 15 g kg -1 BW concentrate treatment than the average of 10 and 20 g kg -1 treatments. Rye-Ryegrass Plot Study Treatments were the factorial combinations of two regrowth intervals (3 and 6 wk) and three N fertilization levels (0, 40, and 80 kg N ha -1 per season) evaluated in two seasons, winter (January-February) and spring (March-April). Plot size was 2 x 4 m with a 1-m alley between plots. Treatments were replicated three times in a randomized complete block design. The 6-wk cutting interval plots received the entire N application for that season (winter or spring) at the beginning of the 6-wk growth period, while the 3-wk plots received half the seasonal rate at the beginning of each of the two 3-wk growth periods per season. Data were collected during January through April 2003 and 2004. Herbage accumulation, botanical composition, and nutritive value were measured. In addition, an in situ technique was used to determine CP fractions and disappearance kinetics of DM and CP. Botanical composition of rye-ryegrass swards was affected by season of the year with the proportion of rye being greater during the winter and proportion of ryegrass greater during spring of the wetter year. Nitrogen fertilization and regrowth interval did not affect botanical composition. Herbage accumulation increased linearly with increased levels of N fertilization. More herbage accumulated with the 6-wk than the 3-wk regrowth interval in spring, but during winter, regrowth interval did not affect total accumulation. Accumulation was greater in spring than winter regardless of regrowth interval.

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131 The effect of regrowth interval on herbage CP and in vitro digestible organic matter (IVDOM) concentrations varied with season. There was no difference in IVDOM during the winter for the two regrowth intervals, however, lesser IVDOM concentrations were observed for the 6-wk than the 3-wk regrowth interval during the spring. Herbage CP concentrations were greater for the 3-wk than the 6-wk regrowth interval during both winter and spring, but the difference between intervals was much greater in spring. Thus, regrowth interval is of much greater importance in determining forage nutritive value and herbage accumulation during more rapid forage growth in spring than winter, and during spring, greater N fertilization levels and shorter regrowth periods than in winter may be necessary to maintain the nutritive value of the forage. There was a significant effect of N fertilization level on the CP fractions of rye-ryegrass. Greater N fertilization increased the concentration of Fraction A and decreased Fractions B and C. More Fraction A at greater fertilization levels has the potential to result in less efficient use of N by ruminants that can lead to greater N losses through the urine. No lag time was observed for in situ CP disappearance indicating that the CP present in rye-ryegrass herbage is highly soluble. Greater N fertilization increased the potential CP disappearance linearly. When considering utilization of rye-ryegrass herbage for early weaned calves, the greater total CP and Fraction A concentrations in rye-ryegrass with increased levels of N fertilization implies that there may be an excess of ruminal degradable protein under these circumstances. Utilization of high energy, low protein supplements may be a useful management practice to improve synchronization of N and energy availability in the

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132 rumen and increase the efficiency of rumen degradable protein utilization by EW calves grazing cool-season grass pastures. Tifton 85 Bermudagrass Plot Study Treatments were the factorial combinations of two regrowth intervals (2 and 4 wk) and three N fertilization levels (0, 40, and 80 kg ha -1 per 4-wk period) evaluated in two periods in each of 2 yr. Treatments were replicated three times in a completely randomized design. Plot size was 2 x 4 m with a 1-m alley between plots. Herbage was harvested to a 15-cm stubble. Herbage accumulation data were reported by period, and data from a given period are based on one harvest of the 4-wk treatment and two harvests of the 2-wk treatment. For laboratory analyses, samples used to determine herbage accumulation were analyzed for IVDOM and N concentration. Crude protein fractions and DM and CP disappearance kinetics were determined using an in situ technique. Herbage accumulation of Tifton 85 bermudagrass increased linearly as the N fertilization level increased and was greater for the 4than the 2-wk regrowth interval in 1 of the 2 yr. Increasing N fertilization from 0 to 80 kg ha -1 resulted in a linear increase in CP and IVDOM concentrations, and herbage harvested at 4 wk presented lesser CP and IVDOM when compared with 2-wk regrowth. Crude protein Fraction A was not affected by N fertilization level or maturity, however, there was a linear increase in Fraction B and linear decrease in Fraction C observed as N fertilization level increased. Forage harvested at 4 wk had a greater proportion of Fraction C than forage harvested at 2 wk. In general, the proportion of the more degradable fractions (A and B) was greater for less mature and more heavily fertilized herbage.

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133 Level of N fertilization did not affect in situ lag time in DM disappearance, rate of disappearance, or potential disappearance; however, DM rate and potential disappearance were less with a longer regrowth period. Potential CP disappearance increased linearly as N fertilization level increased. The in situ lag time for CP was shorter for forage harvested at 2 than at 4 wk of regrowth. The in situ CP lag time for Tifton 85 harvested at 2 wk was comparable to lag times reported in the literature for cool-season grasses. In situ CP disappearance increased from 270 to 380 g kg -1 at 0 h to 720 to 790 g kg -1 at 72 h incubation for the N fertilization levels and regrowth intervals tested in this study. As it matured, IVDOM and CP of the warm-season grass decreased. In order to supply better nutritive value forage to ruminants, greater N fertilization and shorter regrowth intervals may be considered. Longer regrowth intervals and/or low N fertilization decreases ruminal degradable CP concentration in the forage, and when fed to ruminants, is more likely to result in a deficient supply of rumen degradable protein. Diets that are low in rumen degradable protein will have a greater negative impact when fed to ruminants with high CP requirements, e.g., EW calves and dairy cows. Use of N fertilization and a shorter regrowth interval are easily implemented management practices to improve nutritive value and potential rumen degradable protein in Tifton 85 swards. Their use in production systems will depend upon the economics of fertilization and the potential yield reductions and lower stocking rates associated with shorter regrowth intervals. Implications of the Research The results of this series of studies are useful in guiding development of pasture-based feeding systems for EW calves. During winter the high nutritive value of cool-season grasses reduces the need for concentrate supplement. At supplement levels above

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134 10 g kg -1 BW, the calves grazing cool-season pasture reduced forage intake by 1.0 kg per kg of supplement fed. This high substitution rate diminished the potential for favorable economic impact and resulted in the 10 g kg -1 BW supplement level being the most practical treatment tested. The extremely high CP concentration and CP degradability of the cool-season grasses leads to the suggestion that the supplement fed need not be high in rumen degradable CP, but that it be relatively high in TDN and perhaps contain some rumen undegradable CP. Additional research with cool-season pasture systems is needed in several areas. First, it is recommended that lower levels of supplement be evaluated, perhaps 0, 5, and 10 g kg -1 BW. Also, supplement composition should be further evaluated, specifically different energy sources and different proportions of rumen degradable and undegradable CP. It was anticipated that higher levels of supplement would be more critical to performance of calves during summer because of the lower nutritive value of the warm-season grass. There was greater economic benefit associated with the 15 g kg -1 BW supplement treatment compared to the average of the 10 g kg -1 and 20 g kg -1 BW levels. Based on the results of the in situ study, N concentration of total CP and rumen degradable CP in the supplement are more important for calves grazing Tifton 85 than rye-ryegrass because of the proportion of CP that is immediately available and that which is potentially degradable are less for Tifton 85. Future research with warm-season pasture system with EW calves may focus supplement energy sources as well as sources of rumen degradable protein to increase efficiency of energy utilization. In addition, high nutritive value warm-season forages,

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135 e.g., pearl millet [Pennisetum glaucum (L.) R. Brown] and rhizoma peanut (Arachis glabrata Benth.), may be evaluated as alternatives to Tifton 85, especially for North Florida. In situation where high rates of concentrate are fed, the use of ionophores and implants may be tested to determine if they increase the efficiency of concentrate utilization. In summary, these studies have shown that relatively high levels of performance can be achieved by EW calves, on practical, pasture-based feeding systems. Supplementation programs need to be tailored to complement the forage component of the diet, and these studies have provided information that will be usefull in deciding the amount and composition of supplement to be fed to calves grazing cooland warm-season grasses.

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APPENDIX A DATA TABLES A Table A-1. Weather data for Years 2002, 2003, and 2004 in Gainesville FL. Month Rainfall Average Ambient Air Temperature 2002 2003 2004 30-yr Average 2002 2003 2004 30-yr Average ------------------mm-------------------------------------o C---------------------Jan. 99 46 30 83 12.7 8.2 11.0 13.4 Feb. 25 158 149 99 12.4 13.4 12.3 14.3 Mar. 50 243 51 93 17.4 18.7 16.5 17.7 Apr. 62 23 26 75 21.9 19.2 17.9 20.6 May 33 43 15 106 23.1 24.0 23.9 24.1 June 135 265 194 168 24.8 25.2 25.7 26.7 July 249 144 131 180 25.7 25.9 26.0 27.4 Aug. 165 123 281 203 25.0 25.6 25.8 27.4 Sep. 133 126 442 142 25.0 23.9 24.6 26.3 Oct. 63 84 81 59 22.1 20.7 21.4 22.0 Nov. 95 60 57 52 14.0 17.4 17.0 17.7 Dec. 128 26 43 81 10.8 10.1 14.8 14.5 Total 1237 1341 1487 1341 136

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137 Table A-2. Herbage mass double sampling equations and r2 for the rye-ryegrass pastures year Month Equation r 2 Jan. y=354 x 265 0.89 Feb. y=150 x + 278 0.89 Mar. y=206 x 180 0.94 2003 Apr. y=154 x + 287 0.81 Jan. y=243 x + 347 0.86 Feb. y=230 x + 322 0.87 Mar. y= 141 x + 364 0.86 2004 Apr. y=277 x + 538 0.86 When x=height (cm); y=kg DM ha -1

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138 Table A-3. Total chromium output of Captec controlled release device in early weaned calves submitted to total feces collection. Year 2003. Days Calf # 1 2 3 4 5 6 7 Avg -----------------------------------------------g d -1 ----------------------------------------------1 0.084 0.094 0.070 0.152 0.114 0.092 0.173 0.111 2 0.085 0.036 0.105 0.109 0.121 0.030 0.075 0.080 3 0.167 0.064 0.085 0.085 0.102 0.094 0.159 0.108 4 0.099 0.077 0.139 0.093 0.104 0.093 0.126 0.105 Avg 0.109 0.068 0.100 0.110 0.110 0.077 0.133 0.101 Table A-4. Total chromium output of Captec controlled release device in early weaned calves submitted to total feces collection. Year 2004. Days Calf # 1 2 3 4 5 6 7 Avg -----------------------------------------------g d -1 ----------------------------------------------1 0.156 0.119 0.128 0.112 0.132 0.115 0.141 0.129 2 0.105 0.096 0.096 0.105 0.150 0.163 0.138 0.122 3 0.089 0.108 0.134 0.179 0.126 0.106 0.135 0.125 4 0.103 0.101 0.122 0.137 0.122 0.159 0.137 0.126 Avg 0.113 0.106 0.120 0.133 0.133 0.136 0.138 0.125

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139 DATA EWC; OPTIONS PS = 59 LS =125; /* TRT = treatment PAST = pasture YR = year OMI = organic matter intake FOO = fecal output observed IVOMD SUPINTAK = supplement intake, OM basis digsup = digestibility of supplement, OM basis */ INPUT TRT PAST YR OMI FOO IVOMD SUPINTAK digsup; supdig = 0.72; FRGDIG = IVOMD; DO FRGINTAK =1 TO 40 BY .05 UNTIL (DIFF < .01); TOTINTAK = FRGINTAK + SUPINTAK; EXPDIG = (FRGINTAK*FRGDIG + SUPINTAK*SUPDIG)/TOTINTAK*100; ADJDIG = (59.71 -.8948*EXPDIG + .01399*EXPDIG**2)/100; FOP = TOTINTAK*(1-ADJDIG); DIFF = FOO-FOP; END; CARDS; ; proc print; run;

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APPENDIX B DATA TABLES B Table B-1. Herbage evaluation double sample equations and r 2 Month Pre/Post Equation r 2 pre 168x+2278 0.74 May* post 168x+2278 0.74 pre 191x+1753 0.65 June post 287x+878 0.78 pre 260x+1045 0.66 July post 234x+1693 0.66 pre 273x+1358 0.65 2003 August post 226x+1668 0.56 pre 179x+781 0.82 May post 214x+479 0.80 pre 220x+385 0.69 June post 995x+2327 0.51 pre 287x+220 0.79 July post 328x-106 0.78 pre 227x+824 0.81 2004 August post 261x+170 0.82 Pre and post grazing used the same calibration curve When x=height (cm); y=kg DM ha -1 140

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141 Table B-2. Total chromium output of Captec controlled release device in early weaned calves submitted to total feces collection. Year 2003. Days Calf # 1 2 3 4 5 6 7 Avg -----------------------------------------------g d -1 ----------------------------------------------1 0.108 0.143 0.149 0.153 0.165 0.1625 0.173 0.150 2 0.142 0.120 0.117 0.120 0.094 0.156 0.154 0.129 3 0.147 0.123 0.109 0.161 0.148 0.167 0.182 0.148 Avg 0.132 0.129 0.125 0.145 0.136 0.162 0.170 0.143

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BIOGRAPHICAL SKETCH Joao Mauricio Bueno Vendramini was born on 24 December 1972, in Araraquara, Sao Paulo State, Brazil. He is a son of Arnaldo and Lais P.B. Vendramini and has one brother, Jose Olavo. Joao is married to Maria Lucia Silveira, soil scientist, currently working as a chemist in the Soil and Water Science Department at the University of Florida. He graduated from Agriculture Technical High School UNESP in December 1989 with a Technician in Agriculture degree. In 1991, he received the Engenheiro-Agronomo B.S. degree from University of Sao Paulo ESALQ. He worked from 1994 to 2001 at AWL Administration Company administrating six properties specializing in beef cattle, sugarcane, and crops. During 1996-1999, he worked on a Master of Science program in animal science and forages concomitantly with his job, receiving a Master of Science degree from University of Sao Paulo ESALQ in March 1999. In January 2002, he was accepted into a Doctor of Philosophy degree program in agronomy, where he served as Graduate Research Assistant. He is currently a PhD candidate working in the area of forage management under the direction of Dr. Lynn E. Sollenberger. Joao is a member of the American Society of Agronomy, American Society of Animal Science, and American Registry of Professional Animal Scientists. 159


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SUPPLEMENTATION EFFECTS ON EARLY WEANED CALVES GRAZING
COOL- AND WARM-SEASON GRASSES















By

JOAO MAURICIO BUENO VENDRAMINI


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA


2005

































Copyright 2005

by

Joao Mauricio Bueno Vendramini


























To the memory of my grandfather, Jose Pereira Bueno





For being an inspiration for me and my brothers















ACKNOWLEDGMENTS

The author wishes to express his sincere gratitude to Dr. Lynn E. Sollenberger,

chairman of the advisory committee, for his guidance, support, and patience throughout

the graduate program. Appreciation and gratitude are extended to Dr. John D. Arthington,

co-chair of the advisory committee, for his support, enthusiasm, and friendship. Thanks

are extended to other members of the advisory committee, Dr. Carrol Chambliss

(deceased), Dr. Adegbola Adesogan, Dr. Todd Thrift, and Dr. Rob Kalmbacher, for their

contributions to the degree program in the form of suggestions and constructive criticism.

Also, the author thanks Dr. Jerry Bennett, department chairman, and Dr. David S.

Wofford, graduate coordinator, for the opportunity to study in the Agronomy

Department.

Special appreciation is due to Dr. John Moore, Dr. John Fike, and Dr. Bisoondat

Maccon for their contribution and help with intake statistical models.

The contribution of several individuals was valuable to the successful completion

of the research project. To fellow agronomy graduate students Jose Dubeux, Sindy

Interrante, and Lawton Stewart thanks are extended for their help and friendship during

the graduate program. Appreciation is extended to fellow graduate students from different

departments and institutions that collaborated in the research project, Nathan and

Wimberley Krueger, Marcia Grise, Ana Claudia Rugieri, Helder Santos, Flavio Silvestre,

Guto Castro, and Reinaldo Cooke. Special thanks to Agronomy Department staff Claude

Jones and Dwight Thomas, at the Forage Evaluation Field Laboratory, for their valuable









help in the field during the sample collection, animal weighing, and overall management

of the experimental area. Also, thanks go to Richard Fethiere for his help during the

sample analyses at the Forage Evaluation Support Laboratory.

Thanks are due to the families of Jose, Georgia and Victor Dubeux, Marcelo and

Aline Carvalho, Denis and Ann Vitorelli, Gabriel, Paolete, Julia, and Alice Couto, and

others that made us feel at home during graduate school in Gainesville.

Last but not least, the author is grateful to his wife, Maria Lucia, parents, Amaldo

and Lais, and brothers, Jose Olavo and Paulo Emilio (deceased), for their love, patience,

and support.
















TABLE OF CONTENTS

page

A C K N O W L E D G M E N T S ................................................................................................. iv

LIST OF TABLES ....................................................... ............ ....... ....... ix

LIST OF FIGURE S ......... ..................................... ........... xii

ABSTRACT ........ .............. ............. ...... ...................... xiv

CHAPTERS

1 IN TRODU CTION ................................................. ...... .................

2 LITER A TU R E REV IEW ............................................................. ....................... 4

Early Weaning ..............................................4
Forage System s in Southern U SA ........................................ .......................... 8
B erm udagrass .................................................................... 9
G general .................................... ...... ...... ... 9
Fertilization, Yield, and Nutritive Value............... ...........................................10
G razing Studies ............................................................. .... ... .......... 11
C ool-Season A annual G rasses............................................. ............................. 12
G general ......................................................... ...... ..... ......... 12
Fertilization, Yield, and Nutritive Value............... ...........................................13
G razin g Stu d ies .................................................. ................ 15
Supplementation .................................... ............................... .........18
P ro te in .....................................................1 9
Energy ................................................... ...... ......... ................. 21
Interactions between Supplements and Forages ..............................................23
B eh av ior ................................................................... 2 5
Intake ............................ ............. .. ..... ... ............. ......... 26
Measurements of Forage Intake of Grazing Animals........................................27
N Fractioning and in situ Methodology ............................................. ...............32
In situ Procedure .................................. ..................... ..................33
In situ Disappearance of Warm- and Cool-season Grasses...............................35
Other Methods to Estimate Ruminal Protein Degradation..............................37









3 SUPPLEMENTATION EFFECTS ON FORAGE CHARACTERISTICS AND
PERFORMANCE OF EARLY WEANED CALVES GRAZING RYE-
RYEGRA SS PA STU RE S ................................................ .............................. 39

In tro d u ctio n ................................................... .................. ................ 3 9
M material and M methods ........................................................... .. ............... 41
Pasture Sam pling .................. ............................... .......... ....... ...... .. 43
A nim al R response V ariables ........................................ .......................... 45
E conom ic A analysis ............ ...................................................... .... ........ ......47
Statistical A nalysis................................................... 48
Results and Discussion .................................... ...... .. ...... ............. 50
P asture R responses ........................... ........ ............................ .... ...... ...... 50
A nim al R response V ariables ........................................ .......................... 59
E conom ic A analysis ............ ...................................................... .... ........ ......65
Sum m ary and C onclu sions .............................................................. .....................65

4 SUPPLEMENTATION EFFECTS ON FORAGE CHARACTERISTICS AND
PERFORMANCE OF EARLY WEANED CALVES GRAZING TIFTON 85
BERM U D A GRA SS PA STU RE S ................................................... .....................67

In tro d u ctio n ................................................... .................. ................ 6 7
M material and M methods ........................................................... .. ............... 69
Pasture Sam pling .................. .......................... ....... ........ ............ 71
A nim al R response V ariables ........................................ .......................... 71
E conom ic A analysis ............ ............................ ...................... ............ .... ..72
Statistical A naly sis........... .................................................................. ....... .. .... 72
Results and Discussion .................................... ...... .. ...... ............. 73
P asture R espon ses ......................... ............................ .. ......... .... ............73
A nim al R response V ariables ........................................ .......................... 80
E conom ic A analysis ............ ............................ ...................... ............ .... .. 85
Sum m ary and C onclu sions .............................................................. .....................86

5 BOTANICAL COMPOSITION, HERBAGE ACCUMULATION, NUTRITIVE
VALUE, AND CRUDE PROTEIN FRACTIONATION OF RYE-RYEGRASS
HERBAGE AT DIFFERENT REGROWTH INTERVALS AND N
FER TIL IZ A TIO N LE V E L S............................................................ .....................87

Introdu action ...................................... ................................................. 87
M material and M methods ........................................................... .. ............... 90
Laboratory A nalyses .................. ......................... .. .. .... .. .. ........ .... 91
In situ Disappearance Procedure ...................................... ....................... 92
Dry Matter and Crude Protein Degradation Kinetics............ ................93
Statistical A n aly ses.......... ............................................................ .. .... .. .... .. 94
Results and D discussion ................... .......... ......... .... ............ ... ......... ......... 96
Botanical Composition, Herbage Accumulation, and Nutritive Value..............96
Protein Fractionation of the Forage........... ............. ...................... 99









Degradation Parameters of DM and CP ...................................................... 104
Sum m ary and C onclu sions ......... ................. ....................................................... 106

6 HERBAGE ACCUMULATION, NUTRITIVE VALUE, AND CRUDE
PROTEIN FRACTIONS OF TIFTON 85 SWARDS AT DIFFERENT CUTTING
INTERVALS AND N FERTILIZATION LEVELS ...............................................108

In tro d u ctio n ......................................................................................................... 1 0 8
M material and M ethods .................................. ............ ...... ........................ 110
Dry Matter and Crude Protein Degradation Kinetics ............ ............... 112
Statistical A n aly ses......................................................... ...................... ...... 1 12
Results and Discussion ............................ .. .................... .......... 14
Herbage Accumulation and Nutritive Value....................................................114
Protein Fractionation of the Forage................... ....... ........... .. .............. 117
Disappearance Parameters of DM and CP .............. .. ......... .. 119
Sum m ary and C onclu sions ........... ......... ...... ...................................................... 124

7 SUMMARY AND CONCLUSIONS................................................ 126

Rye-ryegrass Grazing Study ................................... ..............127
Tifton 85 Bermudagrass Grazing Study ........................................ ...............128
R ye-R yegrass Plot Study ................................ ......... ..................... ............... 130
Tifton 85 Bermudagrass Plot Study ...................................... ............... 132
Im plications of the R research .................................. .............................. ....... 133

APPENDIX

A D A T A T A B L E S A ............................................................................... ...... ..... 136

B D A T A T A B L E S B ......................................................................... ....................140

L IST O F R E FE R E N C E S ........................................................................ ................... 142

BIOGRAPHICAL SKETCH ............................................................. ............... 159















LIST OF TABLES


Table p

3-1 Ingredient and chemical composition of supplement fed to early weaned calves
on pasture (D M basis). ....................................... ............. .......... 44

3-2 Pregraze herbage mass and herbage accumulation of rye-ryegrass pastures
grazed by early weaned calves supplemented with three levels of concentrate.
(B W = b ody w eight)........... .......................................................... .. .... .... .. ..5 1

3-3 Herbage crude protein (CP) and in vitro digestible organic matter (IVDOM)
concentrations of rye-ryegrass pastures grazed by early weaned calves
supplemented with three levels of concentrate. (BW=body weight).....................52

3-4 Herbage allowance and SR of rye-ryegrass pastures grazed by early weaned
calves supplemented with three levels of concentrate. (BW=body weight) ............56

3-5 Average daily gain (ADG) and liveweight gain (LWG) of early weaned calves
grazing rye-ryegrass pastures and supplemented with three levels of concentrate.
(B W = b ody w eight)........... .......................................................... .. .... .... .. ..6 1

3-6 Total organic matter intake (OMI) and forage OMI of rye-ryegrass pastures
grazed by early weaned calves supplemented with three levels of concentrate.
(B W = b ody w eight)........... .......................................................... .. .... .... .. ..62

3-7 Grazing time during daylight hours of early weaned calves grazing rye-ryegrass
pastures and supplemented with different levels of concentrate............................63

3-8 Blood urea nitrogen (BUN) concentrations of early weaned calves grazing rye-
ryegrass pastures supplemented with three levels of concentrate. (BW=body
w eight) ............................................................... .... ...... ......... 64

3-9 Economic analysis of three concentrate levels on early weaned calves grazing
rye-ryegrass pastures. ..................................................................... ................ .. 65

4-1 Pregraze herbage mass and herbage accumulation of Tifton 85 pastures grazed
by early weaned calves supplemented with three levels of concentrate.
(B W =body w eight)........... .......................................................... .. .... ..... .. 73









4-2 Crude protein (CP) and in vitro digestible organic matter (IVDOM)
concentration of Tifton 85 pastures grazed by early weaned calves supplemented
with three levels of concentrate. (BW=body weight) ...........................................76

4-3 Herbage allowance and stocking rate of Tifton 85 pastures grazed by early
weaned calves supplemented with three levels of concentrate. (BW=body
w eight) ............................................................... .... ...... ......... 79

4-4 Average daily gain (ADG) and liveweight gain (LWG) ha-1 of early weaned
calves grazing Tifton 85 pastures supplemented with different levels of
concentrate. (BW =body w eight) ........................................ ......................... 81

4-5 Total organic matter intake (OMI) and forage OMI of Tifton 85 pastures grazed
by early weaned calves supplemented with three levels of concentrate ................82

4-6 Grazing time during daylight hours of early weaned calves grazing Tifton 85
pastures supplemented with three levels of concentrate. (BW=body weight) .........84

4-7 Blood urea nitrogen (BUN) concentrations of early weaned calves grazing
Tifton 85 pastures supplemented with three levels of concentrate. (BW=body
w eight) ................................................................ .... ...... ......... 85

4-8 Economic analysis of three concentrate levels for early weaned calves grazing
T ifto n 8 5 p a stu re s .................................................................................. 8 5

5-1 Year*season interaction forage effects on botanical composition, crude protein
(CP), and in vitro digestible organic matter (IVDOM) of rye-ryegrass mixtures.. 100

5-2 Season*regrowth interval interaction effects for herbage accumulation and crude
protein (CP) and in vitro digestible organic matter (IVDOM) concentrations of
rye-ryegrass forage mixtures during 2003 and 2004 ............................................101

5-3 N*season interaction effects on crude protein (CP) and in vitro digestible
organic matter (IVDOM) concentrations of rye-ryegrass forage.......................... 102

5-4 Effect of N fertilization levels on DM and CP degradation parameters of rye-
ryegrass forage. .................................................................... 105

6-1 N fertilization effects on herbage accumulation and in vitro digestible organic
matter (IVDOM) of Tifton 85 forage during 42-d periods. ..................................116

6-2 Regrowth interval*N fertilization level interaction effects on CP concentrations
of Tifton 85 forage. ............................................... .... .... .. ........ .. .. 117

6-3 Effect of N fertilization rates on dry matter (DM) and crude protein (CP)
degradation of Tifton 85 forage. ............. .... ............................................ 121









6-4 Effect of regrowth interval on dry matter (DM) and crude protein (CP)
degradation of Tifton 85 forages. ................................. 122

A-1 Weather data for Years 2002, 2003, and 2004 in Gainesville FL..........................136

A-2 Herbage mass double sampling equations and r2 for the rye-ryegrass pastures.... 137

A-3 Total chromium output of Captec controlled release device in early weaned
calves submitted to total feces collection. Year 2003 ............ ........ ...........138

A-4 Total chromium output of Captec controlled release device in early weaned
calves submitted to total feces collection. Year 2004 .........................................138

B-l Herbage evaluation double sample equations and r2...........................................140

B-2 Total chromium output of Captec controlled release device in early weaned
calves submitted to total feces collection. Year 2003 ...........................................141
















LIST OF FIGURES


Figure page

3-1 Year*period interaction effect on pregraze herbage mass of rye-ryegrass
pastures grazed by early weaned calves supplemented with different levels of
concentrate...................................................................... ........... ....... 53

3-2 Rainfall distribution from October to May 2002/03 and 2003/04. ........................53

3-3 Year*period interaction effect on average herbage accumulation of rye-ryegrass
pastures grazed by early weaned calves supplemented with different levels of
concentrate...................................................................... ........... ....... 54

3-4 Year*period interaction effect on crude protein (CP) concentration of rye-
ryegrass pastures grazed by early weaned calves supplemented with different
levels of concentrate ............................................... ........ ................. 57

3-5 Year*period interaction effect on in vitro digestible organic matter (IVDOM)
concentration of rye-ryegrass pastures grazed by early weaned calves
supplemented with different levels of concentrate............................................57

3-6 Year*period interaction effect on herbage allowance of rye-ryegrass pastures
grazed by early weaned calves supplemented with different levels of
concentrate.. ........................... ........... ............................................58

3-7 Year*period interaction effect on average daily gain of early weaned calves
grazing rye-ryegrass pastures and supplemented with different levels of
concentrate.. ........................... ........... ............................................61

4-1 Year*period interaction effect on pregraze herbage mass of Tifton 85 pastures
grazed by early weaned calves supplemented with different levels of
concentrate. .......................................... ............................ 74

4-2 Year*period interaction effect on average herbage accumulation of Tifton 85
pastures grazed by early weaned calves supplemented with different levels of
concentrate...................................................................... ........... ....... 75

4-3 Year*period interaction effect on crude protein (CP) concentration of Tifton 85
pastures grazed by early weaned calves supplemented with different levels of
concentrate. .......................................... ............................ 77









4-4 Period effect on IVDOM concentration of Tifton 85 herbage grazed by early
weaned calves supplemented with different levels of concentrate...........................78

4-5 Year*period interaction effect on average daily gain of early weaned calves
grazing Tifton 85 pastures and supplemented with different levels of
concentrate. .......................................... ............................ 82

5-1 Nitrogen fertilization effects on crude protein (CP) fraction concentrations in
total CP of rye-ryegrass forage..................................... ............................ ........ 103

5-2 N fertilization rate on crude protein (CP) disappearance of rye-ryegrass forage... 106

6-1 N fertilization (Fig. A) and regrowth interval (Fig. B) effects on crude protein
(CP) fraction concentrations in total CP of Tifton 85 forage..............................1.19

6-2 Tifton 85 crude protein (CP) disappearance at three N fertilization levels (A) and
tw o regrow th intervals (B). ............................................ ............................ 123















Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

SUPPLEMENTATION EFFECTS ON EARLY WEANED CALVES GRAZING
COOL- AND WARM-SEASON GRASSES

By

Joao Mauricio Bueno Vendramini

August, 2005

Chair: Lynn E. Sollenberger
Cochair: John D. Arthington
Major Department: Agronomy

Early weaning of their calves increases the likelihood that first calf heifers will

rebreed, but there is little information regarding pasture-based feeding programs for the

weaned calf. Research was conducted in Gainesville, FL, to evaluate the effect of

supplement rates on performance of early weaned (80 d of age) beef calves grazing

annual ryegrass (Lolium multiflorum)-rye (Secale cereale) mixtures from January to April

(winter) and Tifton 85 bermudagrass (Cynodon sp.) from May to August (summer) 2003

and 2004. Three levels of supplement (10, 15, and 20 g kg-1 of calf liveweight) were

evaluated in a completely randomized design with three replications. The supplement

contained 147 and 700 g kg-1 of crude protein (CP) and total digestible nutrients,

respectively. Pastures were rotationally stocked using a variable stocking rate in both

seasons with 21- and 14-d regrowth periods for rye-ryegrass and Tifton 85 bermudagrass,

respectively. Average daily gain increased linearly during the winter (0.74-0.89 kg) and

summer (0.53-0.72 kg) as supplementation rate increased. There was a linear increase in









live weight gain per hectare with increasing supplement rate during the winter (950-1320

kg) and summer (1100-1780 kg). In addition, two plot studies were conducted with the

same species used in the grazing trials to evaluate the effect of N fertilization levels and

forage regrowth interval on in situ CP fractionation of the forage. Treatments were the

factorial combinations of three N rates (0, 40, and 80 kg ha-1) and two regrowth intervals,

3- and 6-wk (rye-ryegrass) and 2- and 4-wk (Tifton 85). Increasing N fertilization levels

increased rye-ryegrass CP Fraction A (readily degradable) (410-590 g kg-1) and

decreased linearly CP Fraction B (slowly degradable) (510-360 g kg-1) and Fraction C

(undegradable) (74-48 g kg-1). On Tifton 85, increasing N fertilization increased linearly

Fraction B (330-455 g kg-1) and decreased linearly Fraction C (283-208 g kg-1). In

addition, greater proportions of fraction C were observed in forage of 4-wk (280 g kg-1)

than 2-wk maturity (200 g kg-1). Results of these studies indicate that pasture-based

feeding systems with modest levels of supplementation (10-15 g kg-1 of liveweight) are

practical options for raising early weaned calves in Florida.














CHAPTER 1
INTRODUCTION

Grasslands cover 4.5 million ha in Florida, most of which are utilized by beef

cattle (Bos sp.; Chambliss, 1999). The beef industry is a very important component of

Florida's agriculture industry. In 2003, Florida had a total of 950 000 beef cows that

accounted for $348 million of income to Florida farmers and ranchers (Florida

Agriculture Statistics, 2004).

The profitability of the Florida cow-calf enterprise is largely dependent on cow

productivity. The low nutritive value of Florida's tropical forages in combination with

reduced pasture production during winter reduces cow reproductive performance. Low

body condition is responsible for lower conception rates and overall cow herd

productivity (Arthington, 2002). These effects are most pronounced in young cows and

heifers which have higher nutritional demands to support both lactation and growth

(Arthington, 2002). Early weaning calves from young cows and first-calf heifers is a

management strategy that has potential to address this problem. At early weaning, cows

stop lactating and their requirement for total digestible nutrients (TDN) decreases by as

much as 49%. As a result they are able to restore body condition rapidly (Arthington,

2002). Bishop and Wettemann (1990), cited by Lusby (1995), reported that 100% of

cows with body condition scores > 5 had begun cycling within 25 d after their calves

were weaned at 45 d of age compared to only 43% of cows with condition score < 5.

Although the benefits of early weaning, e.g., improving reproduction and reducing

nutrient requirements of the cow, have been recognized for many years, producer









adoption of early weaning has been limited by lack of management information about the

weaned calves (Lusby, 1995). The major expense in raising these calves is feed cost

(Lusby and Donald, 1994). Therefore, low-cost alternatives to high concentrate diets are

needed. Forage intake of early weaned calves has not been well documented and varies

greatly between the time of early and normal weaning due to increasing development of

rumen function. The quantity of concentrate supplementation required during this period

for calves on pasture is likewise not well defined. Muehlmann et al. (1997) studied

different species of tropical pastures as the sole source of feed to early weaned calves.

Daily gains were low, averaging 0.2 kg d-1. The National Research Council NRC (1996)

reported that calves of 100-kg live-weight must consume 2.6 kg of dry matter d-1 with

250 g kg-1 of protein and 820 g kg-1 of TDN to achieve 1.0 kg of daily weight gain.

Considering that most forages and all of the C4 grasses are unable to supply these CP and

TDN concentrations, providing the appropriate amount of concentrate supplement to

early weaned calves is critical to achieving high performance. Research to quantify the

effect of various levels of concentrate on forage intake and calf performance is needed to

improve the management of early weaned calves.

The research described in this dissertation consists of four experiments. The first

two experiments were grazing trials with early weaned calves fed three levels of

concentrate supplement while grazing rye (Secale cereale L.)-annual ryegrass (Lolium

multiflorum Lam.) pastures during winter (Chapter 3) and Tifton 85 bermudagrass

(Cynodon spp.) pastures during summer (Chapter 4). The objectives of these studies were

to measure animal responses including average daily gain, stocking rate, live weight gain

per unit of pasture area, and forage intake. Pastures were characterized in terms of









herbage mass, accumulation, allowance, and nutritive value. Economic analyses for the

different rates of concentrate were performed. These data will allow producers to better

assess the profitability of early weaning of beef calves in North-Central Florida.

The last two experiments were plot studies with the same forage species used in the

grazing trials, rye-ryegrass (Chapter 5) during the winter-spring and Tifton 85

bermudagrass (Chapter 6) during the spring-summer. The primary objective was to

characterize the forage N fractions using an in situ technique. This is an important tool

for characterizing the rate and extent of N release in the rumen, potentially resulting in

better supplementation programs that increase animal performance and decrease loss of N

to the environment. Information regarding the N profile of forages is also useful to guide

future research and development of animal nutrition models. Additional forage responses,

including botanical composition and herbage accumulation, were measured in these

studies.














CHAPTER 2
LITERATURE REVIEW

Early Weaning

Early weaning is a management practice used by dairy and beef cattle (Bos sp.)

producers. The definition of early weaning varies, but typically calves weaned < 150 d of

age are considered early weaned. Several authors have reported that increasing the

fertility rates of beef cows is a benefit of early weaning. Lusby and Wetteman (1980)

compared reproductive performance in first-calf heifers following early and normal

weaning of their calves. Conception rate was greater among heifers whose calves were

early weaned (EW) (97%) than among those whose calves were normal weaned (NW)

(59%) during the 64-d breeding season. The average interval from parturition to

conception was reduced from 91 to 73 d by early weaning. According to Arthington and

Kalmbacher (2003), early weaning resulted in heavier first-calf heifers with greater body

condition score at the time of normal weaning. Heifers with EW calves had higher

pregnancy rates than those with NW calves (93 vs. 65%) during a 2-yr study in South

Florida.

Despite improved pregnancy rates, the practice of early weaning calves is still a

challenge for beef cattle producers, in part because of few management options for the

weaned calves. Producers that practice EW could possibly send these animals directly to

the feedlots; however, this procedure has not been the most profitable. The necessity of

keeping the calves during the winter to target the historically greater values of the spring

calf market has resulted in recent studies evaluating feed management of EW calves.









The ability of the young ruminant to sustain itself nutritionally is highly

dependent on the development of a functional rumen. During the first 3 wk of life, the

calves are limited to milk feeding. This limitation directly related to the presence or

absence and activity of digestive enzymes during early development of the

gastrointestinal system. From 3 to 8 wk of age, dry feed consumption may increase to the

level at which it contributes significantly to the energy and protein requirements to the

animal (Davis and Drackley, 1998). There is a large variability among calves for dry-feed

intake after 8 wk of age. This variability probably is related to the rate at which the calf

develops physiologically and its health. However, the calves are able to the majority of

their energy requirement from solid feed after 8 wk (Davis and Clark, 1981).Based on

their physiological development and in order to maintain a 365-d calving interval, calves

should be EW at less than 80 d of age. About 40 d of age may be a practical minimum for

early weaning in beef herds. Calves of at least 40 d of age do not require milk replacers in

the ration and are old enough to eat dry feed.

Mild winters in the southern USA offer an opportunity to raise calves on pasture-

based systems using high nutritive value cool-season annual forages. According to

Arthington (2002), EW calves grazing high quality pastures can grow as fast as NW

calves. The practice of raising EW calves on winter pastures is a feasible alternative,

especially if pasture quality and quantity are superior to hay available for winter feeding

(Barnes et al., 1996). Coffey et al. (2000) stated that calves fed hay plus supplement

gained less weight than calves that grazed winter-annual forages. Weder (1997) reported

an average daily gain of 0.8 kg d-1 for EW calves grazing annual ryegrass. According to

Rouquette et al. (1997), annual ryegrass resulted in individual live weight gains of 1.25









kg d-1 for suckling calves and 1.05 kg d-1 for stocker calves. Early weaned calves grazing

wheat (Triticum aestivum L.) pastures consumed 27% less forage dry matter (DM) during

the first 20 d than during the subsequent 50 d (Paisley et al., 1998). The authors

suggested that initial performance of EW calves grazing winter wheat is limited by low

forage intake. Average daily gain for EW calves grazing wheat pasture was 0.86 kg d-1

during a study with fall-born calves in Oklahoma (Purvis et al., 1996).

Few studies have been conducted with EW calves grazing summer pastures.

Muehlmann et al. (1997) studied tropical pastures as the sole source of feed to EW

calves. Calf performance was low with daily gains averaging 0.2 kg d-1. Smeaton et al.

(1996) reported that concentrate supplementation gave a substantial improvement in live

weight gain (+19 kg) in EW calves grazing on low nutritive value summer pastures. In

south Florida, supplementing EW calves with concentrate at 10 g kg-1 of body weight

resulted in gains of 0.59 and 0.44 kg d-1 on stargrass (Cynodon nlemfuensis Vanderyst)

and atrapaspalum (Paspalum atratum Swallen), respectively (Vendramini et al., 2003). In

a study with different forage species, Harvey and Burns (1989) evaluated gain of EW

calves grazing hill-land pastures (control), 'Tillman' white clover (Trifolium repens L.),

Tillman white clover and tall fescue (Festuca arundinacea Schreb.) mixtures, and pearl

millet [Pennisetum glaucum (L.) R. Brown]. Calves on each forage treatment were

provided ground ear maize (Zea mays L.) ad libitum. Calf daily gains were 0.72, 0.82,

0.57, and 0.92 kg d-1, respectively. Calves grazing millet had the greatest response in gain

and conversion of concentrate to liveweight gain.

According to some authors, EW calves have better feed efficiency than NW calves.

Myers et al. (1999b) conducted a study comparing EW to NW with or without creep









feeding for beef steers. Early weaned steers had greater average daily gains (ADG)

during the entire study than the average of creep-fed or normal-fed steers. Early weaned

steers had a lesser intake (7.29 vs. 7.68 kg d-1) and better feed conversion (0.16 vs. 0.14

kg gain kg-1 feed) than the average of NW calves with and without creep feeding. Lusby

(1995) reported fall EW calves consumed 20% less hay and were 43% more efficient in

converting total digestible nutrients (TDN) to weight gain than NW calves. In a study to

determine the effect of age at feedlot entry, Schoonmaker et al. (2001) reported that EW

calves (110 d of age) were more efficient than NW (202 d of age) and yearling steers

(371 d of age) (0.23, 0.21, and 0.18 kg gain kg-1 feed, respectively). Ringwall (2002)

evaluated EW calves with an average weight at weaning of 70 kg. They gained 0.9 kg d-1

and the feed efficiency was 5.3 kg of feed kg-1 of gain. When weaned at 70 d, EW calves

were 14 kg heavier, had a greater average daily gain, and were 11-d younger at slaughter

than NW calves (Grimes and Turner, 1991).

Considering meat quality traits at slaughter, EW calves had a higher quality grade

than NW calves (Fluharty et al., 2000; Myers et al., 1999b). In a similar study Myers et

al. (1999a) did not find differences for carcass weight, longissimus muscle area, or yield

grade among calves weaned at 90, 152, and 215 d of age. Moreover, there was no

difference in marbling score or percentage of steers grading greater than or equal to

Choice or Average Choice. Fluharty et al. (2000) reported that EW calves fed either 100

or 90% concentrate diets were heavier than NW calves at 210 d. The NW calves had

greater backfat thickness at 210 d but no difference in longissimus muscle area when

compared to EW calves fed a 60% concentrate diet.









Forage Systems in Southern USA

Hoveland (1992) estimated that out of- 39 million ha of perennial pastures grown

in the eastern USA almost 30 million ha are located in the South. These pastures are

mainly used for beef cows and calves and stockering of weaned calves.

Seasonal variation in amount and nutritive value of forage are the main challenges

over the course of the growing season. The predominantly warm-season grasses in

Florida are bahiagrass (Paspalum notatum Fliigge) and bermudagrass [Cynodon dactylon

(L.) Pers.]. These perennial, warm-season forages adapted to the Southeast are typically

of lower nutritive value than either cool-season perennials or warm-season annuals

(NRC, 1989).

Warm-season grasses are more efficient in light conversion than cool-season

species, but the major adaptation advantage is their better water-use efficiency, resulting

in higher total forage production when compared with cool-season grasses in tropical and

subtropical climates (Brown and Simmons, 1979). While the division of physiological

activity between mesophyll and bundle sheath cells in warm-season grasses is beneficial

for CO2 fixation and adaptation, the anatomical organization required for warm-season

grasses causes forage quality to generally be lower than for cool-season grasses (Akin,

1986). Warm-season grasses have relatively larger proportions of vascular tissues, bundle

sheath, and sclerenchyma than cool-season grasses. Some of these tissues in warm-season

grasses are potentially digestible, but because of their chemical composition and plant

anatomical structure the rate of degradation and fermentation is relatively slow (Coleman

et al., 2004). Although warm-season grasses grow well in the summer, their production is

seasonal, being interrupted by cold or dry weather.









Cool-season grass species generally are considered to have high nutritive value

than warm-season grasses mainly because of the amount and arrangement of tissues

(Akin, 1986). Mild winters in Florida give producers the opportunity to cultivate cool-

season annual forages during a period when productivity of warm-season grasses is low.

The importance of the cool-season grasses in the southern USA can be appreciated by

considering the seed market. The annual sale of seeds of winter annuals is enough to

plant 6 million ha of winter pastures (Moser and Hoveland, 1996).

Bermudagrass

General

Cynodon spp. are geographically widely distributed. They occur in greatest

abundance in tropical and subtemperate environments (Taliaferro et al., 2004).

Bermudagrass is one of the most important species in the southeastern USA with 10 to

12 million ha of planted areas for livestock grazing and hay. Many Cynodons have been

used as herbage for livestock, but several are of minor value because of narrow

distribution or characteristics limiting their adaptation. In 1937, USDA-ARS geneticist

Glenn W. Burton initiated a bermudagrass breeding program at the Coastal Plain

Experiment Station in Tifton, GA, with the objective of developing cultivars that were

more productive than the common strains and were capable of supplying highly nutritious

and palatable forage during a greater portion of the year (Burton, 1947). In 1943, the

release of 'Coastal' bermudagrass was a landmark in bermudagrass breeding, because it

was widely accepted by livestock producers in the southeastern USA. In 1993, 'Tifton

85', a hybrid between a South African bermudagrass and 'Tifton 68' stargrass, was

released. Burton et al. (1993) described it as taller, with larger culms, broader leaves, and









darker color than other bermudagrass hybrids. Compared with Coastal, Tifton 85 yielded

26% more DM, was 11% more digestible, and was more succulent at harvest.

Fertilization, Yield, and Nutritive Value

Bermudagrasses require relatively high soil nutrient availability to maintain good

production performance. The major determinants of fertilizer response are climate, native

soil nutrient status, source and rate of applied nutrients, season of applicationss, cultivar,

and defoliation regimen and method.

Nitrogen has the greatest influence on biomass yield and accordingly influences the

amount of other nutrients required to sustain production at specific N levels (Taliaferro et

al., 2004). Bermudagrass hay yield and quality are maximized when N is applied in split

applications at annual levels higher than 400 kg ha1 yr1 (Overman et al., 1992).

According to Wilkinson and Langdale (1974), cited by Taliaferro et al. (2004), Coastal

and other select bermudagrass cultivars produced maximum yields of about 27 Mg ha-1 at

N rates of 1200 kg ha-1, with near linear yield response to N up to about 600 to 700 kg N

ha-1.

Prine and Burton (1956) evaluated the effects of different N rates and harvest

frequencies on Coastal bermudagrass biomass production and nutritive value. There was

a curvilinear increase in herbage accumulation as N rate increased from 0 to 1008 kg N

ha-1 with maximum DM accumulation of 18.4 Mg ha-1 with a 6- to 8-wk harvest interval.

Bermudagrass showed the highest efficiency in production (kg DM kg- N ha-1) at 336 kg

N ha-1. The forage CP concentration ranged from 97 to 190 g kg-1 with N fertilization

rates from 0 to 1008 kg ha-1. Typically, increased CP concentrations have been reported

with increasing N application. Stallcup et al. (1986) reported that CP concentrations in

bermudagrass fertilized with 0 and 50 kg N ha-1 were 114 to 143 g kg-1. Burton et al.









(1963) studied different defoliation intervals of Coastal fertilized with 660 kg N ha-1. The

maximum herbage accumulation was 22 Mg ha-1 using a 6-wk interval. According to the

same author, annual DM production of 20 Mg ha-1 removed 400 to 450 kg N, 30 to 50 kg

P, 260 to 320 kg K, 30 kg S, 30 kg Mg, and 60 kg Ca ha-lfrom the soil.

Johnson et al. (2001b) compared different species of warm-season grasses in

Florida. Bermudagrass produced more forage DM (1,540 43 kg ha-lcutting-1) than

stargrass (1,400 43 kg ha-cutting-1) or bahiagrass (1,300 43 kg halcutting-1) when

harvested every 28 d. Peak herbage accumulation for all species occurred in late June and

July. A quadratic response to N fertilization was noted for in vitro digestible organic

matter digestibility (IVDOM) of bermudagrass, whereas bahiagrass was not affected.

Bermudagrass was more digestible (575 4 g kg-1) than stargrass (546 4 g kg-1) and

bahiagrass (519 4 g kg-1). As fertilization level increased, neutral detergent fiber (NDF)

decreased linearly in all three forages. Total N concentration and the concentration of all

protein fractions for all forages increased as N fertilization increased. In summary,

digestibility, CP concentration, and yield of bermudagrass increased with increasing N

application when plants were harvested every 28 d.

Grazing Studies

Bermudagrass is the most popular warm-season perennial grass for hay production

and pastures in the southern USA (Taliaferro et al., 2004). In North-Central Florida,

Pedreira et al. (1998) compared Florakirk and Tifton 85 bermudagrass in a 3-yr grazing

study. The pastures were continuously stocked and fertilized with 210 kg N ha-1 in four

applications yr-1. Although the average daily gain was similar between species (0.6 kg d-

1), Tifton 85 pastures supported higher average stocking rates (6.0 vs. 4.0 heifers ha-1),

resulting in greater gain (648 vs. 371 kg ha-1). Examination of 3-yr, total-season (169 d)









gains of steers grazing high quality bermudagrass pastures revealed that steer ADGs were

0.90 and 0.88 kg, respectively, from April to July, but only 0.30 and 0.43 kg,

respectively, for Tifton 78 and Tifton 85 pastures from July to October (Hill et al., 1993).

The authors suggested that decreased ADG was the consequence of increased

maintenance requirement of heavier steers and lower nutritive value later in the season.

Rouquette et al., (2003) reported that stocker steers gained 1.03 kg d-1 on Tifton 85

compared to 0.75 kg d-1 on Coastal without supplementation. With supplementation of

1.4 kg d-1 of a 3:1 maize:soybean (Glycine max L.) meal concentrate, steer ADGs were

1.25 kg d1.

Camevalli et al. (1999) maintained Tifton 85 swards heights of 5, 10, 15, or 20 cm

with grazing Santa Inez crossbred lambs (Ovis aries L.) paired by sex and body weight.

Nutritive value of forage was different for sward heights; short swards (5 cm) had the

highest CP and in vitro DM digestibility (IVDMD), but lamb performance was more

dependent on carrying capacity and forage allowance, with lambs performing better on

taller swards (15 to 20 cm).

Cool-Season Annual Grasses

General

Although warm-season grasses dominate most of the pastures in the southeastern

USA, climatic conditions during winter allow for use of cool-season annual grasses,

either sod-seeded or in prepared seedbeds. Reasons for use of cool-season annual grasses

include the extending the grazing period, high nutritive value, compatibility and ease of

establishment in warm-season perennial grass pastures, and tolerance to different

defoliation regimens and stocking rates (Rouquette et al., 1997). Small grains are often a

component of cool-season grass mixtures because they improve early-season forage









production. Of the options available, annual ryegrass-small grain mixtures provide the

earliest and greatest amount of forage if planted early on a well-prepared seedbed;

however, they also are expensive and are best used by lactating dairy cows, stocker

calves, replacement heifers, or limit grazed by beef cows nursing fall calves (Evers et al.,

1997).

Annual ryegrass, a high-yielding, high quality grass, is the most commonly grown

cool-season pasture forage in the southern and southeastern USA from November to May

(Evers et al., 1997). Common annual ryegrass was successfully grown in the Gulf Coast

region of the USA in the 1940s and 1950s. However, crown rust (Puccinia coronata f. sp.

avenae) was a serious problem and reduced forage yields and quality. Plant breeding

efforts in Texas, Mississippi, and Florida resulted in the release of 'Florida Rust

Resistant' annual ryegrass (Chapman and Webb, 1965). Hectarage of annual ryegrass

planted area has increased due to improved crown rust resistance as well as increased

forage and seed yield, and significant gains in winter hardiness (Griffith and Chastain,

1997). Among several cultivars released for southern areas, the tretaploid 'Jumbo' (Reg.

no. CV-220, PI 614099) is noted for its high seed yield, high resistance to crown rust and

gray leaf spot (Pyricularia grisea [Magnaporthe grisea]) and moderate resistance to

Helminthosporium leaf spot disease (Drechslera). Jumbo is late maturing, has larger

stems, leaves, seed heads, and seeds than the diploid 'Surrey', with CP concentration and

IVDOM comparing favorably with Surrey and other ryegrass cultivars (Prine et al.,

2002).

Fertilization, Yield, and Nutritive Value

Robinson et al. (1987) reported annual ryegrass yields in the southern states have

generally increased with N application rates up to 450 kg ha-1. Hovermale (1993)









obtained maximum ryegrass yields with 380 kg N ha-1, however, the maximum yield was

only 6.7 Mg ha-1. Allen et al. (1974) tested different levels of N in oat (Avena sativa L.)-

ryegrass mixed swards. Forage yield was 4.3 Mg ha-1 and 9.0 Mg ha-1 with 0 and 280 kg

N ha-1. Morris et al. (1994) reported herbage accumulation of 10.5 Mg ha-1 when 280 kg

N ha-1 was applied.

In Florida, Dunavan (1975) showed that three equal treatments with ammonium

nitrate (133 kg ha-1 N application-) applied at planting and in January and March

produced higher total yields than a single application of 400 kg N ha-1. Comparing three

N fertilization amounts, the rates of N recovery were 46, 45, and 26% for a single

application of ammonium nitrate to a rye-ryegrass mixture at N rates of 100, 200, and

400 kg ha-l, respectively.

The CP concentration of cool-season grasses is strongly influenced by the available

soil N. Application of N fertilizer to grasses usually increases CP concentration as well as

crop growth. The majority of the increased CP is non-protein N (NPN) in the form of

nitrate and free amino acids (Van Soest, 1982). Crude protein concentrations in annual

ryegrass tend to be high (Haby and Robinson, 1997), commonly averaging 150 to 200 g

kg-1 with no N applied and increasing to 280 g kg1- at N rates of 448 kg ha-1. The CP

concentrations of ryegrass samples collected in mid-January in different locations in

Texas were 228, 321, and 311 g kg-1 for Angleton, Overton, and Knippa, respectively,

(Lippke 1993, cited by Lippke and Ellis, 1997). Redfearn et al. (2002) analyzed nutritive

value of different cultivars of ryegrass. The CP and IVDOM results in January were 234,

229, and 232 g CP kg-1, and 849, 849, and 846 g IVDOM kg-1 for 'Gulf, 'Jackson', and









'Marshall', respectively. In general, CP concentrations of cool-season grasses range from

200 to 300 g kg-1 when well fertilized and in vegetative growth stage.

Lippke and Evers (1986) reported that ryegrass forage in vegetative stages of

growth usually has IVDOM > 700 g kg-1, approaching and sometimes exceeding 800 g

kg-1 in the first weeks of the grazing season (Ulyatt, 1981). In vitro digestibility of annual

ryegrass grown at different N rates has been evaluated in Louisiana (Allen et al., 1974).

The authors reported a 60 g kg-1 increase in digestibility at 448 kg N ha-1 compared with

that at 224 kg N ha-1. The cell wall digestibility and leaf/stem ratio of cool-season grasses

decreases with maturity. Data from Buxton and Russell (1988) for forages sampled

following the beginning of reproductive development indicate a decline in IVDOM from

761 to 552 g kg-1 in orchardgrass (Dactylis glomerata L.) leaves from May to July.

Buxton and Merten (1989) reported that herbage IVDOM of four cool-season grass

species decreased linearly with time during the spring. Plants with a moderate number of

reproductive tillers had an average rate of decline in IVDOM of 3.9 g kg-1 d-1.

Grazing Studies

Cool-season grasses in the southeastern USA usually are productive from late

December to early May. The use of small grains such as oat, wheat, and rye (Secale

cereale L.) in mixtures with ryegrass generally provides forage for grazing during the late

fall as well as during the winter (Rouquette et al., 1997). Small grains grow better from

late December to mid-February and ryegrass results in rapid forage growth during March

to late May, often requiring frequent increases in stocking rate to efficiently use ryegrass

pastures.

According to Rouquette et al. (1997), annual ryegrass is very tolerant of frequent

and severe defoliation and often supports 2000 kg ha-1 body weight for a 75- to 100-d









period in the spring. Depending upon management and climate, stocker calf gain may

range from 450 to 900 kg ha-1 from annual ryegrass pastures during 100 d of grazing.

In early studies, Riewe et al. (1963) compared 'Gulf ryegrass with tall fescue at

different stocking rates. The stocking rates ranged from 2.2 to 3.9 steers ha-1 for ryegrass,

and 1.9 to 4.9 steers ha-1 for tall fescue. The average daily gains for ryegrass were 0.67,

0.49, and 0.28 kg d-1, respectively, for low, medium, and high stocking rates. Maximum

gain per steer and gain per hectare were higher for ryegrass pastures than for tall fescue.

Rye is more productive than other annual cool-season grasses on sandy, well

drained soil of low fertility, while oat, wheat, and barley (Hordeum vulgare L.) do not

grow as well on very sandy soils. Rye produces more forage in fall than spring, while oat

DM production is variable (Phillips et al., 1996). Bransby and Gamble (1993) compared

ADG and LWG from pastures planted to 'Bonel' rye only or to Bonel rye plus ryegrass.

The inclusion of ryegrass resulted in greater stocking rates and ADGs, resulting in greater

LWG. At a stocking rate of five, 350-kg steers ha-l, LWG were 330 and 520 kg ha-l for

rye and rye-ryegrass, respectively. In Louisiana, Feazel (1986) evaluated Marshall and

Gulf ryegrass in pure stand and an 'Elbon' rye-Gulf ryegrass mixtures. The average

stocking rates were similar among the three treatments, 3.5 head ha-1. The LWG ranged

from 522 kg ha-1 for heifers grazing Gulf ryegrass to 688 kg ha-1 for Marshall ryegrass. In

the 3-yr study, there was no advantage to including Elbon rye in mixture with ryegrass. In

North Florida, Bertrand and Dunavan (1974) studied length of the grazing season of

calves (170 kg liveweight) grazing ryegrass and ryegrass-triticale (Triticale hexaploide

Lart.). The inclusion of ryegrass in the mixtures extended the grazing season from 533 to

650 animal days ha-l.









Feazel (1986) evaluated two methods of establishment of annual ryegrass, prepared

seedbed and sod-seeded, on performance of beef steers. Steers grazing prepared seedbeds

gained 1.05 kg d-1 and those grazing sod-seeded stands gained 1.25 kg d-1; however,

because initiation of grazing was 36 d later on sod-seeded pastures, total gain per steer

was 180 and 157 kg for prepared seedbed and sod-seeded pastures, respectively. In field

experiments in Louisiana, rye and ryegrass were sown alone or together into

bermudagrass stubble, or into a prepared seedbed. Dry matter forage yields over seven

cuts between December and May were 6.0 and 7.3 Mg ha-1 for ryegrass and rye-

ryegrass sown into a prepared seedbed and 3.9 and 4.8 Mg ha-1 for ryegrass and rye-

ryegrass sown into a bermudagrass stubble (Eichhom and Venuto, 1999). In some

environments, ryegrass has produced as much if not more ADG and LWG when planted

alone on prepared seedbed as in mixtures of ryegrass and small grains (Bransby, 1996).

Edwards et al. (1978) evaluated various rates of N fertilization (165 to 371 kg N

ha-) of Gulf ryegrass and wheat mixtures. Average daily gain was consistent across

treatments at 0.82 kg d-1. Total liveweight gain ranged from 622 kg ha-1 on pastures that

received 165 kg N ha-1 to 1030 kg ha-1 on pastures that received 371 kg N ha-1. Bransby

(1993) compared rye-ryegrass pastures under continuous and rotational stocking at 2.5,

3.7, and 5.0 (270 kg) steer ha-1 stocking rates. Average daily gain for calves on the

rotationally-stocked pastures was 0.82, 0.65, and 0.63 kg d-1, respectively, for the three

stocking rates compared to 1.05, 0.72, and 0.43 kg d-1 for the continuously stocked

pastures, respectively. Thus, an advantage for rotational stocking was obtained only at

high stocking rates. In Brazil, Restle et al. (2000) studied beef heifers grazing oat-









ryegrass pastures under continuous stocking and fertilized with 200 kg N ha-1. Average

daily gain and gain ha-1 were 0.58 kg d-1 and 429 kg ha-1, respectively.

Results of a 5-yr study were presented by Rouquette et al. (1992) who compared a

range of stocking rates on pure stands of ryegrass plus N and ryegrass-arrowleaf clover

(Trifolium vesiculosum Savi.), sod seeded on Coastal bermudagrass pastures. Cow-calf

pairs grazed the pastures. Average daily gains of suckling calves were similar among

treatments and ranged from 1.0 to 1.4 kg d-1 for ryegrass and arrowleaf pastures at

stocking rates of 5.98 to 2.37 animal units (AU) (680-kg liveweight) ha-1. The most

significant responses to the use of cool-season forages are not only calf ADG but also

weaning weight of fall-born calves. Using a fall calving season, the stocker phase is

essentially eliminated because calves are heavy enough to enter the feedlot immediately

upon weaning (Rouquette et al., 1997).

After an extensive research program with cool-season grasses, Rouquette et al.

(1997) concluded that achieving economic optimum grazing management and use of

cool-season grasses is not an easy task. A knowledge base of forage growth expectations

and the art of managing proper defoliation regimes will allow for the greatest opportunity

for positive economic returns and an acceptable transition from cool-season to warm-

season pastures.

Supplementation

Flexible management of the grazing system is required to maximize animal

performance since forage production and nutritive value are not uniform throughout the

year. The use of supplementary feed can overcome the shortfalls of the forage

component. When energy and protein requirements increase due to lactation, pregnancy,

and growth, part of the roughage component of the diet may need to be replaced by









concentrates (Fontaneli, 1999). Concentrates generally are more digestible than forages

and have much higher fermentation rates. According to Stockdale et al. (1987), several

factors may affect the response to supplement, including quality of the pasture and

supplement, amount of pasture and supplement fed, and degree to which supplemental

feeds replace pasture intake.

Protein

Crude protein concentration of high nutritive value herbage may exceed 250 g kg-1,

however, 650 to 850 g kg-1 of total protein is degradable in the rumen, with 150 to 350 g

kg-1 escaping the rumen. Microbial protein is a major protein source for ruminants;

therefore, optimizing microbial protein synthesis is essential. In a review of protein

supplementation of grazing livestock, Petersen (1987) stated that enhanced forage

utilization occurs in several ways. Protein supplements provide amino acids, carbon

skeletons, and minerals that help satisfy microbial requirements, thereby increasing

microbial growth and/or fermentation. These factors combine to increase intake through

increased microbial activity and rate of passage (Allison, 1985). In addition to increasing

available nutrients in the rumen, protein supplements may also increase the quantity of

protein reaching the small intestine through undegraded or bypass protein. Non-structural

carbohydrates are needed to utilize the forage N in the rumen, to reduce energy cost of

excreting N, and to supply nutrients to the small intestine. Microbial protein synthesis can

become a performance-limiting factor for ruminants consuming highly digestible

ryegrass diets. Hill (1991) observed that in young calves grazing ryegrass, synthesis of

microbial protein was 6 to 10 g 100 g-1 of OM digested in the rumen, and energy/protein

supplements increased the flow of microbial and undegraded dietary protein to the

intestines, and increased forage intake proportionally. In a study with beef steers grazing









rye-ryegrass pasture and receiving free-choice mineral, Grigsby et al. (1991) compared

corn meal and fish meal concentrates. Average daily gain was 1.5 and 1.0 kg d-1 for corn

meal and fish meal, respectively. Supplemental energy may have provided a source of

rapidly fermentable carbohydrate, which was well synchronized with ammonia and

peptide production from forage protein degradation, which in turn may have resulted in a

greater synthesis of microbial protein. Moore et al. (1999) suggested that the ratio of

digestible organic matter (DOM) to CP should be no more than seven to one based on the

observation that animals consuming forages with ratios greater than seven were likely to

respond to CP supplement.

Hill et al. (1990) studied beef heifers grazing ryegrass pastures (243 g CP kg-1) and

supplemented with maize, cottonseed meal, blood meal, feather meal, fishmeal, or

condensed molasses with cottonseed meal. Meal-based supplements were given at 3.5 g

kg-1 of body weight (DM basis). Forage organic matter intake (OMI) (650 g kg-1

digestible) was increased by blood and fishmeal supplements and decreased by

cottonseed and feather meal. It was concluded that blood and fish meal seemed to be

more effective in supplying required protein and essential amino acids that stimulate

intake of forage.

In C4 grasses, protein may be excessive or seriously deficient, depending on the

pasture species, season, and maturity of growth (Poppi and McLennan, 1995). Minson

(1990) compiled a series of studies in which CP was below 62 g kg-1 in the base forage.

There was an average 40% increase in intake due to protein supplement and a 34%

increase due to supplementary urea. Probably, the greatest and most uniform response to

supplemental N occurs when the CP concentration of the base forage is below 70 g kg-1









(Coleman et al., 2004). Bodine et al. (1999) reported the effect of four sources of ruminal

degradable protein and two sources of energy supplementation on steers consuming low-

quality hay (< 70 g CP kg1 OM). Forage OMI and total OMI increased quadratically as

rumen degradable protein increased for both energy supplements. When DOM:CP ratios

of limpograss [Hemarthria altissima (Poir.) Stapf and Hubb.] were between 8 and 10,

cattle responses to CP supplementation were significant (Lima et al., 1999b).

Hammond et al. (1994) suggested assessing blood urea N (BUN) concentrations as

a method to monitor the need for supplements by grazing animals when forage nutritive

value data are not available. Cattle BUN concentrations from 9 to 12 mg dL-1 are

considered to be a transition range below which gain response to protein supplementation

has been positive. Hoffman et al. (1993a) reported a summary of BUN values from

several studies with dairy cows. There was an apparent excess of N in the rumen in

relation to carbohydrates and wastage of highly degradable N in pasture forage when

BUN values ranged from 18.5 to 28.6 mg dL-1. Newman et al. (2002b) observed higher

BUN concentrations (18 vs. 15 mg dL-1) on heifers grazing limpograss either with or

without CP supplementation, respectively.

Energy

Energy supplements for forages generally fall into three categories: starch, sugars,

and fiber (Poppi and McLennan, 1995). Sources of fibrous highly-digestible energy, such

as soybean hulls or wheat middlings, have shown the most consistent response,

presumably because of the good synchrony with NH3 released (Poppi and McLennan,

1995). Johnson et al. (2001a) supplemented mature cows on bermudagrass pastures with

soyhulls or corn at 0.17% BW. There was no difference in dry matter intake (DMI) or

ADG between the treatments. These results suggest that low levels of corn









supplementation did not affect forage intake and animal performance. Supplementing

forage with a starch source, especially at levels > 250 g kg-1 of the total DM tends to

reduce fermentation of the basal forage [Goetsch et al. (1991), cited by Coleman et al.

(2004)]. According to Fieser and Vanzant (2004), when fed at similar levels of OM,

soybean hull supplementation provided an average of 60 g kg-1 greater DOM intake than

corn supplementation.

Ordonez-Tercero et al. (2003) used two energy sources [sugar-cane (Sacharum

officinarum L.), molasses and maize] to test the effect of energy supplementation on

intake and digestibility of the basal diet (elephantgrass) [Pennisetum purpureum Schum.]

and a foliage mix ofBrosimum alicastrum Sw.and leucena [Leucaena leucocephala

(Lam). Dewit] There were no differences in basal diet intake (5.54 kg DM d-1) among

treatments. The potentially degradable DM (337 g kg-1) and OM (364 g kg-1) fractions

were not affected by energy supplementation. Frizzo et al. (2003) studied the effect of

levels of energy supplementation on the productive and reproductive performance of

Charolais heifers maintained in a cultivated pasture of black oat (Avena strigosa Chreb.)

[Avena nuda L.]) plus annual ryegrass. It was shown that supplementation increased

ADG, stocking rate, and LWG. Heifers kept only on pasture had lower body condition

and showed lower estrus percentage than heifers supplemented with 0.7 and 1.4% of

LWG d-1. The estimated intake of DM was lower for the 1.4% BW d-1 supplementation

level. In South Brazil, Rocha et al. (2003) evaluated performance of beef heifers grazing

oat-annual ryegrass mixtures with and without ground maize supplement. The utilization

of oat and ryegrass pasture with supplementation resulted in higher ADG, stocking rate,

and LWG than the mixture with no supplement. Bertrand and Dunavan (1977) studied the









effects of increasing energy supplementation from 0 to 15 g kg-1 BW on performance of

beef steers (180 kg liveweight) grazing ryegrass-rye-white clover (Trifolium repens L.).

The increase in supplementation levels increased ADG (0.89-1.10 kg d-1), stocking rate

(4.09-5.32 head ha-1), and LWG (582-936 kg).

Interactions between Supplements and Forages

When forages are offered free choice and supplemental concentrates are offered in

restricted amounts, forage DMI may either increase, decrease, or remain the same

(Moore, 1992). In many cases, animal responses to supplements are either greater or less

than expected. The deviations between expected and observed performance are usually

explained by associative effects of supplements upon voluntary intake and digestibility of

the total diet (Moore et al., 1999). In general, concentrates will decrease forage intake

when forage quality is high, other nutrients are in balance with energy, and concentrate is

fed in large amounts. Large differences in substitution rates have been reported and the

effects have greater relation to differences among forages rather than to differences

among concentrates (Waldo, 1986). On the other hand, small amounts of concentrate may

increase voluntary intake when forages nutritive value is low, especially when the forage

has a high ratio of TDN to CP (Moore, 1994).

Sarker and Holmes (1974) fed supplement in increments of 2, 4, 6, or 8 kg OM d-1

to non-lactating cows grazing ryegrass. Though total OMI increased with increasing

amounts of concentrate, the average increase in forage intake was 0.46 kg OM kg-1 of

concentrate OM fed. Meijs (1986) fed high-starch supplements or high fiber supplements

to cows grazing predominantly perennial ryegrass (Lolium perenne L.) swards.

Supplement intake was 5.5 and 5.3 kg OM d-1 with forage intakes of 11.5 and 12.6 kg

OM d-1 for high and low starch treatments, respectively. Average forage substitution rate









for animals receiving supplements was 0.45 vs. 0.21 kg herbage kg-1 concentrate for

animals receiving starch and fibrous supplements, respectively. Royes et al. (2001)

evaluated different sources of energy (maize, sugar cane molasses, or soybean hulls) and

feeding rates (0, 1.4, or 2.8 kg DM steer-ld- in growth trials; 0, 150, or 300g kg-lofthe

ration DM in digestion trials) of supplements fed to cattle receiving ammoniated stargrass

hay. Increasing the level of supplementation decreased hay intake but increased total

dietary intake for all diets. Daily gain and feed efficiency of steers improved with

supplementation. Steers supplemented with corn or soybean hulls at 2.8 kg DM d-1 had a

higher ADG (0.92 kg) and gain:feed (0.103) than steers supplemented with molasses

(0.78 kg) at the same level.

According to Galloway et al. (1992), moderate dietary levels of supplement (200-

300 g kg-1) can improve nutrient intake and performance by cattle consuming

bermudagrass. At greater amounts, nutrient digestion, intake, or both, of the forage

portion of the diet can be affected negatively. Cattle grazing forages with DOM:CP ratios

greater than seven to eight are likely to have inadequate ruminally degradable protein and

should respond to N supplemention by increasing intake and animal performance (Moore

and Kunkle, 1999). Wheeler et al. (2002) tested the effects of increasing supplement

protein concentration on performance and forage intake of beef steers consuming

bermudagrass forage. Treatments were no supplement or daily equivalents of 0.2, 0.4,

and 0.6 g of supplemental protein kg-1 of BW. Forage intake increased 16% and total

OMI increased 30% in supplemented compared to unsupplemented steers. Diet OM

digestibility increased 145 g kg-1 in supplemented compared to unsupplemented steers.









A special case of substitution and associative effects is the use of total mixed

rations (TMR) for lactating dairy cows. Such diets may include several sources of forage

and combinations of concentrates. Because TMRs often are based on high quality forage

and include high percentages of concentrates, intake and digestibility of the forage

component is very likely less than expected when forage is fed alone (Moore, 1994). Fike

et al. (2003) studied the effects of two levels of supplementation on forage and total

intake of lactating dairy cows grazing bermudagrass and rhizoma peanut (Arachis

glabrata Benth.) pastures. The substitution of forage OM by supplement OM (kg kg-1)

was 0.48 for rhizoma peanut and 0.06 for bermudagrass. Feeding additional supplement

increased total OMI by 230 and 9 g kg-1 for cows grazing bermudagrass and rhizoma

peanuts pastures, respectively.

Behavior

According to Stobbs (1975), the energy required to prehend forage from the

standing biomass is an insignificant part of the maintenance requirements, but energetic

costs of activities associated with grazing have been estimated to account for 25 to 50%

of an animal's daily energy requirements. Numerous studies have shown that supplying

supplemental protein and (or) energy to cattle consuming low- to moderate-quality

forages can increase animal performance and forage OMI, however, data are limited

showing the effect of supplementation on grazing behavior (Krysl and Hess, 1993).

Adams (1985) postulated that disruption of normal grazing activity resulting from

supplementation regimens could adversely affect forage intake and animal performance.

In research with yearling beef steers grazing ryegrass, the same authors noticed that

although there was an 11.3% decrease in forage intake with supplementation, total daily

grazing time was not altered. Macoon (1999) observed that dairy cows supplemented









with high levels of concentrate spent less time grazing during daylight (135 min) than

cows that received low levels of concentrate (180 min). Supplementation can cause shifts

in daylight grazing behavior patterns; however, there is no indication that supplemented

animals alter their percentage of daytime vs. nighttime grazing compared to

unsupplemented animals (Adams, 1985). Providing protein supplements to cattle grazing

low-quality forages increased harvest efficiency (forage OMI kg-1 BW min- spent

grazing) from 8 to 60% compared with unsupplemented cattle (Barton et al., 1992). In

contrast to protein meal-based supplements, high-starch supplements either failed to alter

or decreased harvest efficiency (Adams, 1985).

Allden and Whittaker (1970) indicated an upper limit of grazing time of- 600 to

720 min d1 for sheep. Similarly, Stobbs (1975) observed that fatigue limits the time that

can be spent grazing to 720 min d-1 in cattle. Numerous studies have reported daily

grazing time to be in the range from 359 to 771 min d1 for cattle over a wide range of

environmental temperatures, supplementation regimens, grazing management practices,

and forage types (Krysl and Hess, 1993).

Intake

Intake is the most important determinant of performance by grazing ruminants

(Lippke, 2002). As much as 60 to 90% of the variation in digestible energy intake may be

due to animal variability, with 10 to 40% due to diet digestibility (Crampton et al., 1960).

The most important mechanisms controlling intake in ruminants are metabolic,

distention, and behavioral. Conrad et al. (1964) examined results from 114 trials with

lactating cows and reported the situations where physical and physiological factors were

regulating feed intake. Intake of diets having digestibility between 500 and 670 g kg-1

was thought to be limited by physical factors such as digestibility of the feed and its rate









of passage through the digestive tract. Intake of diets having a digestibility > 670 g kg-1

was limited primarily by physiological control mechanisms. Intake of perennial warm-

season grasses is considered to be limited by physical (distention) mechanisms due to

their high fiber concentration and low digestibility. Van Soest (1965) reported negative

correlations between cell wall concentration and voluntary intake.

Factors controlling ruminant intake should be assumed to function through multiple

interactions, and the simplistic view should be abandoned that only physical factors limit

the intake of roughage diets and that only chemical or physiological factors limit intake

of concentrate diets (Fisher, 2001; Moore, 1994). Hodgson (1982) stated that under

grazing conditions, ingestive behavior (i.e., bite mass, bite rate, and grazing time) might

be a more important determinant of forage intake than physical and metabolic controls.

Low forage availability and accessibility are situations where ingestive behavior may be

an important controlling factor on forage intake. The DMI on pastures is a result of a

complex interaction among pasture canopy characteristics, ingestive behavior, and

grazing time (Burs and Sollenberger, 2002).

Forage intake is forage species-specific and altered by plant maturity and

morphology. The eating behavior of ruminants has been clearly characterized as

selective, with strong preference for green leaf and against dead and stem tissues

(Minson, 1990). Greater selectivity by the animal for leafy tissue from within the bounds

of forage on offer will generally result in a higher quality diet than that on offer and

greater DMI (Burns et al., 1991).

Measurements of Forage Intake of Grazing Animals

The ability to measure or predict DMI and nutritive value of forages on pastures

has been the impetus for studies of the plant-animal interface (Burs and Sollenberger,









2002). A number of techniques have been adopted to estimate intake because it is not

practical to measure intake on pastures directly (Lippke, 2002). There are two general

approaches to the indirect estimation of pasture intake. According to Moore and

Sollenberger (1997) the techniques may be classified as follows: 1) estimates for

individual animals by fecal output and forage digestibility or ingestive behavior, and 2)

estimates for groups of animals by disappearance of herbage mass or calculation from

animal performance. Each method employs different assumptions which must be met if

the estimates are to be valid.

To avoid errors associated with the estimation of forage intake on pastures,

researchers have cut and carried green pasture herbage to confined animals kept in

confinement. Though this approach affords a great degree of precision, it may be highly

inaccurate because it reduces the opportunity for diet selection. Experimental results are

likely most affected when swards are highly heterogeneous, and when environmental

factors, sward density, and forage accessibility have large effects on grazing behavior.

Intake calculations based on fecal output and forage digestibility have been used

extensively. Fecal output is the measure of interest because it can be used to calculate

intake using the following equation: Intake = fecal output / (100 diet digestibility).

Early attempts to estimate intake from total feces collection in individual animals

included use of fecal collection bags. In addition to the potential for loss or urine

contamination due to poor design or lack of fit, the bags also have the potential to stress

the animal and to alter intake by changing grazing behavior.

The alternative is to estimate fecal output using marker technology. Functionally,

markers are classified as being internal to the feedstuff or external, i.e., added to the









feedstuff or dosed separately to the animal (Lippke, 2002). Markers are reference

compounds used to investigate both chemical and physical digestive processes (Owens

and Hanson, 1992). Characteristics of an ideal external marker were outlined by Owens

and Hanson (1992) and include the following traits: 1) it should be unabsorbable, 2) it

should not affect or be affected by animal or microbial digestive processes, 3) its flow

should closely mimic that of the material it marks, and 4) it must be analyzable with a

specific and sensitive methodology. No single marker currently meets all of these criteria.

Chromium oxide has been the most widely used external marker for estimation of

intake during the last 50 yr (Huston et al., 1999). The primary problem associated with

this marker is that it moves through the digestive tract independently of the undigested

particles of the diet, and fecal concentrations of chromium may exhibit strong diurnal

variation. Measuring daily DM intake with Cr requires a continuous period of Cr intake

and frequent spot feces collection during different hours on the days following dosing.

This additional handling of the animals is undesirable, particularly when it has potential

to disturb established patterns of grazing behavior.

An alternative method of calculating fecal output from a single dose of Cr has been

presented by (France et al., 1988). Animals are dosed once with labeled feed fractions,

and numerous fecal samples are collected over a period of time long enough for the label

source to clear the animal. A nonlinear equation relating time after dosing to fecal Cr

concentration is used to generate parameters for the estimation of fecal output (Pond et

al., 1989).

According to Buntinx et al. (1992), controlled-release Cr capsules offer the

potential of reduced labor, coupled with the possibility of preventing adverse effects on









animal behavior from repeated daily dosing, or from the presence of fecal bags during

total fecal collection. Controlled-release devices greatly reduce the within-day coefficient

of variation for fecal concentration of Cr in sheep and cattle when compared with daily

dosing (Adams et al., 1991). However, total fecal collection must be used on a subset of

animals to correct for possible payout differences from manufacturer specifications

and/or subsampling influences on chromium recovery.

Dosed and plant alkanes are options as external and internal markers, respectively.

Mayes et al. (1984) showed that dosed even-chain length n-alkanes and naturally

occurring odd-chain plant n-alkanes could be used as fecal markers to determine forage

intake. Forage species contain variable quantities of alkanes, and the concentration of

odd-chain alkanes may be too low in some tropical forages for them to be used as an

internal marker (Laredo et al., 1991). The problems associated with species-specific

alkane concentrations and analytical determination of the n-alkanes make the technique

less likely to be used in a wide range of environments.

Associated with the use of inert markers in estimating DM intake of grazing

animals is the requirement of knowing the digestibility of the animal's diet. Obtaining

samples that represent the diet of the dosed animals can frequently be the major factor

limiting the accuracy of the method. Surgically altered animals and "hand plucked"

samples are the two most-used methods to obtain samples of the diet in grazing studies.

After a representative sample is collected, the accuracy of the marker technique resides

with the method used to estimate the absolute digestibility of the diet (Burns et al., 1991).

In vitro dry matter digestibility has been the most-used method because it is cheap,

precise, and in most of the cases, well correlated with in vivo digestibility. Although in









vitro dry matter digestibility (IVDMD) has perhaps been most widely used to predict

digestible DM, a variety of internal markers also have been used.

Ingestive behavior has been suggested as a method of estimating daily DM intake

(Coleman et al., 1989). The parameters to be measured are bite mass, bite rate, and

grazing time. Mechanical methods for short-interval measurements have been reviewed,

but recent advances have occurred through automation to aid quantification. Bite rate

may be recorded by observing, listening, or by electronic devices, however, it is

necessary to discriminate between biting and non-biting jaw movements. Artificial

swards can be used in measurement of bite weight, or esophageal extrusa can be collected

over a short period of time. Finally, grazing time can be recorded visually or by using

electronic devices. Errors associated with these devices and especially with extrapolation

of intake during short measurement periods to daily intake are the most common.

If the pastures are uniform and under intensive management, forage disappearance

can provide a reasonable estimation of forage intake. Forage mass is measured before and

after grazing, and the forage that disappears is considered to have been consumed. The

technique is very labor intensive because a large number of sites have to be measured to

provide an acceptable estimation of herbage mass. The difficulty of achieving accurate

estimations of herbage mass, as well as trampling, defoliation by insects, and senescence,

make the herbage disappearance technique not reliable enough to be recommended for

routine estimates of pasture intake (Moore and Sollenberger, 1997).

The animal performance technique, also called "reverse energy calculations", is the

calculation of intake by using, in reverse, accepted energy requirements for maintenance,

growth, and lactation (McMeniman, 1997). For a given rate of performance, the daily









amount of energy that must be consumed in order to achieve that level of performance

can be obtained from tables and equations. Accurate determinations of changes in body

weight are required when using this technique. Digestibility is the main determinant of

forage energy concentration and efficiency of energy utilization (Moore and

Sollenberger, 1997). Digestibility of the forage can be obtained by the methods reported

before, and concentration of digestible energy (DE) is calculated by assuming 4.4 Mcal

DE kg-1 DOM. Concentration of metabolisable energy (ME) is calculated by multiplying

DE concentration by 0.82 and net energy for maintenance and for gain are calculated

from ME concentration by (NRC, 1996). The main constraint of this technique is that

factors other than forage energy concentration, such as nutrient imbalances and

environmental effects may influence the efficiency of conversion of net energy for gain

to animal product.

N Fractioning and in situ Methodology

Diet formulation systems have been developed that attempt to balance diets for

both ruminal protein degradability and carbohydrate availability with the goals of

maximizing microbial protein synthesis and minimizing protein wastage as ammonia

(Nocek and Russell, 1988). Rumen protein degradability is one of the most important

qualitative factors determining the protein value of a feed. Protein degradability

determines the supply to the rumen microbes of both ammonia and branched-chain amino

acids, as well as the supply to the small intestine of the rumen-undegraded feed protein as

a potential source for amino acid absorption.

The protein concentration in feeds may be expressed on a CP basis. The CP is

composed of two fractions: rumen degradable protein (RDP) and rumen undegradable

protein (RUP). The RDP consists of non-protein N and true protein while the RUP is









formed only by true protein. The protein degradation rate will vary depending on the

proportions of NPN, true protein that degrades at variables rates, and unavailable protein

(Pichard and Van Soest, 1977).

In situ Procedure

Numerous methods currently are available for estimating ruminal DM and N

disappearance of feedstuffs. The in situ technique involves the placement of indigestible

bags containing a feedstuff into the rumen of a fistulated animal for various time intervals

and the measurement of the amount of DM and N that disappears from the bags over time

(Ster and Satter, 1984). According to Nocek (1988) there is no better way to simulate

the rumen environment within a given feeding regime, although in the ruminal

environment, the feed sample in the bag is not subject to the total ruminal experience:

i.e.; mastication, rumination, and passage.

The in situ procedure can be used to quantify the protein fractions, the instantly

degradable Fraction A; the undegradable fraction after an extended rumen incubation,

Fraction C; and Fraction B, the slowly degradable fraction, obtained by difference

(%Fraction B= 100 % Fraction A % Fraction C). Although the technique allows the

quantification of the different N fractions of the forage as well as the rate of digestion of

B, it cannot be used to quantify the rate at which A is degraded. Nocek (1985)

demonstrated that a certain portion of the test feed escapes the bag prior to ruminal

degradation. This fraction is generally assumed to be readily available to rumen microbes

and digested at a rapid rate. Potentially degradable nutrient fractions have been described

by first-order kinetic rate constants (Mertens, 1973). Primary assumptions are that the

pools in question are homogeneous and that the substrate remaining will be degraded as a

linear function of time in the rumen. Orskov and McDonald (1979) and other researchers









have developed mathematical models to fit the estimated ruminal degradability of

feedstuffs. The most common approach included the use of non-linear regression.

Mertens and Loften (1980) proposed the inclusion of lag time for protein degradation in

the models used for fitting N disappearance. If a delay appears at the beginning of the

disappearance of Fraction B, a lag component should be included in the model

(McDonald, 1981).

Various aspects of the in situ technique interact and can influence the interpretation

of the results. Bag porosity is a compromise between limiting the influx of rumen

contents not associated with the test feed and allowing influx of microbes to degrade the

test feed, while at the same time limiting the efflux of undegradable feed particles

(Nocek, 1988). Uden and Van Soest (1984) showed cell wall digestion of timothy

(Phleum pratense L.) increased with increasing bag pore size (20, 37, and 53 tm).

Digestion of DM was similar for pore size ranging from 40 to 102 [am, and estimated

rumen N availability was greater for bags with pore size of 6 or 20 [am (Uden and Van

Soest, 1984).

Feed particle size is another important aspect in the in situ disappearance

determination. Generally, longer and coarser material is associated with slower rates of

digestion and greater variation. However, finely ground materials are subject to greater

mechanical losses from the bags (Nocek, 1988). The same author states different

feedstuffs can require different particle sizes, and the particle size for cereal grains,

fibrous products, forage, and silages ranges from 2 to 5 mm.

Sample weight is also an important consideration. The optimum sample weight is

that which provides enough residue at the end of the extended rumen incubation for









chemical analysis without over-filling the bag so as delay bacterial attachment, increase

lag time, and underestimate digestion rates (Nocek, 1988). For forages, Van Hellen and

Ellis (1977) recommends 10 mg cm-2ofbag surface area, whereas Uden and Van Soest
-2
(1984) recommend < 6 to 7 mg cm-2

The host animal diet is the major factor determining quantity and types of microbes

and therefore the rate and extent of digestion of dietary nutrients. de Faria and Huber

(1984) evaluated in situ DM digestion of corn silage, alfalfa (Medicago sativa L.) silage,

and grass hay in diets containing three different protein (81, 113, 133 g kg-1) and three

different acid detergent fiber (ADF) concentrations (390, 299, and 210 g kg-1). Neither

protein nor ADF of the host animals' diet had a significant effect on DM digestibility for

any forage tested, however, specific sampling time by forage interaction was noted.

The animal species most commonly used for in situ experiments are cattle and

sheep. Siddons and Paradine (1983) compared sheep and steers fed similar diets at

maintenance. Sheep had higher ruminal ammonia N than steers, lower VFA, and similar

ruminal pH and rumen dilution rate.

In situ Disappearance of Warm- and Cool-season Grasses

High quality cool-season pastures and many tropical legumes would be expected to

have high ruminal ammonia losses when their CP concentrations exceeded 150 g kg-1

because of the high concentration of ruminal degradable protein (Poppi and McLennan,

1995). According to Beever (1984), 800 to 900 g kg-1 of ryegrass CP can be degradable

in the rumen. Van Vuuren et al. (1991) stated that OM and CP in situ total degradability

decreased with increasing ryegrass sward maturity and with decreasing rate of N

application. Salaun et al. (1999) studied the relation between N fertilization and rumen

CP degradability of perennial ryegrass. They concluded that reducing N fertilizer









application decreased the theoretical degradability in the rumen due to decrease in the

NPN. The total N disappearance was 600 and 710 g kg-1 for plots fertilized with 250 and

550 kg N ha-1. Assis et al. (1999) did not find significant difference of CP Fraction A in

Tifton 85 bermudagrass with levels of N fertilization from 0 to 400 kg ha-l. Yan and

Agnew (2004) tested the effective degradability of perennial ryegrass N assuming a

ruminal outflow rate of 0.02, 0.05, or 0.08 % h-1. The objective was to use these data to

develop prediction equations for N degradability in C3 grass silages. At 0.02 % h-1

ruminal outflow, total N degradability was 934 g kg-1. The N degradability was

negatively related to ADF, NDF, and lignin concentrations but positively related to CP

concentration.

Warm-season grass CP tends to be more slowly degraded in the rumen than that of

cool-season grasses (Minson, 1990). Akin and Burdick (1975) concluded that

digestibility differences between C3 and C4 grasses were associated with the parenchyma

bundle sheath (PBS) of C4 grasses. The C4 plants have a lower soluble protein

concentration (260 to 300 g kg-1) than C3 plants (330 to 480 g kg-1), which is thought to

be mainly due to differences in Rubisco concentration and location. Further, PBS tissue

from C4 plants, the primary location for Rubisco, is slowly or incompletely digested in

the rumen (Akin and Burdick, 1975).

Redfearn et al. (1995) studied the rates of ruminal protein disappearance for

switchgrass (Panicum virgatum L.), big bluestem (Andropogon gerardii Vitman), and

smooth bromegrass (Bromus inermis Leyss.) and suggested that generalizations regarding

ruminal protein degradability should not be made among forage species. The total protein

disappearance was 847, 795, and 974 mg g-1 for switchgrass, big bluestem, and smooth









bromegrass, respectively. Mandebvu et al. (1999) compared in situ disappearance of

Tifton 85 and coastal bermudagrass. There was no difference in the potentially digestible

fractions (A+B) among the cultivars with approximate values of 600 g kg-1. Assis et al.

(1999) evaluated the in situ N disappearance of Cynodon spp, Tifton 44 and Tifton 85

bermudagrasses, and 'Puerto Rico' stargrass. Tifton 85 had the smallest concentrations of

the potentially digestible CP fractions (A+B), but it had the largest B fraction

concentration. Johnson et al. (2001b) reported that with no fertilization, the Fraction A

concentration was 212 (bahiagrass), 310 (bermudagrass), and 279 g kg-'(stargrass) of the

total N, but upon application of 160 kg N ha-1 per cutting, Fraction A represented 285

(bahiagrass), 400 (bermudagrass), and 421 g kg-'(stargrass) of the total N. Newman et al.

(2002a) fractionated the N content of limpograss at different grazing heights. Pastures

grazed to 20 cm had the greatest proportion of soluble N and the least portion of

undegradable CP. They concluded that low total CP in the DM, resulting in low

concentrations of CP Fractions A and B in the rumen, and the long lag phase for

degradation of the B fraction, may contribute to reported protein deficiencies of cattle

grazing limpograss.

Other Methods to Estimate Ruminal Protein Degradation

Although in vivo and in situ procedures for estimating degradability of protein are

not without drawbacks, they are typically considered to most accurately reflect reality.

However, labor requirements, cost, time, variability, and the need for access to fistulated

ruminants limit the feasibility of widespread use of these procedures for routine analysis

(Mathis et al., 2001). In vitro estimation of rumen-degradable protein is appealing

because it addresses many of the limitations of in situ and in vivo procedures and would

be relatively easy to standardize (Ster et al., 1997). While numerous in vitro methods are









currently available for estimating ruminal protein degradation of feedstuffs, assays based

on Streptomyces griseus are probably the most commonly used (Roe et al., 1991).

In the Streptomyces griseus method, feeds are incubated in a borate-phosphate

buffer at pH 7.8 to 8.0 to facilitate optimum enzyme activity, and enzyme is added at a

level of 6.6 units of enzyme g-1 sample DM (Krishnamoorthy et al., 1983). Streptomyces

griseus is a broad spectrum, commercially available protease that has both exopeptidase

and endopeptidase activity, as in the rumen.

Ruminal protein degradation also has been estimated by the ficin method, which

breaks down protein by proteolytic ficin (Ficus glabrata) used at 8.24 units g-1 of sample

DM in a phosphate buffer at pH 6.5 (Poos-Floyd et al., 1985). Finally, the neutral

protease with amylase method makes use of an endopeptidase, Bacillus subtilus, at 9.6

units g-1 sample DM, in a sodium citrate buffer, in addition to an enzyme that contains

endo-P-glucanase and a-amylase activities (Assounami et al., 1990).

Roe et al. (1991) compared the three in vitro methods of ruminal protein

degradability with the in situ methodology. None of them resulted in feed protein

degradation curves that had a consistent relationship with those generated by the in situ

technique. However, relationships were found between protein degradability estimates

obtained by the neutral proteases with the amylase method at specific time points and

those measured in situ.














CHAPTER 3
SUPPLEMENTATION EFFECTS ON FORAGE CHARACTERISTICS AND
PERFORMANCE OF EARLY WEANED CALVES GRAZING RYE-RYEGRASS
PASTURES

Introduction

Grassland covers 4.5 million ha in Florida and most of this area is utilized by

beef cattle. Florida's beef cattle industry is based on cow-calf enterprises with a total of

950 000 beef cows. Sales of cows and calves accounted for $348 million of income to

Florida farmers and ranchers in 2003 (Florida Agriculture Statistics, 2004).

The profitability of the Florida cow-calf enterprise is largely dependent on the

cow's productivity. The combination of low quality warm-season forages and forage

quantity limitations during winter reduce cow reproductive performance. Early weaning

the calves of young cows and first-calf heifers is a management strategy that has potential

to address this problem. At early weaning, cow requirements for total digestible nutrients

(TDN) decreases by as much as 49%. As a result they are able to restore body condition

rapidly (Arthington and Kalmbacher, 2003). Heifers whose calves were weaned early had

higher pregnancy rates (94%) than those whose calves were weaned at the normal time

(65%) during a 2-yr study in South Florida (Arthington and Kalmbacher, 2003).

Although the benefits of early weaning, i.e., improving reproduction and reducing

nutrient requirements of the cow, have been recognized for many years, the factor

limiting practical application of early weaning has been management of the early weaned

calves. Mild winters in the southern USA offer an opportunity to raise calves on pasture

systems that include high nutritive value winter-annual forages (Arthington and









Kalmbacher, 2002). Annual ryegrass (Lolium multiflorum Lam.), a high-yielding, high

quality grass, is the most commonly grown cool-season pasture forage in the southern

and southeastern USA (Evers et al., 1997). The use of small grains such as oat (Avena

sativa L.), wheat (Triticum aestivum L.), and rye (Secale cereale L.) in mixtures with

ryegrass generally extends the grazing season (Rouquette et al., 1997). Small grains grow

better from late December to mid-February, and ryegrass is most productive during

March to late May.

The practice of raising early weaned calves on winter pastures is a feasible

alternative, especially if pasture quality and quantity are superior to hay available for

winter feeding (Barnes et al., 1996). Weder (1997) reported average daily gain of 0.8 kg

d-1 for early weaned calves grazing annual ryegrass, however, the limited ruminal

capacity of early weaned calves at young ages may limit forage intake and result in low

animal performance. Early weaned calves grazing wheat pastures consumed 27% less

forage dry matter (DM) during the first 20 d exposed to pastures than during the

subsequent 50 d (Paisley et al., 1998). The authors suggested that initial performance of

early weaned calves was limited by low forage intake. Moreover, calves grazing high

quality cool-season pasture may have high ruminal ammonia losses, if forage crude

protein (CP) concentrations exceed 150 g kg-1 (Poppi and McLennan, 1995). Loss of N

from the rumen is costly due to significant energetic expenditure associated with urea

synthesis and excretion. Non-structural carbohydrates are needed to utilize the forage N,

to reduce energy cost of excreting N, and to supply nutrients to the small intestine.

Microbial protein synthesis can become a performance-limiting factor for ruminants

consuming ryegrass diets. Rocha et al. (2003) evaluated beef heifers grazing oat-annual









ryegrass with and without ground maize (Zea mays L.) supplement. Supplementation

resulted in higher average daily gain, stocking rate, and production per unit of land area.

An effective supplementation program is necessary to overcome potentially low

forage intake and maximize the efficiency of utilization of cool-season grasses in early

weaned calf systems. The objective of this study was to evaluate the effect of different

levels of concentrate on performance, stocking rate, gain per hectare, and forage intake of

early weaned calves grazing rye-ryegrass pastures in North-Central Florida.

Material and Methods

The study was located at the Beef Research Unit, 18 km northeast of Gainesville,

FL (30 N). Data were collected during the 2003 and 2004 winter-spring seasons. The

soils at the research site are Adamsville fine sand (Aquic Quartzipsamments,

hyperthermic, uncoated) and Pomona sand (sandy, siliceous, hyperthermic Ultic

Alaquod). These soils have from poor to somewhat poor drainage with rapid

permeability. Prior to initiation of winter grazing, mean soil pH (1:2 soil:deionized H20

ratio) was 6.5, and Mehlich-I (0.05 MHC1 + 0.0125 MH2SO4) extractable P, K, Mg, and

Ca in the Apl horizon (0- to 15-cm depth) were 52, 19, 117, 761 mg kg-1, respectively.

Starting dates for the trial were 28 Jan. 2003 and 15 Jan. 2004. Grazing ended on

14 May 2003 and 29 Apr. 2004. Monthly rainfall totals and temperature data during the

trial are presented in Table 1 (Appendix A).

In early October 2003 and 2004, the area was sprayed with 3.0 L ha-1 of glyphosate

(Monsanto Co., St. Louis, MO; isopropylamine salt [10 g kg-1] of N-phosphonometyl

glycine) to kill bahiagrass (Paspalum notatum Flugge). Three weeks later, the cool-

season forages were planted using a no-till drill. The seeding rates were 20 kg ha-1 of

'Jumbo' ryegrass and 80 kg ha-1 of a mixed-blend rye ('Grazemaster'). All pastures









received an initial application of 40 kg N ha-1, 17 kg P ha-1, and 66 kg K ha-1, 3 wk after

planting. Additional applications of 40 kg N ha-1 were made in late December, early

February, and early March in both years.

Calves were weaned on 2 Jan. 2003 and 5 Jan. 2004 at -90 d of age and 100 kg of

body weight (BW). They were held in dry lot for 7 d with access to bermudagrass

[Cynodon dactylon L. (Pers.)] hay (ad libitum), 1 kg of pre-conditioning medicated

concentrate d-1, and water. Early weaned calves were Angus-sired (crossbred-cows sired

by Angus bulls) products from primiparous and multiparous cows in 2003 and 2004,

respectively. Calves were dewormed with Ivermectin 10 g kg-1 concentration (Ivomec,

Merck & Company, Rahway, New Jersey, USA) on 10 Apr. 2003, and 12 Feb. and 29

Apr. 2004.

Pasture size was 0.2 ha, and each pasture was subdivided into four paddocks.

Pastures were rotationally stocked with a 7-d grazing and 21-d resting period. Two early

weaned calves (1 steer and 1 heifer) were assigned as testers to each experimental unit

such that BW was equal (+ 5 kg) across all experimental units. "Put and take" early

weaned calves of comparable age and weight to the testers were used to maintain similar

herbage allowance across experimental units. The literature does not define specific

herbage allowance targets for the species in this study, thus our primary objectives in

managing pasture quantity were to avoid i) low herbage allowances that would limit calf

ADG and ii) variation among experimental units in herbage allowance within a year that

could be confounded with the responses to supplement treatment. It was also recognized

that variable growth conditions, especially due to rainfall, could result in year differences

in herbage allowance.









Treatments were three levels of a commercial pelleted concentrate, 10, 15, and 20 g

kg-1 of calf BW, offered daily. Each treatment was replicated three times in a completely

randomized design, so there were a total of nine experimental units in the study. In 2004,

for observation purposes, two experimental units were added to the study in which the

EWC did not receive concentrate supplementation. The concentrate composition is

described in Table 3.1. A salt-based trace mineral mix was supplied free choice

throughout the grazing season.

Pasture Sampling

In each 28-d grazing cycle, herbage mass was determined twice using a double-

sample technique (on Paddocks 1 and 3). Indirect estimates of herbage mass were done

using a 0.25-m2 aluminum disk plate meter, and direct measures were done by clipping

herbage in the same area to a 3-cm stubble. Twenty disk settling height measures were

taken the day before the paddock was grazed and twenty were taken at the end of the

grazing period. Twenty, 0.25-m2 sites, representing the range of herbage mass on the nine

pastures, were selected for disk calibration once a month. Disk height was measured and

the herbage at these sites was then clipped to a stubble height of 3 cm, dried at 600C in a

forced-air drier to constant weight, and weighed. A calibration equation was developed

using these double sampling data by regressing actual herbage mass on disk height. To

predict herbage mass for any given pre- or post-graze sampling date, the average of the

20 disk heights measurements was calculated and entered into the calibration equation.

The regression equations and r2 for each double sampling date are shown in Table 2

(Appendix A).









Table 3-1. Ingredient and chemical composition of supplement fed to early weaned
calves on pasture (DM basis).
Ingredients Concentration (g kg-1)
Corn meal 25.0
Wheat middling 400.0
Soybean meal (48% CP) 25.0
Cottonseed meal (41% CP) 50.0
Soybean hulls 392.5
Cottonseed hulls 32.5
Calcium carbonate 12.5
Salt 6.3
Quadra-plex 1.3
Molasses 50.0

Chemical compositionT (g kg-lDM)

Dry matter 900.0
Ash 62.3
Crude protein 146.9
TDN 700.0
Calcium 7.6
Phosphorus 5.7
Salt 6.3
Sodium 3.7
Magnesium 2.5
Potassium 12.6
Sulfur 1.4
Cobalt (mg kg-1) 3.36
Cooper (mg kg-1) 30.48
Iodine (mg kg-1) 0.12
Iron (mg kg-1) 200.12
Manganese (mg kg-1) 111.84
Selenium (mg kg-1) 0.15
Zinc (mg kg-1) 103.13
t Provided by the manufacturer (Lakeland Animal Nutrition)
Pregraze herbage mass during a grazing cycle was calculated as the average of

Paddocks 1 and 3. Herbage accumulation for a given paddock was calculated by

subtracting postgraze herbage mass of cycle n-1 from pregraze herbage mass of cycle n.

Average herbage allowance was computed as the ratio of average herbage mass

[(pregraze + postgraze) / 2] to early weaned calf liveweight exposed to the experimental









unit during the grazing period (Sollenberger et al., 2005). Stocking rate (SR) was a

response variable (variable SR study) and was expressed as animal units (AU; 1 AU=

500 kg of liveweight 0.75) ha1.

Hand-plucked samples were used to estimate nutritive value of the grazed portion

of the canopy. Samples were taken at 20 locations per paddock on the day prior to

grazing and herbage was removed to the stubble height at which the previous paddock

was grazed. These samples were dried at 600C in a forced-air drier to constant weight,

ground in a Wiley mill to pass a 1-mm stainless steel screen, and taken to the laboratory

for analyses. Nitrogen concentration was measured using a modification of the aluminum

block digestion technique (Gallaher et al., 1975). Concentration of CP in herbage DM

was calculated as N x 6.25. In vitro digestible organic matter (IVDOM) concentration

was determined by the two-stage procedure of Tilley and Terry (1963) modified by

Moore and Mott (1974).

Rumen degradable protein and rumen undegradable protein were estimated by the

in vitro method proposed by Roe et al. (1991). Hand-plucked samples from 15 g kg-1 BW

supplement treatment pastures were analyzed. Samples were incubated in a

buffer/protease solution for 48 h, the residue recovered through filtering, and it was

analyzed for CP concentration. Rumen degradable protein is that which was digested

during 48 h, and rumen undegradable protein is the difference between the original CP

content and the rumen degradable protein.

Animal Response Variables

Total organic matter intake (OMI) was computed based on fecal output and diet

digestibility, viz., total OMI= OM output of feces / (1- [OM digestibility/ 100]). Fecal

output was estimated using a controlled release Cr marker (Captec New Zealand Limited,









Auckland, New Zealand). The devices were administered orally to tester calves on 25

Mar. 2003 and 26 Feb. 2004. Four fresh fecal samples were obtained during the morning

on Days 11, 13, 15, and 17 after administration.

In addition, four early weaned calves fitted with the controlled release chromium

marker were used for total fecal collection from Days 11 to 17 after dosing. These four

calves were confined in metabolic crates and fed freshly harvested rye-ryegrass ad

libitum and were supplemented with 15 g kg-1 BW concentrate. The total daily fecal

output from each calf was collected, weighed, mixed and a subsample analyzed for DM

and Cr concentration. Total Cr output was calculated by multiplying the fecal Cr

concentration by total daily fecal weight. The total daily Cr output was used to estimate

total fecal output of the tester calves by relating total daily Cr output and Cr

concentration in the feces OM. Individual daily total Cr output is described in Tables 3

and 4 (Appendix A).

Fecal samples were dried at 600C in a forced-air oven and ground in a Wiley mill to

pass a 4-mm stainless steel screen. Chromium concentration in the feces was assayed by

atomic absorption spectrophotometry following the procedure described by Williams et

al. (1962). The samples were analyzed by d of collection in duplicate and analyses were

repeated for samples where the difference of Cr concentration between duplicates

exceeded 100 g kg-1

Total OMI was calculated based on fecal output estimations and IVDOM of the

respective pasture herbage. Forage OMI was the difference between total OMI and the

known amount of concentrate OM fed. The fecal output estimated with the marker did

not equal the fecal output predicted based on estimated forage and supplement









digestibilities. For this reason, an interactive SAS Institute Inc. (1991) program was used

to adjust the total OMI based on the equations developed by Moore et al. (1999). This

adjustment accounted for the differences in total diet digestibility due to associative

effects from concentrate and forage (Appendix A).

To calculate the forage intake, the following assumptions were made:

1) Digestibility of forage and supplement were determined by IVDOM

2) Digestibility of forage was affected by the level of supplement intake, as

determined by the equations described by Moore et al. (1999)

Diurnal grazing time of calves was evaluated on a 28-d interval. Calves were

observed and actual grazing time recorded from 0700 to 1800 h. These observations were

performed on the first day of the grazing period in Paddock 2 of each grazing cycle.

The weight of the animals was recorded on a 28-d interval at 0900 h. The change in

unshrunk weight of the tester animals was used to calculate average daily gain (ADG).

Liveweight (LWG) per ha in each 28-d period was determined based on the ADG of the

testers multiplied by the number of calves within the pasture during that period and

adjusted to a hectare basis.

In 2004 only, blood was collected from the jugular vein on a 28-d interval. Samples

were collected into 9-mL, Na-heparinized syringes (Luer Monovette, LH, Sarstedt,

Inc., Newton, NC) and placed on ice. Blood was centrifuged (2000 xg rcf for 30 min) and

plasma was separated and frozen at -200C on the same day. Blood urea N (BUN) was

determined using a kit (Kit B-7551-120, Pointe Scientific, Inc., Detroit, MI) and read on

a plate reader at 620 nm.

Economic Analysis

The data and assumptions used in the economic analysis were:









Concentrate cost = $0.22 kg-1

Calf price = $2.2 kg-1

Days of grazing = 100 d

Where LW = calf liveweight per hectare, formulas used to calculate the results

were:

Concentrate cost ha-1 = LW*Concentrate level*Days of grazing*Concentrate cost

Income ha-1 = gain ha-'*calf price

The return was the difference between concentrate cost ha-1 and income ha-1.

The objective of the economic analysis was to compare the specific treatments used

in this study; it should not be extrapolated to a wide range of production systems.

Statistical Analysis

All responses were analyzed by fitting mixed effects models using the PROC

MIXED procedure of SAS (SAS Institute Inc., 1996). Replicate and its interactions were

considered random effects. Single degree of freedom orthogonal polynomial contrasts

were used to test concentrate effects. Treatments were considered different when P

<0.05. Interactions not discussed in the Results and Discussion section were not

significant (P>0.05). The means reported in this text are least squares means.

For pasture and pasture-animal interface variables, the model used was:

Yyk = t + A, + Tj + Pk+ (AT), + (AP),k+ (TP)jk + (ATP)jk + eUk

Where Yyk is the dependent variable

[t is the overall mean

T, is the concentrate effect (main plot)

A, is the year effect (sub-plot)









Pk is the period effect (sub-sub-plot)

(AT), is the year*concentrate interaction

(AP),k is the year*period interaction

(TP)jk is the concentrate*period interaction

(ATP)jk is the year*concentrate*period interaction

e,k is the error

Main plot (concentrate) error was replicate*concentrate; sub-plot (year) error was

replicate*year; sub-sub-plot (period) error was the residual.

For the animal variables, the model used was:

Yjkl = p + A, + T, + Pk+ Si + (AT)j + (AP),k+(PS)kl +(AS),1 + (TP)jk +(TS),1

+(ATP),k +(ATS),i +(TPS)jkl +(ATPS)kl + eJkl

Where Yykl is the dependent variable

t is the overall mean

T, is the concentrate effect (main plot)

A, is the year effect (sub-plot)

S, is the sex effect (sub-sub-plot)

Pk is the period effect (sub-sub-sub-plot)

(AT), is the year*concentrate interaction

(AP),k is the year*period interaction

(AS),i is the year*sex interaction

(TP)jk is the concentrate*period interaction

(TS),1 is the concentrate*sex interaction

(PS)kl is the concentrate*sex interaction









(ATP)jk is the year*concentrate* period interaction

(ATS),i is the year*concentrate*sex interaction

(TPS)jkl is the concentrate*period*sex interaction

(ATPS),kl is the year*concentrate*period*sex interaction

e,jk is the error

Main plot (concentrate) error was replicate*concentrate; sub-plot (year) error was

replicate*year; sub-sub-plot (sex) error was replicate*sex; sub-sub-sub-plot (period) error

was the residual.

Results and Discussion

Pasture Responses

There were no differences in pregraze herbage mass among the different levels of

concentrate (Table 3-2). There was a significant year*period interaction that occurred

because herbage mass was greater during January to March 2004 than in 2003, but there

was no year effect for April (Fig. 3-1). The greater rainfall from November 2002 to

March 2003 (Fig. 3-2) resulted in pastures with excessive soil moisture and likely

reduced the efficiency of utilization of N fertilizer causing lesser herbage mass during the

first 3 mo of the 2003 experimental period. The soils at the research site are characterized

by poor to somewhat poor drainage and are susceptible to flooding under conditions of

excessive rainfall. In contrast, limited rainfall in March and April 2004 likely resulted in

lesser mass during April than in January through March.

Macoon (1999) reported average herbage mass on rye-ryegrass pastures of 1.4 Mg

ha-1 for dairy cows at a SR of five cows ha-1. In that study, there was a significant effect

of month of sampling. Herbage mass was greater (1.3 Mg ha-)) in March than in January

(0.8 Mg ha-1). According to Rouquette et al. (1997), small grains grow better from late









December to mid-February and ryegrass has rapid forage growth during March to late

May, and as a result stocking rate often needs to be increased to efficiently use ryegrass

pastures. In the current study, this type of response was observed only in 2003.

Table 3-2. Pregraze herbage mass and herbage accumulation of rye-ryegrass pastures
grazed by early weaned calves supplemented with three levels of concentrate.
(BW=body weight)
Concentrate Herbage mass Herbage accumulation
-----------g kg-BW---- -----------Mg ha------- ---------kg ha- d----

10 1.6 31
15 1.5 27
20 1.7 35

Polynomial contrast NS' NS
P value > 0.18 > 0.44
SEM 0.1 3.0
SNS = not significant

Herbage accumulation was similar among treatments, with an average of 31 kg DM

ha-d-1 (Table 3-2). These rates are comparable to those reported by Assmann et al.

(2004). In their study in southern Brazil, ryegrass pastures were grazed by beef steers (3

AU ha-1), fertilized with 100 kg N ha-1, and herbage accumulation was 37 kg DM ha-d-.

For heavily-stocked (5 cows ha-1) rye-ryegrass pastures grazed by lactating dairy cows,

Macoon (1999) observed 22 kg DM ha-d-1.

There was a significant year*period interaction for herbage accumulation (Fig. 3-

3). This interaction occurred because herbage accumulation in April 2003 was much

greater than in 2004, while in other months there were no differences. Excessive rainfall

during November and December 2002 (220 mm vs. 30-yr average of 133 mm) resulted in

standing water in some pastures and decreased herbage accumulation during the first 2

mo after planting. In April and May 2003, soils retained sufficient moisture despite fewer









rainfall events to provide a better growth environment for the plants. In contrast, the

decreased herbage accumulation in April 2004 reflects lower than normal rainfall in

March (51 vs. 30-yr average of 93 mm) and April (26 vs. 30-yr average of 75 mm) of that

year (Fig. 3-2).

Table 3-3. Herbage crude protein (CP) and in vitro digestible organic matter (IVDOM)
concentrations of rye-ryegrass pastures grazed by early weaned calves
supplemented with three levels of concentrate. (BW=body weight)
Concentrate CP IVDOM
----------g kgBW----------- ----------g kgDM---------- ----------g kgOM-----------

10 273 841
15 282 837
20 283 838

Polynomial contrast NS' NS
P value >0.13 >0.51
SEM 4.0 3.0
SNS = not significant

There was no concentrate level effect on herbage CP and IVDOM (Table 3-3). The

CP concentrations were greater in 2004 (302 g kg-1) than 2003 (257 g kg-1) in all periods

likely because the very high rainfall during parts of the 2003 season. An anaerobic root

environment caused by excess rain can decrease absorption of N, resulting in lesser N

concentration in the plants (Salisbury and Ross, 1991). Also, heavy rain likely leached

some N out of the root zone.






53


OYear 2003 *Year 2004


10
S2
'~J








Jan


eb

Feb


b


Mar
Months


Fig. 3-1. Year*period interaction effect on pregraze herbage mass of rye-ryegrass
pastures grazed by early weaned calves supplemented with different levels of
concentrate. Year effect within period; **, P< 0.01; NS= not significant.
Period effect within year; means are different if followed by different letters
(P< 0.05).


o 2002/03
* 2003/04


rI^KL


Oct Nov. Dec. Jan.


7E7C


Feb. Mar. Apr. May


Months


Fig. 3-2. Rainfall distribution from October to May 2002/03 and 2003/04.


E 150

S100

50

0


25










|DYear 2003 *Year 2004
90

80

S70
O 60

50 NS

40
S30 NS NS

20

b b b a

Jan. Feb. Mar. Apr.
Months



Fig. 3-3. Year*period interaction effect on average herbage accumulation of rye-ryegrass
pastures grazed by early weaned calves supplemented with different levels of
concentrate. Year effect within period; **, P< 0.01; NS= not significant.
Period effect within year; means are different if followed by different letters
(P< 0.05).

There was a year*period interaction for CP concentration (Fig. 3-4). The herbage

CP concentration was least in January compared to other months because of maturity.

There was greater length of growth period of this forage herbage (-80 d from planting)

relative to regrowth grazing (21 d). Additionally, the lowest CP observed in this study

was in January 2003 following very heavy November and December rains. The greater

CP concentrations in February and March are associated with the dates of N fertilization

on the pastures (early February and early March).

There also was year*period interaction for IVDOM (Fig. 3-5). There was a

significant year effect on IVDOM concentration in January only. The IVDOM was not

affected by climatic conditions to the same degree as herbage mass, accumulation, and

CP concentrations were. The IVDOM decreased in both years, from January to April,

primarily because of the presence of reproductive tillers after February. Highest IVDOM,









unlike CP, occurred in January. Other studies (Chapter 5) have shown cool-season grass

IVDOM to be relatively insensitive to length of regrowth period during winter.

The CP and IVDOM concentrations were similar to the average CP (285 g kg-1)

and IVDOM (756 g kg-1) values reported by Arthington and Kalmbacher (2002) for

ryegrass pastures grazed by early weaned calves. Redfearn et al. (2002) showed CP and

IVDMD concentrations to be 232 and 846 g kg-1, respectively, for ryegrass pastures

grazed by beef steers.

Fractionation of CP showed ruminal-degradable protein concentration to be 730 g

kg-1 CP and ruminal-undegradable protein concentration to be 270 g kg-1 CP (data not

shown). These very high concentrations of ruminal degradable protein have the potential

to result in significant loss ofN in ruminants. Excessive ruminal-degradable protein

consumed by the animal is absorbed by the rumen epithelium and excreted as urea in the

urine. The NRC (1996) reported lower values for the ruminal degradable protein of

ryegrass hay (650 g kg-1). Using the in situ methodology, Michalet-Doureu and Ould-Bah

(1992) observed similar ruminal degradable protein concentration of 740 g kg-1 on

ryegrass hay.

Herbage allowance was similar among the treatments (Table 3-4). This was

achieved by increasing SR on high concentrate treatment pastures to account for

substitution of concentrate for forage. The average herbage allowance through the

experimental period was greater in 2004 (0.8) than in 2003 (0.4) due to superior fall

growth preceding the initiation of grazing in the second year.









Table 3-4. Herbage allowance and SR of rye-ryegrass pastures grazed by early weaned
calves supplemented with three levels of concentrate. (BW=body weight)
Concentrate Herbage allowance SRT
-----------g kg-BW-------- ----- kg DM kgBW------- -----------AU ha-------

10 0.62 5.5
15 0.59 5.9
20 0.66 6.5

Polynomial contrast NSW L
P value > 0.38 < 0.01
SEM 0.02 0.13
t AU = 500 kg BW.75

NS = not significant, L = linear

Phillips (1989) reported that to prevent a decline in individual dairy cow

performance, herbage allowance should allow for dry matter intake of at least 40 g OM

kg-1 BW d-1. McCartor and Rouquette (1977) studied the effect of different levels of

herbage allowance on performance of beef calves grazing pearl millet [Pennisetum

glaucum (L.) R. Br.] and concluded that animal performance declines as herbage

allowance levels fall below 1 kg DM kg1 BW. According to Le Du et al. (1979), the

maximum intake of cool-season grasses occurs when herbage allowance is at least twice

the herbage intake. The average herbage allowance observed in this study was always

more than twice the early weaned calves' forage intake.

There was no correlation (r2 = 0.03) between herbage allowance and ADG (Table

3-5) of the calves implying that factors other than herbage allowance were influencing

animal performance.








57



D Year 2003 0 Year 2004


250


. 200

S,,


Jan.


Feb.


Months Mar.
Months


Apr.


Fig. 3-4. Year*period interaction effect on crude protein (CP) concentration of rye-
ryegrass pastures grazed by early weaned calves supplemented with different
levels of concentrate. Year effect within period; **, P< 0.01; *, P< 0.05; NS=
not significant. Period effect within year; means are different if followed by
different letters (P< 0.05).


O Year 2003 Year 2004
900
**-


850




S800




750


700


NS


Jan


Feb


NS

-I-i


Mar


NS

-'7_


Apr


Months


Fig. 3-5. Year*period interaction effect on in vitro digestible organic matter (IVDOM)

concentration of rye-ryegrass pastures grazed by early weaned calves
supplemented with different levels of concentrate. Year effect within period;
**, P< 0.01; NS= not significant. Period effect within year; means are
different if followed by different letters (P< 0.05).


**


***










There was a significant year*period interaction on herbage allowance (Fig. 3-6).

There was no difference in herbage allowance during January and February 2003.

Herbage allowance was less in March than January, but there was no difference between

February and March. During April, herbage allowance was greater than in the other

months. This was a result of the greater herbage accumulation (Fig. 3-1) during that

month. In 2004, herbage allowance was greatest in January and decreased through April.

High herbage allowance in January 2004 was due to excellent forage growth during fall

2003 and conservative stocking decisions to start the experiment in 2004.

O Year 2003 Year 2004
14

12




08

06

04
b bc C a
02

0
Jan. Feb. Months Mar. Apr.

Fig. 3-6. Year*period interaction effect on herbage allowance of rye-ryegrass pastures
grazed by early weaned calves supplemented with different levels of
concentrate. Year effect within period; **, P< 0.01; NS= not significant.
Period effect within year; means are different if followed by different letters
(P< 0.05).

Horn et al. (1995) conducted a 3-yr experiment to evaluate the influence of energy

supplements on performance of stocker cattle grazing wheat pasture. They concluded that

energy supplements, such as maize and soybean [Glycine max (L.) Merrill] hulls, when

fed at 7.5 g kg-1 BW allowed a 33% increase in stocking rate compared to









unsupplemented calves. In this study, there was 18% increase in SR when the supplement

level was increased from 10 to 20 g kg- BW (Table 3-5).

Animal Response Variables

There was a linear increase in ADG and LWG as supplementation level increased

(Table 3-5). The average daily gain reported for calves supplemented at the 10 g kg1 BW

concentrate level in this study was the same as that reported by Arthington and

Kalmbacher (2003) for calves grazing ryegrass in south Florida. There were no effects

due to sex or year on ADG, however, there was a year*period interaction (Fig. 3-7). The

decrease in ADG during February and March 2003 was associated with an infestation of

endoparasites that was not detected by visual examination. After the depression in ADG

was observed in March, feces samples were collected, examined, and a high incidence of

parasites detected. Calves were individually treated with 6 ml of Ivermectin (10 g kg-1)

and a large increase in ADG was observed during April.

The ADG and LWG of calves that did not receive concentrate supplementation

were 0.3 kg d-1 and 650 kg ha-1, respectively. These values were well below the ADG and

gain per ha of calves supplemented with 10 g kg-1 BW in concentrate, 0.74 kg d-1 and 950

kg ha-l, respectively. Although the forage had nutritive value similar to the concentrate,

the low DM concentration (150-180 g kg-1) of the forage may be limited the total OMI of

the animals resulting in decreased performance.

The linear increase in LWG was a function of the linear increase in both SR and

ADG as concentrate level increased. Horn et al. (1995) observed greater LWG on steers

grazing wheat pastures supplemented with concentrate at 7.5 g kg-1 BW (153 kg ha-1)

than for the unsupplemented calves (103 kg ha-1). Restle et al. (2000) studied beef heifers

grazing oat-ryegrass pastures fertilized with 200 kg N ha-l under continuous stocking.









Average daily gain and LWG were 0.58 kg d-1 and 429 kg ha-1, respectively, less than the

values in our study. The results of the current study agree with the literature on the effects

of supplementation on beef calves grazing cool-season grass pastures, i.e., energy

supplementation leads to substitution of forage by concentrate, allowing higher stocking

rates and gains per unit land area.

There was no difference in total OMI among treatments; however, there was a

linear decrease in forage OMI with increasing levels of supplement (Table 3-6). Total

OMI was similar to that reported by Paisley et al. (1998) for early weaned calves grazing

wheat pastures (28 g kg-1 BW). In general, forage intake is decreased by feeding

concentrate when forage quality is high, other nutrients are in balance with energy, and

concentrate is fed in large amounts (Moore, 1994). Carey et al. (1993) supplemented beef

steers grazing tall fescue (Festuca arundinacea Schreb.) pastures with different sources

of energy. Forage intake was less for steers supplemented with energy than for control

steers, but total intake (forage + supplement) did not differ among treatments. Cravey

(1993) reported substitution ratios of 1:0.93 (forage:concentrate) when steers grazing

wheat pastures were fed energy concentrate at 10 g kg-1 BW. The substitution rate of

concentrate OM (0.8, 1.2, and 1.6 for 10, 15, and 20 g kg-1 treatment respectively) for

forage OM in the current study was 1:1 and consistent across treatments, confirming

observations in the literature of substitution of forage by concentrate intake at similar

proportions by calves receiving high amounts of energy supplement and grazing high

nutritive value cool-season pastures.







61


Table 3-5. Average daily gain (ADG) and liveweight gain (LWG) of early weaned calves
grazing rye-ryegrass pastures and supplemented with three levels of
concentrate. (BW=body weight)
Concentrate ADG LWG1

-----------g kgBW--------- ---------- kg d-1----------- -----------kg ha------------

10 0.74 950

15 0.81 1080

20 0.89 1320

Polynomial contrast L L

P value < 0.01 < 0.01

SEM 0.03 42

SGain per ha during a 106-d period.


L = linear


O Year 2003 Year 2004


08


o06


04


02


0


NS


T


Jan.


Feb.


Months


Fig. 3-7. Year*period interaction effect on average daily gain of early weaned calves
grazing rye-ryegrass pastures and supplemented with different levels of
concentrate. Year effect within period; **, P< 0.01; NS= not significant.
Period effect within year; means are different if followed by different letters
(P< 0.05).

The greater ADG at higher concentrate levels despite similar total intake may be a

function of the different sources of nutrients provided by the concentrate that resulted in









better N and energy synchronization in the rumen and increased microbial protein

production. In addition concentrate had more rumen-undegradable protein than the

forage, potentially leading to more amino acids being absorbed post rumen and

decreasing the levels of ammonia absorbed by the rumen epithelium. Absorption and

excretion of excess ammonia is a process that spends energy that otherwise could be used

to improve animal performance. Calves that received greater concentrate rates spent less

time grazing during daylight (Table 3-7) likely reducing their energy requirement for

maintenance. This could also be a factor contributing to greater ADG for calves receiving

greater levels of concentrate, despite total OMI being the same across treatments..

Table 3-6. Total organic matter intake (OMI) and forage OMI of rye-ryegrass pastures
grazed by early weaned calves supplemented with three levels of concentrate.
(BW=body weight)
Concentrate Total OMIt Forage OMI
---------g kg-BW-------- -------------- (% BW)---------

10 2.6 1.8
15 2.4 1.3
20 2.7 1.1

Polynomial contrast NSW L
P value > 0.68 < 0.01
SEM 0.01 0.01

STotal OMI intake data adjusted according to Moore et al. (1999).

NS = not significant, L = linear

Grazing time differed among the calves receiving different levels of concentrate.

Calves supplemented with 10 g kg-1 BW concentrate spent more time grazing during

daylight hours than calves receiving 15 and 20 g kg-1 BW (Table 3-7). There was no sex

effect on grazing time.









In research with yearling beef steers grazing ryegrass, Adams (1985) reported that

although there was an 11% decrease in forage intake by supplemented calves compared

with control calves, total daily grazing time was not altered. Macoon (1999) observed

that dairy cows supplemented with high levels of concentrate spent less time grazing (135

min) than cows that received low rates of concentrate (180 min). Cowan et al. (1977)

observed decreased grazing time of 23 min d1 for each additional kg of concentrate fed to

cows grazing grass-legume pastures. Increased grazing time in the current study suggests

that calves supplemented with lower rates of concentrate grazed longer during the day in

an attempt to meet their nutritional requirements. According to Macoon (1999), lactating

cows receiving low levels of concentrate tended to increase the night-time grazing as

well.

Table 3-7. Grazing time during daylight hours of early weaned calves grazing rye-
ryegrass pastures and supplemented with different levels of concentrate.
Concentrate Grazing time
-----------g kgBW----------- ----------------------------min----- ----------

10 284
15 230
20 234

Polynomial contrast Q
P value < 0.08
SEM 17

SQ = quadratic.

There was no effect of three levels of concentrate supplementation on calf BUN

concentrations (Table 3-8). Blood urea N levels between 11 and 15 mg dL-1 were

associated with maximum rates of gain for growing steers (Byers and Moxon, 1980), thus

it is likely that the calves had an adequate protein supply during the entire experimental









period. Increased solubility and degradability of dietary protein can lead to increased

ruminal ammonia concentrations resulting in increased BUN concentrations (Hammond

et al., 1994). Protein in intensively fertilized cool-season grasses is highly degradable and

thus easily fermented to volatile fatty acids and ammonia in the reticulo-rumen.

Ammonia that is not captured is absorbed and excreted in the urine (Van Vuuren et al.,

1991). It was expected that calves receiving greater amounts of concentrate would have a

lesser BUN concentration because the concentrate had lower CP and ruminal degradable

protein concentrations than the rye-ryegrass pastures. There was a numerical suggestion

of lesser blood urea N for calves receiving the 20 g kg-1 BW concentrate compared to the

15 g kg-1 level, however, no statistical difference was observed.

Table 3-8. Blood urea nitrogen (BUN) concentrations of early weaned calves grazing rye-
ryegrass pastures supplemented with three levels of concentrate. (BW=body
weight)
Concentrate BUN
---------- g kg- BW---------------------------------mg dL---------------------

10 14.1
15 15.1
20 13.0

Polynomial contrast NS'
P value > 0.26
SEM 1.0
SNS = not significant

According to the NRC (1996), the ruminal-degradable protein requirement for 120

kg calves is 300 g d-1. Based on the estimations of forage and concentrate intake for

calves receiving the 10 g kg-1 BW supplement and the data for CP fractionation, ryegrass

provided 380 and concentrate 100 g of ruminal degradable protein d-1. Therefore, the

estimated ruminal degradable protein consumed was 60% greater than the requirement.









The excessive rumen degradable protein consumed by the calves was not expressed in

increased BUN concentrations that were in the range considered adequate for growing

calves.

Economic Analysis

There was a linear increase in supplement cost and income with increasing

supplement levels; however, there was no difference in return among the treatments

(Table 3-9). Provided that there are no economic incentives to producing heavier calves,

these data do not justify use of supplement levels above 10 g kg1 BW.

Table 3-9. Economic analysis of three concentrate levels on early weaned calves grazing
rye-ryegrass pastures.
Concentrate Concentrate cost Income Return
-1 -1
--------g kg-1 BW------ -------------------------- $ ha-- -------------------

10 600 2100 1500

15 970 2370 1400

20 1430 2900 1470

Polynomial contrastT L L NS
P value < 0.01 < 0.01 > 0.86

SEM 25 91 92

L = linear; NS = not significant

Summary and Conclusions

Herbage mass, accumulation, allowance, and nutritive value of rye-ryegrass

pastures did not differ among levels of concentrate. Seasonal variation in forage

responses were the result of the climatic conditions during the trial, especially the very

high rainfall that occurred during the 2002/03 season. It resulted in lower herbage mass

and CP during that year.









Total OMI did not vary among the three supplementation levels, but forage OMI

decreased linearly as supplement level increased. Concentrate substituted for forage

intake in a 1:1 ratio. To enable greater forage intake, calves receiving the 10 g kg-1 BW

concentrate level spent more time grazing during daylight hours than calves receiving the

15 and 20 g kg-1 BW treatments.

Average daily gain, SR, and LWG increased linearly with increasing levels of

concentrate. Increased ADG at the same level of intake implies better synchronization of

energy and N in the rumen, and possibly an effect associated with the greater amounts of

ruminal undegradable protein provided by the concentrate. Stocking rate increased

because of the substitution of concentrate for forage in the 15 and 20 g kg-1 BW

treatments.

There was a trend for BUN concentration to be lower for the 20 than 15 g kg-1 BW

treatment, likely the result of the lower CP and ruminal degradable protein concentration

in the concentrate compared to the forage. The BUN concentrations of all treatments

were in the range considered adequate for growing cattle.

The economic analysis implies that there is no economic benefit of increasing the

supplementation levels above 10 g kg-1 BW, unless a greater price is paid for heavier

weight calves. In practice, the optimum supplementation level will be dependent upon

forage availability, cost of the concentrate, and calf price. Given the pasture conditions,

and the concentrate and calf pricing in this study, the 10 g kg-1 BW supplement level is

recommended. Future studies should evaluate lesser levels, e.g., 5 g kg-1, to determine if

more economical gains can be achieved with less supplement.














CHAPTER 4
SUPPLEMENTATION EFFECTS ON FORAGE CHARACTERISTICS AND
PERFORMANCE OF EARLY WEANED CALVES GRAZING TIFTON 85
BERMUDAGRASS PASTURES

Introduction

There are 39 million ha of perennial pastures grown in the eastern USA and almost

30 million ha are located in the South. These pastures are mainly used for beef cows and

calves and stockering of weaned calves (Hoveland, 1992). Among several species found

in the southeastern USA, bermudagrass [Cynodon dactylon (L.) Pers.] is one of the most

important with 10 to 12 million ha of planted for livestock grazing and hay.

Johnson et al. (2001b) compared different species of warm-season grasses in

Florida and showed that bermudagrass produced more forage DM (1540 43 kg ha-1

cutting-1) than stargrass (Cynodon nlemfuensis Vanderyst) (1400 43 kg ha-1 cutting-) or

bahiagrass (Paspalum notatum Flugge) (1300 43 kg ha-1 cutting-'). Moreover,

bermudagrass was more digestible (575 4 g kg-1) than stargrass (546 4 g kg-1) and

bahiagrass (519 4 g kg-1). In 1993, 'Tifton 85', a hybrid between a South African

bermudagrass and 'Tifton 68' stargrass was released. Burton et al. (1993) described it as

taller, with larger culms, broader leaves, and darker color than other bermudagrass

hybrids. Compared with 'Coastal', Tifton 85 yielded 26% more dry matter, was 110 g

kg-1 more digestible, and more succulent at harvest.

Warm-season grasses have relatively larger proportions of cell-wall material. Cell

wall is potentially digestible, but because of the chemical composition and anatomical

structure of warm-season grasses the rate of degradation and fermentation is relatively









slow (Coleman et al., 2004). Although forage nutritive value of C4 grasses is generally

inferior to that of C3 grasses, animal production can be high on C4 grasses because of

their high DM production potential. Pedreira et al. (1998) studied Tifton 85 bermudagrass

in a 3-yr grazing trial. The pastures were continuously stocked with a fertilization

program of 210 kg N ha-1 yr-1. Tifton 85 pastures supported six heifers ha-1, resulting in

live weight gain (LWG) 648 kg ha-lduring spring-summer in North-Central Florida.

The use of supplementary feed is a management practice to overcome the

nutritional shortfalls of the forage component of the diet. When energy and protein

requirements are high, e.g., for lactation, pregnancy, and growth, part of the roughage

component may need to be replaced by feeding of concentrates (Fontaneli, 1999).

Rouquette et al. (2003) reported that stocker steers gained 1.03 kg d-1 on Tifton 85

compared to 0.75 kg d-1 on Coastal bermudagrass without supplementation. With

supplementation of 1.4 kg d-1 of a 3:1 maize (Zea mays L.):soybean [Glycine max (L.)

Merr.] meal concentrate, steers grazing Tifton 85 gained 1.25 kg d1.

According to Galloway et al. (1992), moderate dietary levels of supplement (200-

300 g kg-1 of the diet DM) can improve nutrient intake and performance by cattle

consuming bermudagrass. At greater amounts, nutrient digestion, intake, or both of the

forage portion of the diet can be affected negatively. Wheeler et al. (2002) tested the

effects of increasing supplement protein concentration on performance and forage intake

of beef steers consuming bermudagrass forage. Treatments were no supplement or daily

equivalents of 0.2, 0.4, and 0.6 g of supplemental protein kg-1 of body weight (BW).

Forage intake increased 16% and organic matter (OM) intake increased 30% in

supplemented compared to unsupplemented steers. Diet OM digestibility increased 145 g









kg-1 and total digestible OM intake increased 490 g kg-1 in supplemented compared to

unsupplemented steers.

The success of the Florida cow-calf enterprise depends upon maintaining high

calving percentage. First-calf heifers and other young cows may experience low

conception rates due to stress of lactation. Early weaning of calves of young cows is one

approach to reduce cow nutrient requirement, increase body condition, and increase the

likelihood of rebreeding. Management practices for raising the early weaned (EW) calf

have not been widely studied. In particular there is little information to guide

supplementation programs for EW calves grazing C4 grass pastures. In a study done in

south Brazil, Muehlmann et al. (1997) reported average daily gains (ADG) of 0.2 kg d-1

for EW calves grazing a bermudagrass-native warm-season grasses mixture. Vendramini

et al. (2003) observed that supplementing EW calves with concentrate at 10 g kg-1 BWof

body weight resulted in gains of 0.59 and 0.44 kg d-1 on stargrass and atrapaspalum

(Paspalum atratum Swallen), respectively. Additional information is needed regarding

the amount of supplement required by EW calves grazing C4 grass pastures. The

objective of this study was to evaluate the effect of different levels of concentrate on

ADG, stocking rate (SR), LWG, and forage intake of EW calves grazing Tifton 85

bermudagrass pastures in North-Central Florida.

Material and Methods

The research site was the Florida Beef Research Unit, 18 km northeast of

Gainesville, FL (300 N). Dates for the trial during the 2 yr were 14 May through 13 Aug.

2003 (86 d) and 18 May through 10 Aug. 2004 (86 d). Monthly rainfall and temperature

data during the trial are presented in Table 1 (Appendix A).









The soils at the research site are Adamsville fine sand (Aquic Quartzipamments,

hyperthermic, uncoated) and Sparr fine sand (Grossarenic Paleudult, hyperthermic, loamy

siliceous). These soils are moderately well drained with rapid permeability. Prior to

initiation of summer grazing at the site, mean soil pH (1:2 soil:deionized H20 ratio) was

5.5, and Mehlich-I (0.05 MHC1 + 0.0125 MH2SO4) extractable P, K, Mg, and Ca in the

Apl horizon (0- to 15-cm depth) were 44, 17, 40, 328 mg kg-1, respectively.

Bermudagrass pastures were established in August 2002 and fertilized with 40 kg

ha- of N, 17 kg ha- of P, and 66 kg ha- ofK 1 mo after establishment. In April 2003 and

2004, the pastures received 40 kg N ha-l, 17 kg P ha-l, and 66 kg K ha-1. An additional 80

kg N ha- was split in two applications of 40 kg ha- in early June and early July of both

years.

Treatments were three levels of a commercial pelleted concentrate, 10, 15, and 20 g

kg- of the calves body weight (BW), offered daily. The concentrate composition was

described in Table 3.2 (Chapter 3). Each treatment was replicated three times in a

completely randomized design so there were a total of nine experimental units in the

study. In 2004, for observation purposes, two experimental units were added to the study

in which the EWC did not receive concentrate supplementation.

The EWC were described in Chapter 3. At the start of this trial, they were 200-d

old with an average liveweight of 190 kg. Calves grazed rye (Secale cereale L.)-annual

ryegrass (Lolium multiflorum Lam.) pastures from January to April. Calves received the

same supplement treatment in summer as they did during the winter-spring period. At the

initiation of grazing of the Tifton 85 pastures, calves within a supplement level were

randomly allocated to the three replicates of that treatment. They were vaccinated with









Ultrabac 8 and Bovashield 4 (Pfizer Animal Health New York, NY) and dewormed with

ivermectin (Ivomec, Merck & Company, Rahway, New Jersey, USA) 10 g kg-

concentration on 23 July 2003 and 14 July 2004.

Pasture size was 0.15 ha subdivided in three paddocks for rotational stocking. The

grazing period was 7 d and the rest period was 14 d. Two EW calves (1 steer and 1

heifer) were assigned as testers to each pasture and "put and take" EW calves were used

to maintain herbage allowance at -1 kg forage DM kg-1 of calf BW.

Pasture Sampling

In each grazing cycle (21 d), herbage mass was determined on Paddock 2 using a

double sampling technique. The procedure of herbage sampling was similar to the one

described in Chapter 3. A total of 40, double-samples were taken every 3 wk to calibrate

the disk meter. This greater number than used during the winter was necessary because

C4 grasses usually have a greater heterogeneity of herbage bulk density through the

different heights of the canopy (Sollenberger and Bums, 2001). Thus, separated pre- and

post-graze calibration equations resulted in more precision of the height/weight

relationship than combining them into one. The regression equations and r2 of the pre-

and post-graze double sampling technique are shown in Table 1 (Appendix B).

The methodology used to determine herbage allowance, stocking rate, and forage

nutritive value were the same as those described in Chapter 3.

Animal Response Variables

Total organic matter intake (OMI) and forage OMI were estimated only in 2003.

The controlled-release devices used in the trial were designed for sheep or small calves.

When used by 200-kg calves, most of the calves regurgitated the device before the fecal

collection period. The data are presented only for 2003 and reflect results from only 8 out









of 18 calves dosed with the controlled-release device. The total Cr output calculated from

total fecal collection from three calves is presented in Table 2 (Appendix B). The Cr

analyses procedure and total and forage OMI determinations were described in Chapter 3.

Grazing time of EW calves during daylight hours was evaluated every 21 d. Calves

were observed and time spent grazing was recorded from 0700 to 1900 h. These

observations were performed on the first day of the grazing period in Paddock 2 in each

grazing cycle.

The weight of the animals was recorded on a 21-d interval at 0900 h. The change in

unshrunk weight of the tester animals was used to calculate ADG. The LWG in each 21-d

period was determined based on the ADG of the testers multiplied by the number of

calves within the pastures during that period and adjusted to a hectare basis. Blood urea N

(BUN) sampling and determination were conducted as described in Chapter 3.

Economic Analysis

The assumptions used in the economic analysis were:

* Concentrate cost = $0.22 kg-1
* Calf price = $2.20 kg-1
* Live weight (LW) = SR*500 kg
* Days of grazing = 90 d

The formulas used to calculate the results were described in Chapter 3.

Statistical Analysis

The models, procedure, and software used for statistical analyses were described in

Chapter 3.









Results and Discussion

Pasture Responses

There was no difference in pregraze herbage mass and herbage accumulation

among the treatments (Table 4-1).

Table 4-1. Pregraze herbage mass and herbage accumulation of Tifton 85 pastures grazed
by early weaned calves supplemented with three levels of concentrate.
(BW=body weight)
Concentrate Herbage mass Herbage accumulation
-----------g kg-1 BW------- ---------Mg ha-------- --------kg had---

10 4.9 101
15 4.8 95
20 4.7 84

Polynomial contrast NS' NS
P value 0.20 0.11
SEM 0.1 15

SNS = not significant

Using continuous stocking, Pedreira et al. (1998) reported greater yields of Tifton

85 than Florakirk bermudagrass early in the grazing season. Herbage accumulation was

73 kg DM ha-ld-for Tifton 85 vs. 44 kg DM ha-ld- for Florakirk. Under continuous

stocking, average herbage mass during a 3-yr study with Tifton 85 grazed by beef steers

in Georgia was 2.8 Mg ha-l, and ADG decreased when herbage mass was below 2.5 Mg

ha-1 (Hill et al., 1993). Greater average herbage mass was maintained in our study to

provide opportunity for the calves to select forage that was high in nutritive value and

maximize animal performance.







74


OYear 2003 MYear2004
70

** NS
60




40




20- b C ab a

10-

00-
May Jun. Jul. Aug.
Months

Fig. 4-1. Year*period interaction effect on pregraze herbage mass of Tifton 85 pastures
grazed by early weaned calves supplemented with different levels of
concentrate. Year effect within period; **, P< 0.01; NS= not significant.
Period effect within year; means are different if followed by different letters
(P< 0.05).

There was a significant year period interaction effect on herbage mass (Fig. 4-1).

The lesser herbage mass present in the first 2 mo of 2004 can be attributed to very low

rainfall that occurred during May (15 mm vs. the 30-yr average of 106 mm).

Herbage accumulation also varied throughout the grazing season ranging from 45

to 121 kg ha-1 d-1 in 2003 and from 51 to 133 kg ha- d-1 in 2004 (Fig. 4-2). According to

Dubeux et al. (2003), rotationally stocked pastures of bahiagrass had greater herbage

accumulation than continuously stocked pastures. The use of a rotational stocking may be

one of the factors responsible for greater herbage accumulation in this study compared to

those of Hill et al. (1993) and Pedreira et al. (1998) who stocked their pastures

continuously.










DYear 2003 *Year 2004
160

140

120



80 -
aE
U 60 -

40

20 C b a


Jun. Jul. Aug
Months

Fig. 4-2. Year*period interaction effect on average herbage accumulation of Tifton 85
pastures grazed by early weaned calves supplemented with different levels of
concentrate. Year effect within period; **, P< 0.01; NS= not significant.
Period effect within year; means are different if followed by different letters
(P< 0.05).

Herbage CP and IVDOM concentrations were not affected by treatment and

averaged 184 g kg-1 and 624 g kg-1, respectively (Table 4-2). These concentrations are

greater than those reported by Pedreira et al. (1998) (120 g kg-1) and Hill et al. (1993)

(140 g kg-1) for Tifton 85 pastures stocked continuously. Fike et al. (2002) used rotational

stocking with similar resting period (21 d) and N fertilization program (45 kg N ha-1 on

21 May, 8 June, and 7 August) on Tifton 85 pastures. They reported CP and IVDOM

values (140 and 600 g kg-1, respectively) lower than those observed in this study.

Masticate samples from Tifton 85 pastures had CP and IVDMD concentrations of 180

and 650 g kg-1 (Hill et al., 1998). With a 35-d regrowth interval, Mislevy and Martin

(1998) reported CP of 170 g kg-1 DM for hand-plucked samples of stargrass pastures

grazed by beef steers.









Table 4-2. Crude protein (CP) and in vitro digestible organic matter (IVDOM)
concentration of Tifton 85 pastures grazed by early weaned calves
supplemented with three levels of concentrate. (BW=body weight)
Concentrate CP IVDOM
-----------g kg-1 BW---- -----------g kg-DM--------- -----------g kgOM----------

10 182 626
15 181 610
20 189 636

Polynomial contrast NS' NS
P value 0.49 0.44
SEM 6 8
SNS = not significant

There was a year*period interaction for CP concentration (Fig. 4-3). In both years,

the lower CP during the first month of grazing was attributed to a longer growth period

preceding the first grazing cycle than subsequent cycles. At initiation of grazing in May,

the herbage present reflected growth that had occurred during the spring and not from the

most recent 14 d. Thus, hand-plucked samples included older herbage with more stem

material resulting in lesser CP and IVDOM concentrations. Pedreira (1995) reported

similar results from bermudagrasses pastures grazed by beef heifers. Herbage CP was 70

g kg-1 in June while the average for June through August was 125 g kg-1. Similar results

were presented by Hill et al. (1993), with CP concentrations of 114 (May) and 156 g kg-1

(September) in the first and last month of the grazing study with Tifton 85.










O Year 2003 Year 2004
300


NS

200


150 NS


100


50 C a a b



May Jun. Jul. Aug.
Months


Fig. 4-3. Year*period interaction effect on crude protein (CP) concentration of Tifton 85
pastures grazed by early weaned calves supplemented with different levels of
concentrate. Year effect within period; **, P< 0.01; NS= not significant.
Period effect within year; means are different if followed by different letters
(P< 0.05).

There was no year*period interaction effect on herbage IVDOM, but there were

year and period (Fig. 4-4) effects. Mean herbage IVDOM was greater in 2004 (640 g kg-

1) than in 2003 (580 g kg-1). The herbage IVDOM period response was similar to that of

CP concentration, with the lowest values occurring during the first month of grazing

because of the longer regrowth period of the forage. Herbage IVDOM was greater in

June and decreased in July and August.

The estimated rumen-degradable and rumen-undegradable protein concentrations

were 550 and 450 g kg-1 CP, respectively (data not shown). Similar values for

bermudagrass rumen-degradable protein (570 g kg-1) were found by Basurto et al. (2001).

Mathis et al. (2001) measured bermudagrass ruminal degradable protein from seven

different locations using the enzymatic methodology proposed by Roe et al. (1991). The

average across locations was 610 g kg-1 CP.








78




750
700
650
600
550

450
400
350
300
S250 d a c b
200
150
100
50

May Jun. Jul. Aug.
Months



Fig. 4-4. Period effect on IVDOM concentration of Tifton 85 herbage grazed by early
weaned calves supplemented with different levels of concentrate. Means are
different if followed by different letters (P< 0.05).


There was a linear decrease in average herbage allowance with increased


supplementation levels, although the magnitude of the change was small (Table 4-3). Put


and take animals were used to maintain similar herbage allowance among treatments.


Even though herbage mass was not different among treatments, the greater SR for the 20


g kg-1 BW treatment caused herbage allowance to be slightly reduced. There was no


period or period year interaction on herbage allowance, however, a year effect was


observed with averages across treatments of 1.0 and 0.8 kg DM kg1 BW for 2003 and


2004, respectively. Herbage allowance of at least 1.0 kg DM kg-1 BW was associated


with highest gains of unsupplemented yearling beef heifers on continuously-stocked


Tifton 85 bermudagrass (Pedreira, 1995). Although the herbage allowance in this study


was below 1 kg DM kg-1 BW in 2004, it is important to consider that calves were


supplemented with at least 10 g kg-1BW concentrate. This amount of concentrate









accounts for 33% of the daily energy requirement of these animals, likely resulting in

lower forage allowance requirements.

Table 4-3. Herbage allowance and stocking rate of Tifton 85 pastures grazed by early
weaned calves supplemented with three levels of concentrate. (BW=body
weight)
Concentrate Herbage allowance Stocking rate
-----------g kg-1 BW------------kg DM kg BW ------- -----------AU ha----------

10 0.97 11.1

15 0.95 11.2

20 0.81 13.7

Polynomial contrast L0 L
P value <0.01 <0.01

SEM 0.02 0.4
SAU = 500 kg LW.75

tL=linear

Hernandez Garay et al. (2004) studied the effect of N fertilization and stocking rate

on weanling bulls grazing stargrass. Animal ADG decreased quadratically with

increasing stocking rate from 1.3 to 3.8 AU (500 kg LW) ha-1. The LWG was maximized

at stocking rates of 2.5 AU ha-1 for N fertilization rates of 112 and 224 kg ha-lover a

280-d grazing season. Pedreira (1995) verified a mean seasonal stocking rate of 5.9 AU

ha-1 on Tifton 85 pastures with maximum stocking rate of 9 AU ha-1 in July 1994.

The greater stocking rates in this study are the result of supplementation of the

calves. The substitution of forage by concentrate resulted in more forage available for

grazing, resulting in a linear increase in stocking rates with increased level of

concentrate.









Animal Response Variables

There was linear increase in ADG and LWG as level of supplement fed increased

(Table 4-4). The performance of early weaned calves grazing Tifton 85 and receiving 10

g kg-1 BW in concentrate was similar to the results presented by Vendramini et al. (2003)

for early weaned calves grazing stargrass (0.59 kg d-1) and receiving the same amount of

concentrate. When Harvey and Burns (1989) evaluated performance of EW calves

grazing pearl millet [Pennistum glaucum (L.) R. Br.] with ground ear maize fed ad

libitum, the ADG of the calves was 0.92 kg d1.

In the current study, average daily gain increased only 25% with an increase of

100% in amount of supplement fed. Because the level of gain observed is well below the

genetic potential for these calves, it is likely that energy and/or protein limited growth of

the calves. According to the NRC (1996) simulations calves grazing Tifton 85 with 10 g

kg-1 BW supplementation would gain 0.9 kg of liveweight d-1. The lesser ADG observed

in this study may be related to greater than expected negative associative effect of

concentrate on forage intake. In addition, high temperature and humidity during the

grazing season may have contributed to the relatively poor performance of the calves.

The ADG and LWG of calves that did not receive concentrate supplementation

were 0.33 kg d-1 and 950 kg ha-1, respectively. These values were inferior to the ADG

and gain per ha of calves supplemented with 10 g kg-1 BW in concentrate, 0.52 kg d-1 and

1080 kg ha-1, respectively.

There was a year*period interaction on calf ADG (Fig. 4-5). The greater ADG

observed during May 2003 may have resulted from differences in gut fill during the

transition of the calves from cool-season to warm-season pastures. In 2003, calves were

moved from rye-ryegrass directly to Tifton 85, and a greater amount of fill associated









with consumption of Tifton 85 relative to rye-ryegrass could explain the greater observed

gain in May. In 2004, the calves spent 2 wk grazing warm-season grasses before the

initiation of the warm-season study, decreasing the variation in fill effect between the

first two weight measurements. With the exception of May 2003, there was a tendency of

increasing ADG through the experimental period in both years. This may be related to

generally increasing CP of pastures throughout the experimental period.

Table 4-4. Average daily gain (ADG) and liveweight gain (LWG) ha-1 of early weaned
calves grazing Tifton 85 pastures supplemented with different levels of
concentrate. (BW=body weight)
Concentrate ADG LWG1
-----------g kg-1 BW ---------------- kg d-1------ ------------kg ha--------

10 0.52 1080
15 0.65 1450
20 0.65 1550

Polynomial contrast L L
P value 0.04 <0.01
SEM 0.02 85

SLWG during a period of 86 d.

L = linear

Using a fixed stocking rate, Rouquette et al. (2003) reported that supplementation

with 0.9 kg of a mixture of maize and soybean meal increased LWG of yearling heifers

on Tifton 85 pastures from 521 (unsupplemented control) to 615 kg ha-1. In the current

study, gain ha-1 increased 43% when supplementation level increased from 10 to 20 g

kg-1BW.

There was no difference in total and forage OMI (Table 4-5). Trends were

observed, especially for forage OMI, but missing data due to bolus regurgitation

weakened the statistical tests.







82


D Year2003 Year2004


** T


Jun. Jul. Aug.
Months


Fig. 4-5. Year*period interaction effect on average daily gain of early weaned calves
grazing Tifton 85 pastures and supplemented with different levels of
concentrate. Year effect within period; **, P< 0.01; NS= not significant.
Period effect within year; means are different if followed by different letters
(P< 0.05).

Table 4-5. Total organic matter intake (OMI) and forage OMI of Tifton 85 pastures
grazed by early weaned calves supplemented with three levels of concentrate.
Concentrate Total OMI Forage OMI

---------g kg-1 BW-------- --------------- (% BW)---------

10 3.2 2.3

15 2.8 1.6

20 2.5 1.2

Polynomial contrast NSW NS

P value 0.45 0.21

SEM 0.4 0.4

t Total organic matter intake data adjusted according to Moore et al. (1999).

;NS = not significant

Nelson et al. (1980) reported that intake of Coastal bermudagrass hay was 23.9 g

kg-1 BW for forage harvested at 4-wk regrowth. Hill et al. (1993) studied DM intake of


- 06


S04


02


May


08









freshly harvested Tifton 85 forage at different maturities. Intake varied from 18 to 20 g

kg-1 BW for 3- to 7-wk regrowth. When supplemented with maize at 10 g kg-1 BW, beef

steers reduced the bermudagrass hay intake 0.46 kg for each kg of maize fed.

Burns et al. (1991) compared intake of different species of warm-season grasses in

North Carolina. Bermudagrass (22.3 g kg-1 BW) was the lowest compared with

switchgrass (Panicum virgatum L.) and flaccidgrass (Pennisetumflaccidum Griseb.)

(30.9 g kg-1BW). The authors attributed the lower bermudagrass intake to low forage

digestibility. Wheeler et al. (2002) verified increased DM intake in beef steers grazing

stockpiled bermudagrass pastures when supplemented with concentrate of different CP

concentrations. The intake was 21 and 23 g kg-1 BW for steers supplemented with 16 and

40% CP supplement.

It is expected that calves grazing bermudagrass and consuming over 30 g kg-1 of

the diet as an energy supplement would have decreased forage OMI. The data collected

in this study support this trend. However, the problems experienced with the

methodology limit the conclusions that can be drawn.

Grazing time during daylight hours decreased linearly with increasing

supplementation levels (Table 4-6). These date corroborate the results of Macoon (1999)

who found that dairy cows supplemented with high levels of concentrate spent less time

grazing (135 min) than cows that received low rates of concentrate (180 min). There was

a significant year effect on grazing time with calves grazing 240 and 106 min for Years

2003 and 2004, respectively. The large difference in grazing time between years can be

explained in part by the greater herbage mass present in 2003.









Table 4-6. Grazing time during daylight hours of early weaned calves grazing Tifton 85
pastures supplemented with three levels of concentrate. (BW=body weight)
Concentrate Grazing time
-----------g kg BW------ -------------------min--- --------------

10 193
15 181
20 147

Polynomial contrast LT
P value <0.01
SEM 8

L=linear

There was no effect of increasing supplementation rates on calf BUN (Table 4-7).

Based on the high CP concentration of the forage (180 g kg-1) and concentrate (160 g kg

1), it was expected that calves consuming more forage would have a higher BUN

concentration; however, the forage rumen-degradable protein (550 g kg-1) was less than

concentrate rumen degradable protein (700 g kg-1) (NRC, 1996). As a result, calves

consumed similar total rumen degradable protein, resulting in similar BUN among the

treatments. According to Hammond et al. (1993), cattle BUN concentrations from 9 to 12

mg dL-1 are considered to be in a transition range below which daily response to protein

supplementation has been positive. The calf BUN concentrations in this study suggest

that protein supplementation is unlikely to improve performance of the calves. Further

studies are necessary to clarify the relation of BUN concentrations and performance of

early weaned calves grazing warm-season pastures.









Table 4-7. Blood urea nitrogen (BUN) concentrations of early weaned calves grazing
Tifton 85 pastures supplemented with three levels of concentrate. (BW=body
weight)
Concentration BUN
-----------g kg-1 BW---------------------------------------mg dL--------------------

10 14.8

15 15.4

20 14.0

Polynomial contrast NS1
P value 0.64

SEM 0.9
SNS = not significant

Economic Analysis

Cost and income increased linearly with increasing supplement rate (Table 4-8).

The increase in income was more than compensated for by greater concentrate cost at the

highest supplementation rate. There was a quadratic effect on economic return indicating

that the 15 g kg-1 BW treatment provided greater return than the average of the 10 and 20

g kg-1 treatments.

Table 4-8. Economic analysis of three concentrate levels for early weaned calves grazing
Tifton 85 pastures.
Concentrate Concentrate cost Income Return
1 -1
-------g kg BW------- ----------------------- $ ha--- -----------

10 1100 2380 1280

15 1670 3200 1530

20 2710 3400 700

Polynomial contrast LT L Q
P value <0.01 <0.01 0.04

SEM 74 187 284


t L = linear, Q = quadratic