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| Title Page | |
| Dedication | |
| Acknowledgement | |
| Table of Contents | |
| List of Tables | |
| List of Figures | |
| Abstract | |
| Introduction | |
| Cogongrass [imperata cylindrica... | |
| The influence of herbicide combinations... | |
| The influence of mechanical and... | |
| The influence of stage of development... | |
| Summary and conclusions | |
| Appendices | |
| Literature cited | |
| Biographical sketch |
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Page i Dedication Page ii Acknowledgement Page iii Table of Contents Page iv Page v Page vi List of Tables Page vii Page viii List of Figures Page ix Abstract Page x Page xi Page xii Introduction Page 1 Page 2 Cogongrass [imperata cylindrica (L.) Beauv.] distribution on Florida highway rights-of-way Page 3 Page 4 Page 5 Page 6 Page 7 Page 8 Page 9 Page 10 Page 11 Page 12 Page 13 Page 14 Page 15 Page 16 Page 17 The influence of herbicide combinations and application technology on cogongrass [imperata cylindrica (L.) Beauv.] control Page 18 Page 19 Page 20 Page 21 Page 22 Page 23 Page 24 Page 25 Page 26 Page 27 Page 28 Page 29 Page 30 Page 31 Page 32 Page 33 Page 34 Page 35 Page 36 Page 37 Page 38 Page 39 Page 40 Page 41 Page 42 Page 43 Page 44 Page 45 Page 46 Page 47 Page 48 The influence of mechanical and chemical inputs on cogongrass [imperata cylindrica (L.) Beauv.] control Page 49 Page 50 Page 51 Page 52 Page 53 Page 54 Page 55 Page 56 Page 57 Page 58 Page 59 Page 60 Page 61 Page 62 Page 63 Page 64 Page 65 Page 66 Page 67 Page 68 Page 69 Page 70 Page 71 Page 72 Page 73 Page 74 Page 75 Page 76 Page 77 Page 78 The influence of stage of development and mowing on bahiagrass [paspalum notatum var. saurae parodi ‘Pensacola’] and cogongrass [imperata cylindrica (L.) Beauv.] interference Page 79 Page 80 Page 81 Page 82 Page 83 Page 84 Page 85 Page 86 Page 87 Page 88 Page 89 Page 90 Page 91 Page 92 Page 93 Page 94 Summary and conclusions Page 95 Page 96 Page 97 Appendices Page 98 Page 99 Page 100 Page 101 Page 102 Page 103 Page 104 Page 105 Literature cited Page 106 Page 107 Page 108 Page 109 Page 110 Page 111 Page 112 Biographical sketch Page 113 Page 114 Page 115 Page 116 |
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BIOLOGY, ECOLOGY AND MANAGEMENT OF COGONGRASS [Imperata cylindrica (L.)Beauv.] By TOMMY RAY WILLARD 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 1988 ; ; iii .;. i ii ~iiil;;;..il li',iilEiiiiiiiiii ;il To my loving wife Donna and my daughter Stacey I humbly dedicate this work. No words can adequately describe how you have enhanced the quality of my life. ACKNOWLEDGEMENTS For their support during my tenure at the University of Florida I wish to express my sincere appreciation to my committee members: Dr. William Haller, Dr. Donn Shilling, Dr. David Hall, Dr. Barry Brecke, and Dr. Paul Mislevy. I also wish to thank Dr. Wayne Currey for his guidance during my Masters program and the majority of my Ph.D. studies. I especially acknowledge Dr. Donn Shilling, co-chairman of my committee; without his unending patience and willingness to teach I would not be at this point today. I thank Mr. David Studstill and Mr. Tim Pederson, without whose technical assistance it would have been extremely difficult, if not impossible, to conduct the research presented. During my tenure at the University of Florida I was fortunate to have been associated with graduate students of the highest caliber. The friends I made here will always be remembered. I thank my parents, Mr. and Mrs. Morris Willard for their love and understanding. The values they instilled in me during my youth are responsible for all I have attained. k. iii TABLE OF CONTENTS PAGE ACKNOWLEDGEMENTS.................................... iii LIST OF TABLES................................. ....... vii LIST OF FIGURES............. .................... ..... ix ABSTRACT.............................................. x CHAPTERS I INTRODUCTION................................... 1 II COGONGRASS [Imperata cylindrica (L.)Beauv.] DISTRIBUTION ON FLORIDA HIGHWAY RIGHTS-OF-WAY 3 Introduction.....*** .......................*... 3 Materials and Methods.......................... 7 Results and Discussion......................... 9 III THE INFLUENCE OF HERBICIDE COMBINATIONS AND APPLICATION TECHNOLOGY ON COGONGRASS [Imperata cylindrica (L.)Beauv.] CONTROL............... 18 Introduction..........................,........ 18 Materials and Methods.......................... 21 Cogongrass rhizome biomass sampling.......... 21 'Sequential herbicide application programs.... 22 Tank-mix combinations of imazapyr and glyphosate... ...... ...................... 24 Comparison of low and conventional volume applications of imazapyr and glyphosate.... 25 Ropewick applications of imazapyr and glyphosate................................. 27 Imazapyr longevity bioassay.................. 28 Results and Discussion........*........,...... 32 Cogongrass rhizome biomass sampling......... 32 Sequential herbicide application programs.... 34 Tank-mix combinations of imazapyr and glyphosate.. ...... ............. o.......... 40 Comparison of low and conventional volume applications of imazapyr and glyphosate.... 42 Ropewick applications of imazapyr and glyphosate................................. 44 Imazapyr longevity bioassay.................. 46 IV THE INFLUENCE OF MECHANICAL AND CHEMICAL INPUTS ON COGONGRASS [Imperata cylindrica (L.)Beauv.] CONTROL....... ......... ...................... 49 Introduction............................... 49 Materials and Methods...................... 51 Integration of mowing and disking with herbicide treatments................... 51 The influence of photoperiod on shoot initiation in cogongrass rhizomes...... 55 The influence of cogongrass stage of development and defoliation on glyphosate efficacy ................... 56 Results and Discussion..................... 60 Mowing-herbicide studies................. 60 Disking-herbicide studies................ 65 The influence of photoperiod on shoot initiation in cogongrass rhizomes...... 71 The influence of cogongrass stage of development and defoliation on glyphosate efficacy.................... 73 V THE INFLUENCE OF STAGE OF DEVELOPMENT AND MOWING ON BAHIAGRASS [Paspalum notatum var. saurae Parodi 'Pensacola'] AND COGONGRASS [Imperata cylindrica (L.)Beauv.] INTERFERENCE.......... 79 Introduction............................... 79 Materials and Methods...................... 81 Preliminary Studies...................... 81 Seedling bahiagrass and emerging cogon- grass interference studies............. 83 The influence of mowing on established bahiagrass and emerging cogongrass interference........................... 84 Experimental design and analysis......... 86 Results and Discussion..................... 86 Seedling bahiagrass and emerging cogon- grass interference studies............. 86 The influence of mowing on established bahiagrass and emerging cogongrass interference..............** ........... 90 VI SUMMARY AND CONCLUSIONS......................... 95 APPENDICES A OCCURENCE AND SEVERITY OF Andropogon sp. ON FLORIDA HIGHWAY RIGHTS-OF-WAY................. 98 B OCCURRENCE AND SEVERITY OF Phragmites australis ON FLORIDA HIGHWAY RIGHTS-OF-WAY ............. 99 C OCCURRENCE AND SEVERITY OF Panicum maximum ON FLORIDA HIGHWAY RIGHTS-OF-WAY................. 100 D OCCURRENCE AND SEVERITY OF Sorghum halepense ON FLORIDA HIGHWAY RIGHTS-OF-WAY................ 101 E OCCURRENCE AND SEVERITY OF Pennisetum purpureum ON FLORIDA HIGHWAY RIGHTS-OF-WAY.............. 102 F OCCURRENCE AND SEVERITY OF Brachiaria mutica ON FLORIDA HIGHWAY RIGHTS-OF-WAY................. 103 G OCCURRENCE AND SEVERITY OF Sporobolus indicus ON FLORIDA HIGHWAY RIGHTS-OF-WAY................. 104 H OCCURRENCE AND SEVERITY OF Paspalum urvillei ON FLORIDA HIGHWAY RIGHTS-OF-WAY................. 105 LITERATURE CITED................ ..................... 106 BIOGRAPHICAL SKETCH................................... 113 LIST OF TABLES TABLE PAGE 2.1 Perennial grass weeds frequently occurring on Florida highway rights-of-way....6....... 5 2.2 Right-of-way distance surveyed by district in 1984-85 for perennial grass weeds by the Florida Department of Transportation........ 10 3.1 Cogongrass rhizome biomass collected in Chiefland, FL from 1986 to 1988............. 33 3.2 The effect of sequential herbicide treatments on cogongrass foliage and rhizome dry weight (Experiment 1 1985 through 1987 Area A). 35 3.3 The effect of sequential herbicide treatments on cogongrass foliage and rhizome dry weight (Experiment 2 1986 through 1988 Area B). 36 3.4 The effect of imazapyr and glyphosate tank- mixes on cogongrass foliage and rhizome dry weight...................................... 41 3.5 The effect of carrier volume and imazapyr and glyphosate rate on cogongrass foliage dry weight...................................... 43 3.6 The effect of ropewick applications of imazapyr and glyphosate on cogongrass foliage and rhizome dry weight.............. 45 3.7 The influence of imazapyr dissipation on the inhibition of dry weight (IDW) response of four grass species.......................... 47 4.1 Timing of mechanical and chemical treatments and harvest performed for evaluation of integrated cogongrass control studies....... 54 4.2 The effect of mowing and herbicides on cogon- grass foliage and rhizome dry weight (Experi- ment 1 1985 through 1987)................. 61 vii 4.3 The effect of mowing and herbicides on cogon- grass foliage and rhizome dry weight (Experi- ment 2 1986 through 1988)................. 64 4.4 The effect of disking and herbicides on cogon- grass foliage and rhizome dry weight (Experi- ment 1 1985 through 1987)................. 66 4.5 The effect of disking and herbicides on cogon- grass foliage and rhizome dry weight (Experi- ment 2 1986 through 1988)................. 69 4.6 The influence of photoperiod on shoot initiation from cogongrass rhizomes at one week after planting.......*................. 72 4.7 The influence of time of defoliation on greenhouse-grown cogongrass growth.......... 74 4.8 The influence of defoliation on glyphosate efficacy in cogongrass...................... 76 5.1 The interaction of seedling bahiagrass (B) and emerging cogongrass (C) as measured by relative yield (RY), relative yield total (RYT) and relative crowding coefficient (RCC) of leaf dry weight and height and total pot dry weight at eight weeks after planting.... 87 5.2 The influence of mowing on the interaction of established bahiagrass (B) and emerging cogongrass (C) as measured by relative yield (RY) and relative crowding coefficient (RCC) using leaf dry weight produced by each species..................................... 91 5.3 The influence of mowing and bahiagrass competition on cogongrass height, shoot number and rhizome dry weight............... 92 viii LIST OF FIGURES FIGURE PAGE 2.1 Occurrence and severity of Imperata cylindrica on Florida highway rights-of-way. 12 3.1 Weekly precipitation in Gainesville, FL, 1986 through 1987........................... 38 ix 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 BIOLOGY, ECOLOGY AND MANAGEMENT OF COGONGRASS [Imperata cylindrica (L.) Beauv.] By Tommy Ray Willard December, 1988 Chairman: Dr. W. T. Haller Major Department: Agronomy Field and greenhouse studies were conducted to deter- mine the distribution and management of cogongrass and the effects of this weed and control strategies on desirable vegetation. A survey of 22% of the Florida highway rights- of-way indicated that cogongrass is distributed along major thoroughfares from Lake Okeechobee to the Panhandle region. The greatest infestation level was found in north-central Florida. Field studies conducted on herbicide and application technology for cogongrass control indicated that single herbicide applications would not provide long-term control, but control could be acheived with sequential treatments of imazapyr and glyphosate. Ropewick applications of imazapyr were as effective as conventional spray applications. Imazapyr or glyphosate applied in 46 L'ha-1 water did not increase cogongrass control over 234 L'ha- applications. The half-life of imazapyr in sandy soils was determined to be eight months and residual imazapyr affected seedling growth of four grass species in the following order (most to least affected): ryegrass > browntop millet > bermudagrass > bahiagrass. Field and greenhouse studies, which were conducted to determine the influence of mowing and disking alone and when combined with herbicide treatments, indicated that mowing alone has little effect on cogongrass. Disking cogongrass one time caused an increase in stand density, however moder- ate control was acheived with two diskings. The combination of imazapyr or glyphosate with two diskings provided nearly 100% cogongrass control. Other studies indicated that glyphosate should be applied during the fall to maximize translocation to, and control of, cogongrass rhizomes. Replacement studies indicated that cogongrass emerging from rhizomes competed effectively with, and could displace, seedling bahiagrass. However, cogongrass growth from rhi- zomes was severely inhibited when competing interspecifically with established bahiagrass. Mowing negatively impacted cogongrass rhizome dry weight when competing intraspec- ifically, but shoot number increased. This did not occur under interspecific competition due to the overwhelming xi impact of bahiagrass. Mowing had less impact on bahiagrass than cogongrass. Differences in leaf morphology (cogongrass being erect with little leaf lamina below 10 cm and bahia- grass being more prostrate with significant leaf area below 10 cm) appear to favor bahiagrass under mowed conditions. xii CHAPTER 1 INTRODUCTION. Cogongrass [Imperata cylindrica (L.)Beauv.] is an exotic, aggressive, perennial grass. During the last fifty years it has become a serious pest in pastures, pines, and other uncultivated areas in Florida. It went virtually unnoticed as a pest in this state until approximately 1972, even though it already covered four thousand hectares in south Alabama. By 1982, botanists and weed specialists in Florida were being routinely questioned as to the identifica- tion and control of this grass that had "suddenly appeared". The need for soil stabilization materials and low-input forages in Florida led to the introduction and intentional, as well as unauthorized, dissemination of cogongrass. How- ever, researchers realized too late that the cogongrass spread beyond soil stabilization areas and was not a quality, productive forage (Tabor, 1949). During this same period (1940-50s) authorities in Alabama and Mississippi were destroying plantings of cogongrass on their experiment stations. The Florida Department of Transportation in 1984 funded cogongrass research to be conducted by the Agronomy Depart- ment of the University of Flotida. The goals of this research were to characterize the growth and development of cogongrass in Florida and to use this information to develop environmentally-sound and effective control strategies. The objectives were to: 1) define the distribution of cogongrass on Florida highway rights-of way, 2) evaluate available herbicide and application technology, 3) integrate mechan- ical and chemical control strategies, and 4) to evaluate the competitive interaction of cogongrass and bahiagrass as it is influenced by vegetation management practices. Cogongrass does not discriminate between roadsides and other favorable habitats; therefore, these results should be used not only to guide cogongrass control programs on Florida highway rights-of-way, but could be applied to other infested areas. It may not be possible to eradicate cogongrass from Florida; however, a diligent effort to manage as many infestations as possible could reduce the threat of this species. CHAPTER 2 COGONGRASS [Imperata cylindrica (L.)Beauv.] DISTRIBUTION ON FLORIDA HIGHWAY RIGHTS-OF-WAY. Introduction The Florida Department of Transportation (FDOT) maintains 45,326 hectares of roadside vegetation along 18,500 km of highway Maintenance practices, which include mowing, fertilization, tree and brush removal, wildflower management, and selective herbicide treatments are necessary to create a safe, aesthetically pleasing thoroughfare for the state's large resident and tourist populations (Evink, et al., 1983). Therefore, the FDOT allocates approximately twenty million dollars annually for these activities. The same environmental conditions that are conducive to the establishment of turfgrass species [predominantly bahia- grass, Paspalum notatum var. saurae Parodi 'Pensacola' #2 PASNS, with sporadic populations of common bermudagrass, Cynodon dactylon (L.)Pers. # CYNDA, and centipedegrass Eremochloa ophiuroides (Munro)Hack. # ERLOP] on these Personal communication, 1986, J.A. Lewis, Florida Department of Transporation, Tallahassee, FL. Letters following this symbol are a WSSA-approved computer code from the Composite List of Weeds, Weed Sci. 32,Suppl. 2. Available from WSSA, 309 West Clark Street, Champaign, IL 61820. rights-of-way also favor the establishment of undesirable perennial grasses. Rank growth of these weedy species, which rapidly exceeds the height of desirable vegetation, causes several problems. Motorist safety is jeopardized by. limited sight distance caused by tall vegetation during the growing season and also by smoke from fires fueled by this vegetation during the dry, winter months when the foliage is dessicated. Excessive vegetation increases maintenance costs by reducing mowing efficiency (km per man-hour) and by increasing equipment repair and subsequent downtime. Mowing has provided insignificant control of these species, and has been shown to stimulate seed production and dispersal in some of them (McCaleb and Hodges, 1971). The use of herbi- cides to control weeds and reduce mowing costs has increased as infestations have expanded (Lewis, 1985). Although less tangible, motorist reaction to the poor aesthetics caused by weedy vegetation and the use of herbicides has been negative. In 1981, the FDOT Bureau of Environment cited nine grassy weed species that required additional efforts beyond those needed to maintain turf species (Evink, et al., 1983). Table 2.1 lists the common and scientific names of these grasses. Selective control of these weeds is very difficult, as the weeds and the turf are both warm-season perennial grasses. Specialized equipment, such as ropewick applicators, can be used in some situations to achieve selective control. For ropewick treatments to provide optimal control without turf Table 2.1. Perennial grass weeds frequently occurring on Florida highway rights-of-way. Common Name Bluestems or Broomgrasses Cogongrass Giant Reed Guineagrass Johnsongrass Napiergrass Paragrass Smutgrass Vaseygrass Scientific Name Andropogon virginicus L. I ANOVI Andropogon virginicus L. # ANOVI Andropogon glomeratus (Walt.)BSP. # ANOGL Imperata cylindrica (L.)Beauv. # IMPCY Phragmites australis (Cav.)Trin. ex Steud # PHRCO Panicum maximum Jacq. # PANMA Sorghum halepense (L.)Pers. # SORHA Pennisetum purpureum Schumach. # PESPU Brachiaria mutica (Forsk.)Stapf # PANPU Sporobolus indicus (L.)R. Br. # SPZIN Paspalum urvillei Steud. # PASUR injury, a significant height differential between the weed and the turf is required. Herbicide applications made with a ropewick are slow compared to broadcast applications, more expensive, and may conflict with the mowing protocols of the FDOT management districts (Lewis, 1985). Nonselective control, usually attained with handgun applications, is undesirable from an aesthetic standpoint due to the dessica- tion of large areas of vegetation and could also lead to erosion. These selective and nonselective approaches are intensive in terms of chemical and manpower expenditures. The applications are usually employed only when infestations are large enough to warrant expenditure of manpower and equipment. This approach to weed control has resulted in reduced herbicide efficacy on large infestations (making repeat applications necessary to achieve long-term control) and the continued expansion of smaller untreated infesta- tions. Glyphosate, isopropylamine salt of N-(phosphonomethyl)- glycine, dalapon, magnesium and/or sodium salts of 2,2- dichloropropionic acid, and hexazinone, 3-cyclohexyl-6-(di- methylamino)-l-methyl-1,3,5-triazine-2,4(1H,3H)-dione are the primary chemicals used for control of perennial grass weeds in Florida (Lewis, 1985). However, several applica- tions may be necessary for optimal control, and in the case of cogongrass, improved application and/or herbicide technology is needed. The FDOT surveys its rights-of-way to assess needs and develop programs for roadway and roadside maintenance. Previous surveys have not focused on the identification, distribution, and intensity of perennial grass weed infesta- ..tions. To obtain this information FDOT district maintenance engineers surveyed major rights-of-way for the nine weed species listed in Table 2.1. MATERIALS AND METHODS A survey of Florida highway rights-of-way was conducted in 1984-85 for the occurrence and severity of nine perennial grass weeds. The purpose of the survey was, first, to give district personnel the opportunity to identify the problem species on a county or individual roadway basis and second, to assess the impact of maintenance activities (i.e., mowing, chemical control, or roadside construction) on these species. District maintenance engineers, who were involved in roadway selection and survey format, conducted the survey, and university investigators verified and compiled the data. The criteria for roadway selection were: 1) all limited access roadways including interstate highways and the Florida Turnpike; 2) where possible, at least one north- south and one east-west roadway per county in addition to those selected under criteria 1; and 3) urban sections of roadway were excluded. The survey format included location data (county name, section number, roadway number, and mileage marks) and a zero to three rating scale with a zero rating indicating that the particular species was not present on that section of roadway, a rating of one indicating that the species was present in only sparse infestations and that present maintenance/control practices provided adequate ..control, a rating of two indicating the species was more frequently encountered and present maintenance/control practices provided only partial control, and a rating of three indicating numerous large infestations and/or little control was realized from existing programs. Additional space was provided on the survey form to note any unusual conditions or to more precisely locate major cogongrass and napiergrass infestations. Using this rating scale, cogongrass and napiergrass would receive a rating of three, in most situations, due to the lack of effective control programs. These two species pose an extreme problem for roadway maintenance personnel and are a threat to adjoining agricultural, silvicultural, and residential areas. To gain a perspective of the relative intensity of these two species, those counties with one to five infestations of 100 m2 or greater were assigned a rating of one (low), counties with six to fifteen infesta- tions were assigned a rating of two (moderate), and counties with greater than fifteen infestations were assigned a rating of three (severe). Training seminars were conducted to familiarize personnel with the survey format and identification of each species. Trial surveys of selected roadways were also conducted. Surveyors were given eighteen months to complete their surveys of assigned roadways, which afforded them the opportunity to evaluate mowed or reconstructed areas after sufficient weed regrowth had occurred. Districts IV and VI did not use the survey form as the recording medium. A K-5000 Nu-Metrics Distance Measuring Instrument (DMI)3 and printer were programmed with event numbers corresponding to each species. This instrument continuously monitored miles traveled. As infestations were identified, the operator entered the appropriate event number which was printed with the mileage mark. From these printouts the principle investigator assigned severity ratings for the counties based on the number and size of infestations in a manner similar to that used for cogongrass and napiergrass. It should be emphasized that the data presented is subjective in nature and is limited to those roadways surveyed. Therefore, a species that is represented as not occurring in a particular county may, in fact, occur on other unsurveyed roadways or in off right-of-way areas. Results and Discussion Table 2.2 presents the geographical scope of the survey by district. The 8234 km surveyed was 92.1% of that designated for survey. Though varying with locale and surveyor, the districts using'the survey sheet recorded an 3Nu-metrics Instrumentation, Vanderbilt, Pa, 15486-0471. Table 2.2. Right-of-way distance surveyed by district in 1984-85 for perennial grass weeds by the Florida Department of Transportation. Distance Geographic No. of Surveyed % of District Region Counties km Centerline I SW Gulf Coastal 14 2182 28.1 through S Central II N Central through 18 1709 21.4 N Atlantic Coastal III West Florida 16 1377 18.5 (Panhandle) IV SE Atlantic Coastal 7 1078 21.0 V Central through 10 1415 21.3 Central Atlantic Coastal VI Everglades 2 473 22.5 (Monroe and Dade counties) Statewide Total 67 8234 22.2 1Percentage of centerline distance computed by dividing the kilometers surveyed by two to account for surveying each shoulder. average of one observation for every 10.1 centerline km. In Districts IV and VI where the DMI was utilized, recorded observations averaged one for every 0.5 centerline km. This points out the greater accuracy that can be achieved by using an automatic recording device such as the DMI. The distribution and severity of cogongrass has been reported for Alabama, Mississippi, and Louisiana (Dickens, 1974; Patterson, et al., 1983). The major emphasis of this survey was to determine the range and severity of cogongrass on Florida highway rights-of-way. The distribution maps for the other eight species surveyed are presented in Appencies A through H but will not be elaborated upon further. The distribution and severity of cogongrass on Florida highway rights-of-way is presented in Figure 2.1. Rights-of-way in the north central region (Districts II and V) and the south central region (Districts I and IV) were severely infested with cogongrass. Historical records indicate that these locations correspond to intentional propagation of this species for forage and soil stabilization purposes during the 1940s and 1950s (Dickens and Buchanon, 1971; Tabor, 1949). Points of introduction in Florida include Gainesville (University of Florida Experiment Station), Brooksville (USDA Plant Introduction Station), and Withlacoochee (Soil Conservation Service reclamation area). Once introduced, dispersal of cogongrass can occur by two mechanisms, seeds and rhizomes. Cogongrass produces plummed wind-blown seeds beginning in January (in response DISICT IS DIST I O NONEREPORTED MLow MODERATE ICT DISTRICT I SEVERE DISTRICT L v DISTRICT L IV Figure 2.1. Occurrence and severity of Imperata cylindrica on Florida highway rights-of-way. DISTRICT to the freezing temperatures, Holm, et al., 1977) and continuing until May (personal observation). Dickens and Moore (1974) found that cogongrass seed collected from Grand Bay, Alabama, was viable soon after anthesis and germinated readily under laboratory conditions. They also reported a significant temperature response, with the optimum germina- tion (60%) occurring at 30 C, but less than 10% germination at 20 C. The viability of the seed declined steadily under storage in sealed vials. The significance of cogongrass seed production in Florida is undetermined at this time. Several factors appear to limit the impact that seeds could have on the overall spread of cogongrass on Florida highway rights- of-way. First, the plant begins seed production during the coldest and driest months of the year, conditions which do not favor immediate germination. Secondly, Eussen and Soerjani (1975) reported that flowers produced in response to environmental stress, such as freezing temperatures, seldom contained seed. If seed is produced, research has shown that they do not accumulate in the soil seed bank (Hopkins and Graham, 1984). Thirdly, the orientation of the majority of infestations on highway rights-of-way is parallel to the roadway. Winds funneling through these tree-lined corridors and turbulence caused by vehicles would tend to disperse the seeds within the length of the infestation, as the distance cogongrass seeds travel has been estimated to average only fifteen meters from the seedhead (Holm, et al., 1977). Fourthly, if viable seeds are dispersed beyond the bounds of the infestation area, they are unable to penetrate into adjoining forested areas due to the hairs which catch on vegetation. If they are deposited onto the right-of-way, research has shown that their survival is dependent on con- --tacting moist, bare soil that is free of competing vegetation (Kushwaha, et al., 1983). Lastly, cogongrass seedlings have been reported to be frail and unlikely to survive (Kushwaha, et al., 1983) except under ideal conditions (Dickens, 1976). However, environmental and soil conditions in the phosphate mining area of the state near Bartow (Polk county, District I) could be favorable cogongrass seedlings on off right-of- way areas. This ability to spread by seeds into open areas could also pose a major threat to citrus production on the central Florida ridge. The second method of cogongrass dispersal is by rhizomes. From established infestations it has been reported that cogongrass will expand laterally 25-40 cm per year (Dickens, 1973). Infestations on Florida highway rights-of- way tend to be isolated and occurring at irregular intervals. Roadway construction requires that large quantities of soil be transported to the site to create front and back drainage slopes. This bulk movement of potentially rhizome-contami- nated soil from maintenance unit barrow pits may be a major avenue of dispersal in Florida. This may have occurred on interstate highway 75 (1-75) and U.S. 441/301. These two roadways which traverse Alachua (District II) Marion, Sumter (District V), Polk, Highlands (District I), and Okeechobee (District IV) counties are heavily infested with cogongrass. Alachua and Marion counties are the most heavily infested in the state, on rights-of-way as well as in pastures and pulpwood production areas. These heavy infestations - occurred due to the aforementioned intentional planting of cogongrass followed by intense roadway construction in these counties during the mid to late 1960s (Lewis, 1986). District III reported several isolated infestations. These infestations occurred on highways U.S. 90 and interstate 10 (1-10). During the 1940s cogongrass was removed from university experiment stations without authorization. It was estimated that four hundred hectares were planted in west Florida during this period (Tabor, 1952). There is little doubt that transport of cogongrass occurred on U.S. 90 which was the major east-west corridor during this time, prior to the completion of I-10 in the 1950s. Movement of cogongrass from Florida to Alabama was reported by Dickens (1974). West to east movement of cogongrass may have also occurred via U.S. 90 and 1-10 (which parallels U.S. 90) as these two roadways pass through Grand Bay and Mobile, Alabama, which have been infested with cogongrass since the 1940s (Tabor, 1949). However, observa- tions made in 1988 of cogongrass along 1-10 from Grand Bay to Pensacola, Florida, showed that Alabama infestations ended 15-25 km west of the state line and Florida infesta- tions began approximately 15 km east of the state line. Tallahassee, the state capital (Leon county, District III), is a major transportation hub because of the conver- gence of I-10, U.S. 90, and U.S. 19/27. The interchanges surrounding Tallahassee are heavily infested with cogongrass. .This reinforces the hypothesis that roadway construction and maintenance has played a major role in the distribution of cogongrass in Florida. The dispersion of cogongrass in Florida does not appear to be as rapid as that experienced in Alabama. However, its spread will continue to increase unless control programs are enacted immediately. In the author's opinion, there should be a coordinated effort headed by the Florida Department of Agricultural and Consumer Services that will at least partially subsidize the cost of cogongrass control in all areas of the state. This program should be conducted in a manner similar to that implemented for witchweed [Striga asiatica (L.)Ktze. # STRLU] in North Carolina (USDA, 1970). Efforts should also be made to more strictly regulate the introduction of plant species into the United States under the Federal Noxious Weed Law. There are numerous examples of species that have been introduced in the past for forage or soil stabilization/improvement purposes that have, with time, proven to be unsuitable for the intended purpose or have exhibited characteristics that enable them to success- fully compete with and eventually displace desirable vegeta- tion [e.g., kudzu, Pueraria lbbata (Willd.)Ohwi # PUELO, melaleuca, Melaleuca quinquenervia (Cav.)Blake # MLAQU, and torpedograss, Panicum repens L. # PANRE] (Williams, 1980). Efforts by the Animal and Plant Health Inspection Service (APHIS) have prevented the introduction of potentially hazardous insects and plant pathogens. The effort placed on Identifying, locating, and eventually controlling noxious plant species should receive the same level of diligence. The sedentary nature of weeds and the relatively slow encroachment that they make have allowed many introduced species to become established to the point that they cannot be eradicated. It has been forty years since R. L. Pendleton wrote: Certainly its [cogongrass'] hazard as a poten- tial weed for upland crops in the tropical and subtropical portions of the western hemisphere is a very much more serious threat to agriculture than the small amount of benefit it can possibly be as a forage. The writer feels very strongly that steps should be taken at once to completely eradicate this noxious weed from the western hemisphere. (Pendleton, 1948, p. 1048). If control measures are not implemented in the near future in all states with cogongrass infestations, many hectares of valuable pasture and silvicultural land could become unproductive. CHAPTER 3 THE INFLUENCE OF HERBICIDE COMBINATIONS AND APPLICATION TECHNOLOGY ON COGONGRASS [Imperata cylindrica (L.)Beauv.] CONTROL. Introduction Cogongrass [Imperata cylindrica (L.)Beauv. # IMPCY] is a serious pest throughout the sub-tropical and tropical regions of the world. It ranks as the seventh most trouble- some weed worldwide (Holm, et al., 1977). Banana (Musca x paradisiaca L.), citrus (Citrus species), coconut (Cocus nucifera L.), oil palm (Elaeis guineensis Jacq.), pastures, pineapple [Ananas comosus (L.)Merrill], pine (Pinus species), rubber [Hevea brasiliensis (Willd. ex A. Juss.)Mull. Arg.], and tea [Camellia sinensis (L.)0. Kuntze] are major crops that have been reported to be adversely affected by the presence of cogongrass. With little utility except for thatch and short-term forage production and soil stabiliza- tion, cogongrass research has been geared toward control. However, control of this species has proven to be extremely difficult. Slash and burn and shifting agriculture have resulted in transient control, usually allowing only a year 1Letters following this symbol are a WSSA-approved computer code from the Composite List of Weeds, Weed Sci. 32,Suppl. 2. Available from WSSA, 309 West Clark Street, Champaign, IL 61820. or two of crop production before reinfestation takes its toll. By eliminating natural vegetation that competes effectively with cogongrass and concomitantly distributing seeds and rhizomes, these control strategies have increased the area of cogongrass infestation (Prommool, 1984). With the advent of chemical weed control in the 1950s efforts were made to apply this technology to cogongrass management. Since that time, at least thirty compounds and hundreds of combinations have been evaluated and reported for cogongrass control (Dickens and Buchanon, 1975; SEAWIC, 1988). Of these herbicides, glyphosate, N-(phosphonomethyl)- glycine, dalapon, 2,2-dichloropropanoic acid, and imazapyr, ()-2-[4,5-dihydro-4-methyl-4-(1-methylethyl)-5-oxo-lH- imadazol-2-yl]-3-pyridinecarboxylic acid, have shown the greatest activity on cogongrass with the fewest adverse effects (i.e., bioaccumulation of heavy metals, extended periods of soil sterilization, non-target species injury, or applicator injury). In most situations, long-term control was not acheived from a single application of any of these compounds. Repeat applications have proven to be necessary to directly kill or deplete the rhizome biomass, which is the basis of cogongrass' ability to survive and spread. Various innovations in application technology have also been tested to improve the activity of these herbicides in cogongrass. Low and ultra-low volume (ULV) applications (usually in the range 20 to 100 L'ha-1 carrier) have been reported to enhance the activity of glyphosate in common bermudagrass [Cynodon dactylon (L.)Pers. # CYNDA] (Erickson and England, 1983; Jordan, 1981), quackgrass (Agropyron repens (L.)Beauv. I AGRRE) (Sandberg, et al., 1978), and torpedograss [Panicum repens L. # PANRE] (Baird, et al., 1983). The use of this low-volume technology with glyphosate (Arif, et al., 1986) and imazapyr (Bacon, 1986) on cogongrass has been reported. However, results have been variable and not definitive as to enhancement of activity or the long-term control realized. Ropewick and roller-type applicators have been used for the past ten years for selective control of perennials in susceptible crops such as cotton (Gossypium hirsutum L.), soybean [Glycine max (L.)Merr.], and pastures (Dale, 1981; Wiese and Lavake, 1980). The type of applicator, speed, herbicide concentration, and number of applications have all been reported to influence the efficacy of glyphosate when applied in this manner (Derting, 1981). The concept of control has been frequently defined as the initial effects of the herbicide treatments. In a few cases, excellent control has been reported in as little as two weeks (Al-Juboory and Sarmaly, 1984) and usually within the same growing season. The research that provides the most useful information (i.e., long-term efficacy) about cogongrass control is that which allows sufficient time to elapse following treatment before evaluation, such as those studies conducted by Dickens (1973). The most accurate measure of perennial weed control is the inhibition of regrowth from perenniating organs. This can be accomplished by harvests of foliage regrowth and the amount of productive perenniating tissue remaining after an extended period of time (i.e., one growing season following application). Therefore, the objective of this research was to study the influence of herbicide combinations, both in-tank mixtures and sequential herbicide programs, and application methodology (i.e., low-volume and ropewick applications) for long-term cogongrass control. Materials and Methods Cogongrass Rhizome Biomass Sampling. To determine the influence of time and to characterize the infestation area that was to be used for the research henceforth described, cogongrass rhizomes were harvested from a site in Chiefland, Florida. Sampling of two designated areas was conducted from October, 1986, through April, 1988. Historical information provided by the land- owner indicated that cogongrass was introduced in the 1950s following intentional planting for fire break stabilization within a pulpwood production area. Cogongrass was not a problem in this area until approximately 1975. A lightning strike caused the site to burn following harvest of mature trees, land preparation, and planting of seedling pines. With the competing vegetation removed, cogongrass dominated this eight hectare area within a year. This infestation then spread to adjoining cleared areas and throughout mature pines to cover an area of approximately 20 ha. Other species present were dogfennel [Eupatorium capillifolium (Lam.)Small # EUPCP], partridgepea (Cassia fasciculata Michx. # CASFA), saw palmetto [Serenoa repens (Bartr.)Small # SERRE], and maidencane (Panicum hemitomon Schultes # PANHE). The infestation was divided into two equal size areas. Area A was known to be the older, more established section from which Area B originated. With the exception of January, April, and December, 1987, monthly samples were taken from each of these sections. For each sampling time, six soil cores from each area were taken using a bucket auger 182 cm2 in surface area to a depth of 23 cm. Rhizomes which were not decayed were removed and dried at 100 C for 24 hrs. From these samples rhizome biomass on a per hectare basis was computed. Analysis of variance (Helwig and Council, 1982) was used to test for main factor (area and sampling time) effects and the interaction. Least significant difference (LSD) procedure was used to compare rhizome biomass between areas within a sampling time. Sequential Herbicide Application Programs. To determine the influence of combinations of initial and sequential herbicides on cogongrass control, two experi- ments were conducted over a three-year period at Chiefland, Florida. The area was located in a flatwoods, pulpwood production site. The soil type was a Sparr fine sand (loamy, siliceous, hyperthermic, Grossarenic Paleudults). The herbicides and rates utilized for these studies were selected from a herbicide screening field study conducted in 1983-1984 (data not shown). The herbicides and rates applied were: 1) imazapyr at 0.8 kg (active ingredient) ai'ha-1, 2) glyphosate at 3.4 kg ai'ha-1, 3) dalapon at 16.8 kg ai'ha- 4) sulfometuron, 2-[[[[(4,6-dimethyl-2-pyrimidinyl)amino]- carbonyl]amino]sulfonyl]benzoic acid, at 1.1 kg ai'ha- and 5) untreated control. Initial herbicide applications were made to cogongrass foliage that was 60-90 cm tall, and sequential treatments were made to regrowth that was 60-120 cm tall. Treatments were applied using a C02-pressurized -1 backpack boom-sprayer calibrated to deliver 280 L'ha- at 210 kPa. A three by five factorial design (initial treat- ments of imazapyr, glyphosate, and sulfometuron and sequen- tial treatments of imazapyr, glyphosate, dalapon, sulfomet- uron, or untreated) was utilized with three replications on 1.8 m x 4.6 m plots. The first experiment was located in Area A of the infestation and the second experiment in Area B. Initial and sequential applications for the first experi- ment were made on July 9, 1985, and September 19, 1986, respectively. In the second experiment, initial and sequen- tial applications were made on September 19, 1986, and October 8, 1987, respectively. In January, 1986, (first experiment) and January, 1987, (second experiment) plots were mowed to ground level. This allowed the sequential applica- tions to be made to foliage regrowth without interference from persistent dead foliage. In January, 1987, (first experiment) and January, 1988, (second experiment) a 1.8 m swath was mowed through the plots. From this area foliage regrowth within a 0.25 m2 area was harvested. Rhizome biomass was harvested from within these quadrants by taking .a single soil core in the same manner as that described in the rhizome biomass sampling study. Untreated control plots were also harvested and used to calculate inhibition of foliage and rhizome dry weight. These harvests were made on September 17, 1987, (first experiment) and June 23, 1988 (second experiment). All samples were dried at 60 C for 72 hrs. This substantial regrowth period was necessary in order to provide an accurate assessment of the long-term effect of these herbicide programs on cogongrass. Analysis of variance was used to test for main factor (initial and sequential herbicide treatments and experiment) effects and the interactions. Least significant difference (LSD) procedure was used to separate treatment means. Tank-mix combinations of imazapyr and glyphosate. To determine the influence of tank-mixes of imazapyr and glyphosate on cogongrass control, two experiments were conducted during a three-year period. Single applications were made to cogongrass foliage on July 9, 1985, (first experiment Area A) and on September 16, 1986, (second experiment Area B). Applications were made using the CO2-pressurized boom system previously described. The absolute amount of each herbicide added to the tank-mix was computed by designating 1.1 and 3.4 kg ai'ha-1 of imazapyr and glyphosate, respectively, as the 100% dosage. Using these rates imazapyr + glyphosate mixtures of 0 + 100, 25 + 75, 50 + 50, 75 + 25, and 100 + 0% were formulated. The experimental design utilized was a randomized complete block with three replications on 1.8 m x 4.6 m plots. Plots were mowed in January, 1986, and January, 1987, for the first and second study, respectively. Both foliage regrowth in a 0.25m2 area and a rhizome-soil core from within this area were harvested on September 17, 1987, for the first experi- ment. For the second experiment, only foliage regrowth was harvested on June 23, 1988. Untreated control plots were harvested for both studies and this data was used to compute inhibition of foliage and rhizome dry weight. Analysis of variance was used to test for treatment and experimental effects and for the interaction. Least signif- icant difference (LSD) procedure was used to separate treatment means. Comparison of low and conventional volume applications of imazapyr and glyphosate. To determine the influence of carrier volume on the efficacy of imazapyr and glyphosate, two experiments were conducted during a three-year period. Imazapyr and glyphosate were applied in 46 and 234 L'ha-1. Glyphosate was applied at 1.7 and 3.4 kg ai'ha-1 and imazapyr was applied at 0.4 and 0.8 kg ai*ha-1. In the first experiment, applications were made using a tractor-mounted boom sprayer travelling at 6.4 km'hr-1. To apply 46 L'ha-I the boom was equipped with 11001LP2 flat fan nozzles calibrated at 124 kPa pressure. To apply 234 L'ha-1 the boom was equipped with 110052 flat fan nozzles calibrated at 276 kPa. In the second experiment a C02-pressurized backpack boom sprayer was .utilized. Using this system, 46 L'ha-1 was applied by using TX-62 hollow cone nozzle calibrated at 207 kPa and travelling at 8 km'hr-1 while, 234 L'ha-1 was applied by using 110032 flat fan nozzles calibrated at 221 kPa and travelling at 4.8 km'hr-1. Treatments were applied on July 9, 1985, (first experi- ment) and September 16, 1986, (second experiment). The experimental design utilized was a 23 factorial with three replications. Plots were 3.0 x 6.1 m and 1.8 x 4.6 m in the first and second experiment, respectively. Both studies were conducted in Area B of the infestation and cogongrass was 60-90 cm tall at the time of application. In January, 1987, and January, 1988, plots were mowed. Regrowth foliage and soil-rhizome cores were harvested, as previously described, on September 17, 1987, and June 23, 1988, for the first and second experiment, respectively. Untreated control plots were harvested for use in computing inhibition values. Data were subjected to analysis of variance to test for significance of herbicide, herbicide rate, carrier volume, and experimental effects and the interactions. Since there 2Spraying Systems Co., North Avenue, Wheaton, IL 60188. was no experimental interaction, data for the two experiments experiments were pooled. Treatment means were separated using least significant difference (LSD) procedure after appropriate sorting. Ropewick applications of imazapyr and glyphosate. To determine the influence of ropewick-applied imazapyr and glyphosate on cogongrass control, two experiments were conducted during a three-year period. The effects of concentrations of imazapyr and glyphosate (33 and 50% v/v) applied once or twice (in opposite directions) within a plot were evaluated. The ropewick apparatus utilized was 2.1 m long with a reservior capacity of 17.3 L. Two rows of half-overlapping Pistachio3 rope (nine sections per row each 20 cm long) were attached using rubber bushings within a screw-cap compression fitting, the body of which was glued to the PVC reservior creating 1.8 m of wicking surface. Two sets of ropes were constructed, one for each herbicide. Applications were made by two people carrying the ropewick through the plot at 4.8 km*h-1 with the wicking surface held approximately 150 below perpendicular to the leaf surface. The experimental design utilized was a 23 factorial with three replications in the first experiment and four replications in the second experiment. The plot size in both experiments was 1.8 x 4.6 m. Both experiments were Gulf Rope and Cordage Inc., P.O. Box 5516, Mobile, AL 36605. conducted in Area A of the infestation. Applications of both herbicides were made on July 9, 1985, for the first experiment. In the second experiment, glyphosate was applied October 2, 1986, and imazapyr was applied October 3, 1986. In both experiments, cogongrass was 60-90 cm tall at the time of application. In January, 1987, and January, 1988, plots were mowed. Regrowth foliage was harvested, as previously described, on September 17, 1987, and June 23, 1988, for the first and second experiment, respectively. Untreated control plots were harvested for use in computing inhibition values. Data were subjected to analysis of variance to test for significance of herbicide, herbicide concentration, number of passes, and experimental effects and the interactions. Since there was no experimental interaction, data for the two experiments were pooled. Treatment means were separated using least significant difference (LSD) procedure after appropriate sorting. Imazapyr longevity bioassay. To determine the persistence of imazapyr under Florida conditions and the effects of residual imazapyr on four seedling grass species frequently used for roadside cover, an imazapyr rate titration study and a field soil bioassay study were conducted. Of the four species used, two (Italian ryegrass, Lolium multiflorum Lam., and browntop millet, Brachiaria ramosa (L.)Stapf.) are annuals used to establish rapid cover in reestablishment areas and two (Pensacola bahiagrass, Paspalum notatum var. saurae Parodi, and common bermudagrass) are perenniating species established from seed. The mixture of the four species is dependent on the time of year and the location. Greenhouse experiments were conducted to develop stan- dard curves for inhibition of dry weight of each species. Field soil (containing cogongrass leaf and rhizome matter) from the Chiefland site was placed in 1.0 L cups. Imazapyr was surface-applied at 0, 0.001, 0.01, 0.1, 1.0, and 10.0 kg ai'ha-1 using a CO2-pressurized micro-applicator4 calibrated to deliver 376 L'ha-1 at 276 kPa. A 0.5 cm layer of silica sand was put on the soil surface and 30 seeds of each species (1 species per pot) were planted and covered with and additional 1.0 cm of sand. Plants were watered daily and maintained in a greenhouse with the following environ- mental conditions; day temperature 35 5 C, night tempera- ture 25 5 C, light intensity at noon 1000 pE'm"s- , and a 16-hr light/8-hr dark photoperiod. At thirty days after treatment, the plants were cut at the soil surface and dried at 100 C for 24 hours. Each treatment was replicated four times and the experiment was repeated. Regression analysis was used to determine the best- 2 fitting relationship (based on F-values, r values, and significance of the equation-parameter estimates) between imazapyr rate and inhibition of dry weight (IDW) for each R & D Sprayers, Opelousas, LA 70570. species. Both log-transformed and nontransformed herbicide rates and plant responses were used to determine the best model. The following models were chosen based on this criteria: Equation [1]: Annual ryegrass (AR): Logl0JIDWI = 1.74 + 0.22 (Log10Rate) r2 = 0.96 Pensacola bahiagrass (PB): IDW = -36.1 54.4 (Logl0Rate) 11.1 (Logl0Rate)2 r2 = 0.99 Common bermudagrass (CB): IDW = -69.6 28.2 (Logl0Rate) r2 = 0.94 Browntop millet (BTM): IDW = -5.8 71.1 (Rate) + 6.2 (Rate) r = 0.99 These data were utilized to: 1) determine that browntop millet (based upon the model significance, its intermediate response to imazapyr, and its consistently high germination) would be used in the field soil bioassay, and 2) develop relational models between the response of browntop millet with the other species. These relationships are: Equation [2]: IDWAR = 3.6 + 0.98 (IDWBTM) r2 = 0.85 IDWpB = 34.0 + 1.25 (IbWBTM) r2 = 0.97 IDWCB = -5.9 + 0.9 (IDWBTM) r2 = 0.74 To determine the dissipation rate of imazapyr under Florida conditions following treatment of cogongrass, a 0.1 ha block was treated with 1.1 kg ai'ha- imazapyr on September 16, 1986. Application was made using a CO2 pressurized backpack boom-sprayer calibrated to deliver 235 L'ha- at 221 kPa. From this area four PVC columns (16 cm in surface area and 23 cm long) were extracted at 2, 4, 5, 6, 8, 9, 10, 11, 12 and 14 months after treatment, sealed and stored at 0 C until all samples had been retrieved. Samples were allowed to thaw for 24 hrs at which time the top 15 cm was removed intact and placed vertically in 1.0 L cups. The fact that imazapyr is more or less immobile in soil would indicate that the active ingredient would be in the top portion of these soil columns (American Cyanamide, 1983). Four untreated pots were included to calculate IDW. Thirty browntop millet seeds were planted and covered with 1.0 cm sand. These pots were maintained in a greenhouse with environmental conditions similar to those described for the titration study. Thirty days after planting above-ground biomass was harvested and dried. The response (IDW) data for browntop millet bioassay were used to develop a regression model for predicting IDW of browntop millet at a given time after application. The following equation describes that relationship: Equation [3]: IDWBTM = -91.2 + 7.7 (Months after treatment) r2 = 0.80 From equations [2] a desired response (IDW) is set (in this case, the 0, 25, and 50% IDW) for each of the three species and the corresponding repsonse (IDW) for browntop millet is determined. This response is used in equation [3] to determine the TO (time required to allow imazapyr to dissipate before the IDW of a given species equals 0), T25, and T50 values for each of the species. Results and Discussion Cogongrass Rhizome Biomass Sampling. Table 3.1 presents the rhizome biomass harvested from the two areas of the infestation used for the control studies. There was a significant interaction between area and time of sampling. Overall, Area A, the older section of the infestation, contained a greater amount of rhizome biomass than did Area B. Both areas tended to be cyclic in terms of rhizome production. The greatest quantity, in both areas, was harvested during the late fall in both 1986 and 1987. The least amount of rhizome was present during the late spring and early summer. This phenomenon has been documented for other perennial grasses (Johnson and Buchholtz, 1962; Horowitz, 1972). It has also been shown that during regrowth periods (i.e., in the spring or following mowing) there is a quantitative shift from carbo- hydrate storage to remobilization for leaf production (Potter, et al., 1986; Arny, 1932). There is also a shift from rhizome biomass production during periods of rapid leaf Table 3.1. Cogongrass rhizome biomass collected in Chiefland, FL from 1986 to 1988. Month Year AREA A AREA B LSD October November December February March May June July August September October November January February March April 1986 1986 1986 1987 1987 1987 1987 1987 1987 1987 1987 1987 1988 1988 1988 1988 - - kg'ha- 7355 (387)2 6898 (521) 6734 (318) 7465 (707) 7355 (631) 4623 (537) 7218 (396) 7300 (66) 7163 (275) 8031 (544) 8598 (495) 8159 (258) 7076 (428) 7044 (420) 7181 (333) 6012 (343) 1 Significance levels for rhizome biomass differences between areas within a sampling time using LSD are 0.01 (***), 0.05 (**), and nonsignificant (NS). 2 Mean rhizome dry weight of 6 replicates/area computed from 4170 cubic cm soil cores followed by the standard error of the mean. NS *** 5592 7328 4797 3984 4806 2960 3682 4806 4185 4020 6789 4879 4061 4651 5217 4048 (360) (443) (398) (305) (212) (181) (214) (272) (398) (500) (504) (197) (97) (198) (303) (135) *** area expansion which then reverses after sufficient leaf area has been produced. The efficacy of translocated herbicides (e.g., glyphosate) has been shown to be affected by the stage of development of perennial species (Atkinson, 1985). The best long-term control is attained when perennial grasses are: 1) utilizing rhizome for expansion purposes, thereby not harbouring large numbers of metabolically quiescent nodes and 2) transporting photosynthate more to the perenniating organs than to expanding leaf tissue, which would occur during the latter part of the growing season. Relating these factors and the characteristics of the infestation will be used to explain year-to-year interactions that occurred in the following control studies. Sequential Herbicide Application Programs. There was a significant three-way interaction between initial herbicide treatment, sequential herbicide treatment, and experiment. Therefore, experiments were analyzed separately. Tables 3.2 and 3.3 present the cogongrasss leaf and rhizome inhibition data for the first and second experi- ment, respectively. Overall, control was superior in the second experiment. Two factors may have contributed to this. First, the two areas within the infestation were shown to be significantly different in terms of rhizome biomass, with Area A (first experiment) containing a greater quantity (Table 3.1). There was also a greater amount of leaf material present in Table 3.2. The effect of sequential herbicide treatments on cogongrass foliage and rhizome dry weight (Experiment 1 - 1985 through 1987 Area A). Initial Application (kg ai'ha1) Sequential Imazapyr Glyphosate Sulfometuron Application (0.8) (3.4) (1.1) (kg ai'ha -1 FDW1 RDW2 FDW RDW FDW RDW ----% Inhibition -- - Imazapyr (0.8) Glyphosate (3.4) Dalapon (16.8) Sulfometuron (1.1) 100 82 64 55 100 -37 88 -42 -17 -25 -48 28 -71 Untreated LSD(0 1)Initial 20 30 LSD(0.1)Sequential 25 Cogongrass foliage dry weight (regrowth) harvested from 0.25 square m quadrants at 12 months after sequential herbicide application. 2 Cogongrass rhizome dry weight harvested from 4170 cubic cm soil cores at 12 months after sequential herbicide application. Inhibition values computed using untreated plots which contained 53.5 and 3.9 g of foliage and rhizome, respectively. Table 3.3. The effect of sequential herbicide treatments on cogongrass foliage and rhizome dry weight (Experiment 2 - 1986 through 1988 Area B). Initial Application (kg ai'ha1) Sequential Imazapyr Glyphosate Sulfometuron Application (0.8) (3.4) (1.1) (kg ai'ha-1) FDW1 RDW2 FDW RDW FDW RDW - % Inhibition3 Imazapyr 98 68 100 78 89 68 (0.8) Glyphosate 94 59 98 29 100 15 (3.4) Dalapon 84 55 100 60 43 -49 (16.8) Sulfometuron 68 78 91 64 12 -29 (1.1) Untreated 96 66 83 69 48 28 ----- ---- ---- -- -- -- --m - ---- -- -- -- LSD(0.1)Initial 13 13 LSD(0.1)Sequential 16 16 1Cogongrass foliage dry weight (regrowth) harvested from 0.25 square m quadrants at 10 months after sequential herbicide application. 2Cogongrass rhizome dry weight harvested from 4170 cubic cm soil cores at 10 months after sequential herbicide application. 3Inhibition values computed using untreated plots which contained 10.8 and 3.3 g of foliage and rhizome, respectively. the untreated controls of Area A (first experiment). There- fore, a denser, more established stand would be more diffi- cult to control. Second, the weather patterns occurring during the two experiments were different (Figure 3.1). During the first experiment, a dry summer 1985 was followed by normal precipation through 1986. In the second experiment, there was adequate rainfall through the summer and fall 1986, but was followed by a below normal rainfall through the summer and fall 1987 except during the month prior to the sequential applications. Weather patterns could have adversely affected herbicide efficacy in the first experiment. First, the effect of the initial applications may have been reduced by drought-stressed cogongrass which was demonstrated by the relatively poor control obtained from single applications. Second, adequate rainfall following the sequential application favored cogongrass regrowth (which is evident from the 53.5 g of foliage regrowth in the untreated control plots). In the second experiment, initial applications to vigorously growing cogongrass were quite efficacious. This is demonstrated by the excellent control obtained from single applications of imazapyr and glyphosate. Though the sequential applications were made during an overall dry period, the rainfall during September and early October was adequate to maintain cogongrass growth. After this time, rainfall was low which resulted in suppressed cogongrass growth. This evident from the 10.8 g of foliage regrowth in Gainesville, Florida Weekly Precipitation CM 1986 12.71 1 U, 7.6 - 5.1 -o. -- 2.5 , Jan Feb Mar Apr May Jun Jul Aug Sept Oct leyv Dec Total Precipitation 132.9 cm Gainesville, Florida Weekly Precipitation cm 1987 Jan Feb Mar Apr May Jun Sl Aug SCp OC Nov Dec Total Precipitation 115.5 cm Figure 3.1. Weekly precipitation in Gainesville, FL, in 1986 and 1987 (Adapted from McCloud and Hill, 1987 and McCloud and Hill, 1988). ~rr r the untreated control plots. The combination of a less dense stand (Area B) and the efficacious initial herbicide treatments followed by less than favorable conditions for regrowth following the sequential applications resulted in better overall control in this experiment. In both experiments two compounds, imazapyr and glyphosate, were most effective. However, single applica- tions provided adequate control only in the second experi- ment. The response of cogongrass has been shown to be impacted by the overall soil-water status. Arif (1979) found that cogongrass control acheived with glyphosate was greater during the dry season (unless there was serious water stress) than during the rainy months. This may explain the improved glyphosate activity in the second experiment. In the first experiment, single applications of these herbicides had no effect on rhizome growth and consequently poor control of foliage regrowth was obtained. The use of dalapon as a sequential treatment provided some additional control in the first experiment when glyphosate or sulfometuron were used as an initial treatment, but none when imazapyr was used. In the second experiment, dalapon provided no additional control when used as a sequential treatment. Sulfometuron apparently had no adverse effect on cogongrass. In fact, there was significant stimulation of rhizome growth in both experiments. This may have occurred not because of a direct chemical effect on cogongrass, but from the control of broadleaf species that may have been competing with cogongrass. These experiments indicated that near 100% control of cogongrass on a long-term basis may be consistently achieved, regardless of extraneous field and climatic factors, only with an initial and sequential herbicide .program based on imazapyr and glyphosate. Tank-mix combinations of imazapyr and glyphosate. Table 3.4 presents cogongrass control data obtained from tank-mixes of imazapyr and glyphosate. There was a significant interaction between treatment and experiment, therefore experiments were analyzed separately. Overall, control of foliage regrowth was superior in the second experiment. This occurred possibly for the same reasons as those proposed for the sequential herbicide programs study. In the first experiment no clear trend is apparent for the level of control achieved and the herbicide components. This indicates that in-tank antagonistic interactions did not occur. In the second experiment control of foliage regrowth was greater from tank-mixes with 50% or more imazapyr. The results of the two experiments may indicate that these two herbicides were additive in their effect on cogongrass, with imazapyr being more active. These data also indicate that the use of imazapyr + glyphosate ratio tank-mixes did not alleviate the need for multiple applications to consistently achieve long-term control. Table 3.4. The effect of imazapyr and glyphosate tank-mixes on cogongrass foliage and rhizome dry weight. Tank-mix Treatments Imazapyr + Glyphosate Experiment 1 1985-87 (Area A) FDW1 RDW2 Experiment 2 1986-88 (Area B) FDW - kg ai'ha-1 - kg a i'ha 1.1 0.8 0.6 0.3 0.8 1.7 2.5 3.4 3nhibition - % Inhibition 60 36 82 - - LSD(0.1) 33 40 9 Cogongrass foliage dry weight (regrowth) harvested from 0.25 square m quadrants at 26 and 21 months after treatment for the first and second experiment, respectively. 2 Cogongrass rhizome dry weight harvested from 4170 cubic cm soil cores at 26 and 21 months after treatment for the first and second experiment, respectively. Inhibition values computed using untreated plots which contained 81.0 and 13.5 g of foliate and rhizome, respectively in experiment 1 and 17.3 g of foliage in experiment 2. - - Comparison of low and conventional volume applications of imazapyr and glyphosate. There was not a significant interaction between experi- ment and the other factors (volume, rate and herbicide). However, experiments were significantly different (P < 0.05). The differential rainfall discussed previously could have caused the overall increased control acheived in the second experiment, but the efficacy of the treatments was not differentially affected. There was a significant two-way interaction between herbicides (imazapyr or glyphosate) and carrier volume (46 or 234 L'ha1 ) for the inhibition of cogongrass foliage regrowth. These data are presented in Table 3.5. Imazapyr provided significantly greater control than -1 glyphosate at both rates when applied in 234 L'ha1 water. For both herbicides, rate did not influence control. However, in the case of glyphosate, the half rate was not significantly different from the untreated control on an absolute foliage dry weight basis, but the higher rate did cause a significant reduction in foliage dry weight as compared to the untreated control plots. In either case, control obtained using glyphosate was poor. Imazapyr provided a greater level of control when applied in 234 L'ha-1 volume application than when applied in 46 L'ha-. This may indicate that in dense stands of cogongrass the greater foliage coverage attained with this volume is more important than the enhanced activity that has been reported 43 Table 3.5. The effect of carrier volume and imazapyr and glyphosate rate on cogongrass foliage dry weight. Imazapyr Carrier Volume (L'ha-1) (L'ha ) Glyphosate -i kg ai'ha1 0.4 0.8 1.7 3.4 1 -% Inhibition 333 732,3 234 483 712,3 122 Inhibition values computed using untreated plots which contained 24.8 g of foliage regrowth/0.25 square m quadrant. 2Means within a given volume and herbicide rate indicates that herbicides are significantly different using LSD(0.1). Means with a given herbicide and herbicide rate indicates that volumes are significantly different using LSD(0.1). -102 to occur with ultra-low volume applications. This study again points out the necessity of multiple applications to achieve long-term cogongrass control. Ropewick applications of imazapyr and glyphosate. There was no significant interaction between experiments and the tested factors (herbicide, concentration, and number of passes). However, experiments were signifi- cantly different (P < 0.05). There was a significant three- way interaction between herbicides (imazapyr or glyphosate), herbicide concentration (33 or 50% v/v), and number of passes for the inhibition of both cogongrass foliage regrowth and rhizome dry weight. These data are presented in Table 3.6. Control of cogongrass rhizomes with glyphosate was unaffected by the concentration or the number of passes made. This may have occurred due to the solutions being too concentrated. Boerboom and Wyse (1988) speculated that the reason for poor control of Canada thistle [Cirsium arvense (L.)Scopoli] achieved using ropewick-applied glyphosate was that the concentrations being applied were greater than the optimum level needed to maximize translocation to the roots. In essence, by overdosing the leaf tissue, glyphosate provides more contact activity to the detriment of systemic activity. Therefore, the use of glyphosate at concentra- tions greater than 33% would be economically unsound. In fact, the use of concentrations lower than 33% may provide as much, if not more, control of cogongrass. Table 3.6. The effect of ropewick applications of imazapyr and glyphosate on cogongrass foliage and rhizome dry weight. Imazapyr Glyphosate .% Solution # Passes FDW RDW2 FDW RDW - % Inhibition3 33 1 67 -175 55 17 33 2 68 115 65 31 50 1 764 245,6 384 28 50 2 784 455,6 604 27 Cogongrass foliage dry weight (regrowth) harvested from 0.25 square m quadrants. Cogongrass rhizome dry weight harvested from 4170 cubic cm soil cores. Inhibition values computed using untreated plots which contained 43.6 and 5.8 g of foliage and rhizome, respectively. Means within a given % solution and number of passes indicates that herbicides are significantly different using LSD(0.1) 5Means within a given herbicide and number of passes indicates that % solutions are significantly different using LSD(0.1). 6Means within a given herbicide and % solution indicates that number of passes are significantly different using LSD(0.1) Imazapyr at the 50% concentration provided better control of cogongrass foliage regrowth than did glyphosate at this concentration. Unlike glyphosate, imazapyr efficacy on rhizome growth was significantly affected by concentration .and number of passes. The highest level of control was achieved at 50% v/v applied twice. This would seem to indicate that imazapyr adsorption and/or translocation is not as sensitive to high concentrations as glyphosate. The use of two passes at the 50% concentration provided the highest level of control of any of the treatments. Although 100% control was not acheived from either herbicide, multiple ropewick applications of glyphosate or imazapyr may be a viable alternative for cogongrass control in situations where broadcast applications are not desirable. Imazapyr longevity bioassay. Data from the imazapyr bioassay are presented in Table 3.7. The most tolerant of the four species tested was bahiagrass with an 150 rate of 1.40 kg ai'ha-1. Common bermudagrass and browntop millet were intermediate in their response and annual ryegrass was the most sensitive. The half-life of imazapyr in sandy soils of Florida was found to be approximately eight months. Research has shown that under temperate conditions imazapyr persists from three months to one year, while under tropical conditions biological activity persists for three to five months (American Cyanamide, 1983). The fact that the applications were made late in the growing Table 3.7. The influence of imazapyr dissipation on the inhibition of dry weight (IDW) response of four grass species. 1 50 Species kg ai'ha 2 T T T 0 T25 T50 Months After Treatment Annual ryegrass (AR) 0.65 11.4 8.0 4.7 Pensacola bahiagrass (PB) 1.40 8.3 5.7 3.1 Common bermudagrass (CB) 0.70 12.7 9.1 5.5 Browntop millet (BTM) 0.70 11.8 8.6 5.4 1Imazapyr rate required to cause 50% IDW. 2 Dissipation time (months) required to cause 0, 25, and 50% IDW of each species. season (September 16) should be taken into consideration as degradation by soil microbes would have been reduced during the cold, dry winter months. However, research has shown that the major route of imazapyr degradation is via photo- lysis (American Cyanamide, 1983). Photolysis of imazapyr in the soil would probably be reduced by the dense and persistent cogongrass foliage during the winter months. Therefore, the conditions under which this study was conducted would have tended to increase the persistence of imazapyr. Therefore, if imazapyr were to be used for cogongrass control on highway rights-of-way, reestablishment of this area with one of the four grass species tested would have to be delayed until the late spring following the fall applica- tion. The amount of imazapyr remaining at this time would possibly be reduced by the removal of cogongrass foliage after the first frost, thereby increasing imazapyr photolytic breakdown. However, this would also reduce the amount of imazapyr available for control of cogongrass regrowth. The use of multiple applications (which may be necessary to achieve 100% control) should also be considered. The use of imazapyr as an initial treatment, followed by the use of glyphosate or dalapon as a sequential spot treatment to cogongrass regrowth following turf establishment may provide the highest level of control with the shortest reestablish- ment period. CHAPTER 4 THE INFLUENCE OF MECHANICAL AND CHEMICAL INPUTS ON COGONGRASS [Imperata cylindrica (L.)Beauv.] CONTROL. Introduction In the tropical South Pacific and Southeast Asia cogongrass (known there as alang-alang or lalang-alang) covers several million hectares (Holm, et al., 1977). These developing countries, whose economies are based largely on agriculture, do not have the capital, expertise, or tech- nology necessary to effectively control cogongrass on a large-scale basis (Holm, 1969). Therefore, localized efforts relying on slash-and-burn, grazing, and tillage are the most widely used control methods. Controlled studies on the effectiveness of these techniques have indicated that; 1) shifting agriculture may provide short-term control (long enough to produce one or two.crops-prior to reinfestation), but in the long-term, cogongrass populations are increased, 2) repeated burning followed by grazing will marginally support animal production, but provides little in the way of control, and 3)- intensive tillage has repeatedly shown to be an effective method off bogongrass management, however the C I **<; availability of ilaplmefts, soil type climatic conditions, Said. tera.n limit it use. in southeast Asia (Falvey, et al., ':Ip~l, fl8xiea j 2,833; Prorasool, 1184). ,* .. ... .. .. .. .. ... W.FF; TE: .~: :, ." ..... J .: ....= The recent introduction of several new (and more . efficacious) herbicides in these areas have demonstrated:.the utility of chemical control (Arif, 1979; Bacon, 1986). However, application technology and expertise in the proper, and most effective use of these compounds is limited. Much of the chemical control conducted was (and still is) with dalapon, 2,2-dichloropropanoic acid, paraquat, 1,1'-dimethyl- 4,4'-bipyridinium ion, and glyphosate, N-(phosphonomethyl)- glycine applied with knapsack sprayers or antiquated high- volume spray systems (Hartley; 1949, Keepings and Matheson; 1949; Sandanam and Jayasinghe, 1977; SEAWIC, 1987; Seth, 1970). Multiple applications are expensive, but a require- ment to acheive any significant control and then long-tend control is seldom realized. The integration df til age :and herbicide treatments has been used in the United States :for the control of several perennial weeds. Rhizome johnsongrass [Sorghum halepense (L.)Pers. # SOP A] control programs based on dishing after corn (Zea mays L.) harvest, fellow applications of glyphosate to re growth, and tri lra li F 2,6-dinitto-,N-- ": *" .. 4 A.* dipropyl-4-(tri-fluoromethyl) benzenaWi nr treat ent'at twice .. .. : .. : the normal use rate :: io to soybean. [Gi a;. (L.)Mer;.] 4 4 ....F'" ". ," : i,":""..a. ;. :.::. i .. ..: planting provides excellent a ontol (7LA1O~ 1.87; lonanto, 1988)., Programs ustni il Ie e either _91r following . ..' ,* .. .. ..,...:.. .. .. .. . Letters follow ing thseb a vd code from the oapo.dite, 4Psdst t N .. 2. Availabib frin SSA, 3C9WMw J. 0.201& LX 61820. ." "" S ... .. .... ." A;, 2pr. :t':.j .J.. .."t rr-'.. i'. ... 2. "" "" :" """ ..~ a.'n.: .': :,"i glyphosate applications have been utilized for control of other perennial weed species (Baird and Begeman, 1972; Sandberg and Meggitt, 1977). Mowing has been shown to have only short-term effects Son perennial grass growth (Beard, 1973). However mowing is utilized extensively in Florida for right-of-way and pasture management. If chemical control is to be effectively interfaced with mowing then the combined effects of mowing and herbicides needs to be investigated. The objectives of these studies were to determine the influence of mowing and disking on the efficacy of selected herbicides. Studies were also conducted to determine the influence of light and photoperiod on shoot initation in cogongrass rhizome segments and to determine the influence of cogongrass stage of development and defoliation on the efficacy of glyphosate. Materials and Methods Integration of mowing and disking with herbicide treatments. To determine the interaction of mowing or disking with herbicide treatments on long-term cogongrass control, two studies were conducted from 1985 to 1988 at Chiefland, Levy County1. Florida. Studies were located on an eight hectare cogongrass infestation that had became established during the: mid-l970. The soil type was a Sparr fine sand (loamy, *siliceo per thermic, Grssaarenic Palenulduts),. .' :. :* 1:. *c ; .:.. ;;.. ::1:ib ......... .. :.ii: ." ". .... *. * ,A '.. 1 ... iii.1 , :.4 :" The experimental design was a split-plot with the mechanical treatment (mowing or disking) being the main plot and herbicide treatment as the subplot. Three levels of mowing or disking were used: 1) no mechanical treatment, 2) mowing or disking prior to herbicide treatment, and 3) mowing or disking prior to and subsequent to herbicide treatment. Five herbicide treatments were applied: 1) untreated control, 2) imazapyr, () -2-[4,5-dihydro-4-methyl- 4-(1-methylethyl)-5-oxo-lH-imidazole-2-yl]-3-pyridinecar- boxylic acid, applied at 0.8 kg ai'ha-1, 3) glyphosate applied at 3.4 kg ai'ha-1, 4) dalapon applied at 16.8 kg ai'ha-1, and 5) sulfometuron, 2-[[[[(4,6-dimethyl-2-pyrimi- dyl)amino]carbonyl]amino]sulfonyl]-2-thiophenecarboxylic -1 acid, at 1.1 kg ai'ha1. In 1985 (first experiments) each mechanical-chemical treatment was replicated three times on 1.8 m x 3.7 m plots. In 1986 (second experiments) each treatment was replicated four times on 1.8 m x 4.6 m plots. Mowing treatments were made using a tractor-mounted horizontal impact-mower2 which was set to remove approxi- mately the top half of the foliage then set to cut ground level. The disking treatments were made by first mowing the blocks followed by repeated disking until all vegetation was destroyed and the soil was tilled to a depth of 10-15 cm. Due to the dense rhizome layer, eight to ten passes per block were required. Herbicide applications were made using 2Bush Hog Manufacturing, Inc., Selma, AL. a CO2-pressurized backpack sprayer calibrated to deliver 280 -1 L'ha1 at 207 kPa. On July 5, 1985, initial mowing and disking treatments for the first experiments were implemented. Herbicide treatments were applied on November 15, 1985, to cogongrass that was 30 cm tall in the mechanically treated blocks and 100 cm tall in the undisturbed blocks. Following the first freeze in January, 1986, all blocks were mowed to ground level. On July 29, 1986, the sequential mowing and disking treatments were implemented. For the second experiments in each study, initial mowing and disking treatments were imposed on September 16, 1986, followed by herbicide applications on November 14, 1986. In January, 1987, all blocks were mowed to ground level. Sequential mechanical treatments were completed on June 25, 1987 (Table 4.1). 2 Cogongrass control was determined by harvest of 0.25 m quadrants from the center of each subplot and a soil-rhizome core was extracted from this quadrant using a bucket auger 182 cm2 in surface area to a depth of 23 cm. Rhizomes which were not decayed were removed from these samples. These harvests were made on September 17, 1987, (first experiments) and on June 23, 1988 (second experiments). Foliage and rhizome samples were dried at 60 C for 72 hrs. Foliage and rhizome dry weights were converted to percent inhibition based on the untreated controls. Data were subjected to analysis of variance (Helwig and Council, 1982) to test for single factor effects and interactions. In both the mowing -.. "., .'-:. .... :: ,. 5 ':: '';:" 4. Table 4.1. Timing of mechanical and chemical treatments and harvests performed for evaluation of integrated cogongrass control experiments. Operation SPerformed Experiment 1 1985-1987 Initial Mowing and Disking Herbicide Applications Mowing for Removal of Dead Foliage Sequential Mowing and Disking Foliage Regrowth and Rhizome Harvest 7/5/1985 11/15/1985 1/15/1986 7/29/1986. 9/17/1987 Experiment 2 1986-1988 9/16/1986 11/14/1986 1/15/1987 6/25/1987 6/23/1988 I . F- * 1*. '* *'* ** :... .... *' '> ..* :. .,." . I ". ., T- : .. ; ;, ,: .: ... 5. ** : : .... : .* : :- ..; : .:- y . .. .: .. ". .* 1 J ". :. .. .i ":... :. ; "," "::" : ... . "',i .. .:,. ; ,. ... . ; . .:."*k . "ii : ..s -, I E. .. iPt :.." ,E i.*; ; ::;^ : .*. ** :-. -t . *" "Ii .- ; *.. ; *'*.: , S:... .. ... .:. :. . S:. .. .. ;...: ..... :. ". .. '" :.,. .'.." ; " i :: ".. .*' .: ." H. : .. ... "i : :.... '." :. '. :. ** .. *" ". .. .:.' :.. *** : *:.* : ..' ..." : : :... ..*' ..^ ^ A S .,. .. . .; .. ~ ~ ~ ~ .. ':E .. ." :. ..:. .; :!., .. . .* * : .; 'I :: : : ", / ::; : :.. ; :: .'" : : .. : :iii .' .; : E .. -- :* 1 rr; .1. ;;- and disking studies there was a significant interaction (P < 0.05) between experiment and treatment, therefore experiments were analyzed separately. Treatment means were separated using least significant difference (LSD) procedure. Influence of photoperiod on shoot production in cogongrass rhizomes. Growth chamber experiments were conducted to determine the effect of photoperiod on shoot initiation in cogongrass rhizomes. Rhizomes, harvested from an infestation at the University of Florida Agronomy Farm in 1987, were washed, de-scaled, and cut into six-node sections which showed no evidence of injury (i.e., pathogen or insect damage). No attempt was made to differentiate sections from various positions on the rhizome. Five or six rhizomes sections were placed on filter paper3 in petri dishes, 2 ml of deionized water was added, and petri dishes were covered. Dark controls were prepared first and immediately wrapped in aluminum foil. This procedure took approximately ten minutes during which time the dark controls were exposed to light. Preliminary studies indicated that this amount of exposure was insufficient to initiate shoots (data not shown). Water was added to petri dishes during the light portion of the photoperiod on the fourth day after planting. Dark controls were watered at night in a darkened room. The conditions in the growth chambers were 30 2 C and 3Whatman Paper Ltd., Maidstone, England. fluorescent lights were used to obtain a light intensity of 200 pE'm-2s-. Two growth chambers were utilized for each study. The first study evaluated continuous light and a 16-hr light/8-hr dark photoperiod. The second study .evaluated a 9-hr light/15-hr dark photoperiod and a 8 + 1-hr light(8 hr with a 1 hr light period during the dark period)/15-hr dark photoperiod. The third study evaluated a 16-hr light/8-hr dark photoperiod and a 9-hr light/15-hr dark photoperiod. Light treatments were replicated (5 petri dishes within the growth chamber). Dark controls were evaluated in each study and studies were repeated. One week after photoperiod treatments were imposed the following data were collected: number of productive segments, shoot number per segment, and total node number. Based on the dark controls only, all three studies (each conducted twice) were not significantly different (P > 0.05), thus all the data were combined and analyzed. Analysis of variance procedure was used to test for the significance of photoperiod on shoot initiation. Duncan's Multiple Range test was used to separate treatment means. The influence of cogongrass stage of development and defoli- ation on glyphosate efficacy. Greenhouse studies on cogongrass growth and development were conducted in 1987 and 1988 to characterize the effects of defoliation at various times after planting. Cogongrass was propagated in a growth chamber from rhizome segments that were harvested from an infestation in Gainesville, Florida in 1987. Rhizomes were cleaned, de-scaled, and cut into six-node segments that were free of insect or pathogen injury. Rhizomes were planted horizontally in trays filled with 2 cm of vermiculite and covered with an additional 2 cm of vermiculite. Rhizomes were watered daily. Growth chamber conditions were 30 2 C with a 16-hr light/8-hr dark -2. -1 photoperiod. Fluorescent lights provided 2001uE'm- s- Two weeks after planting, nodes that had produced a shoot 5 cm long were cut from the six-node segments. These shoots were planted into 2.8 L pots filled with 2.4 L of soil mix containing 50% fumigated4 Arredondo fine sand (loamy, siliceous, hyperthermic, Grossarenic Paledults) and 50% commercial potting mix5 on a volume basis. Three shoots were planted in each pot and fertilized with 100 ml of 5X Hoagland's solution one week after planting (Hoagland and Aronon, 1950). Plants were watered three times per week. Greenhouse conditions were as follows: day temperature 27 - 5 C, night temperature 15 5 C, and light intensity at noon of 1000 uE'm 2s-1 Four pots were harvested every three weeks after planting. At these harvest periods four other pots were defoliated to ground level. Defoliated pots were harvested three weeks later. Harvests were carried out by; clipping leaf tissue at ground level and measuring leaf Brom-o-gas (methyl bromide + chloropicrin), Great Lakes Chemical Corp., West Lafayette, IN 47906. Metro-mix 200, Gracewood Horticultural Products, Cambridge, MA 02140. area6, composite root-rhizome biomass was washed and dried at 100 C for 24 hrs and weighed, as was leaf tissue. Leaf and root-rhizome dry weight were converted to ratios based on total plant dry weight. This study was conducted twice. To determine the influence of stage of development and defoliation on glyphosate efficacy in cogongrass, greenhouse studies were conducted during 1988. Cogongrass propagation was identical to that used in the growth analysis study. Three stages of cogongrass were used in these experiments. Three-week-old plants (Stage A), six-week-old plants (Stage B), and plants that were defoliated at three weeks and allowed to regrow for three weeks (Stage C). To allow glyphosate applications to be made at the same time to all stages, cogongrass for stages B and C was planted first, then three weeks later cogongrass for stage A was planted (also at this time stage C plants were defoliated). Glypho- sate was applied three weeks after the second planting. Glyphosate solutions were prepared using technical glyphosate (96.5% acid7). The surfactant MON-08187 (poly- oxyethylene tallow amine) was added at 0.5% (v/v). 8 Solutions were applied using a single nozzle (TJ-60 8002E ) CO2-pressurized micro-applicator calibrated to deliver 860 Licor Model 3100 Area Meter, Licor, Inc., Lincoln, NE 7Monsanto Agri. Prod. Co., St. Louis, MO 63617. 8Spraying Systems Co., North Avenue, Wheaton, IL 60188. R & D Sprayers, Inc. Opelousas, LA 70570. L'ha- at 276 kPa. Glyphosate was applied at 0.1, 0.3, 0.6, and 1.1 kg (acid equivalent) ae'ha- (first experiment) and 0.01, 0.06, 0.1, 0.3, and 0.6 kg ae'ha-1 (second experiment). Untreated plants were also included in both experiments. Treatments were replicated four times within each cogongrass stage. At the time of application four pots from each stage were harvested. The data collected from these harvests were leaf area and dry weight and composite rhizome-root biomass. Samples were dried at 100 C for 24 hrs. At three weeks after application leaf tissue was cut off at ground level. Plants were allowed to regrow for three weeks, at which time leaf and root-rhizome tissue was harvested and dried. Data was converted to percent inhibition of dry weight change over the treatment period. Data were subjected to analysis of variance procedure to test for single factor effects and interactions. There were no significant inter- actions (P > 0.05) between experiments and treatments or experiments and cogongrass growth stages, therefore data from the two experiments were combined. There was a significant interaction (P < 0.05) between treatment and cogongrass stage. Therefore, regression models (% inhibition versus transformed glyphosate rate) were developed for each growth stage separately. The best models were chosen based on the significance level (P < 0.05) of the model, the equation components and the r2 value. Results and Discussion Mowing-herbicide studies. Data on the effects of mowing and herbicides on cogon- .grass are presented in Tables 4.2 and 4.3. The experiments are presented separately due to a significant (P < 0.05) experiment by treatment interaction. This interaction may have been a result of locational and/or environmental factors enhancing the effect of effiacious treatments while having little, if any, effect on treatments that provided little control. The area in which the first experiment was conducted contained five times more foliar biomass (95.8 g vs. 15.7 g'0.25 m-2) and two times more rhizome biomass -3 (15.0 g vs. 5.2 g'4170 cm-3) than the area in which the second experiment was conducted. On a yearly basis, rain- fall for 1986 was normal with no extended dry periods (McCloud and Hill, 1987 and McCloud and Hill, 1988). Rainfall in 1987 was 28 cm below normal with three extended periods of near-zero rainfall during the growing season. In the first experiment, there was a significant (P < 0.05) interaction between mowing and herbicide treatments on inhibition of foliage dry weight (Table 4.2). Mowing had a significant impact when no herbicides were applied or when dalapon or sulfometuron were applied. With these two herbicides, as the number of mowings increased control (% inhibition) of foliage dry weight (FDW) also increased. The use of mowing prior to sulfometuron treatment apparently Table 4.2. The effect of mowing and herbicides on cogon- grass foliage and rhizome dry weight (Experiment 1 1985 through 1987). Mowings1 0 1 2 Rate Rate-1 2 3 Herbicide kg ai'ha FDW RDW FDW RDW FDW RDW -- % Inhibition4 - - Imazapyr 0.8 94 44 89 61 95 42 Glyphosate 3.4 86 33 85 42 97 49 Dalapon 16.8 65 12 74 -3 90 24 Sulfometuron 1.1 11 -2 61 -2 79 52 Untreated 0 0 36 13 65 26 LSD(0.1)Mowings 7 18 LSD (0.1)Herbicides 9 24 1 0 mowings indicates herbicides applied to undisturbed cogongrass, 1 mowing indicates mowing prior to herbicide application, and 2 mowings indicates mowing prior and subsequent to herbicide application. 2Cogongrass regrowth foliage dry weight harvested from 0.25 square m quadrants at 14 months after second mowing operation. Cogongrass rhizome dry weight harvested from 4170 cubic cm soil cores at 14 months after second mowing operation. Inhibition values computed using unmowed and untreated plots which contained 95.8 and 15.0 g of foliage and rhizome, respectively. increased herbicide efficacy. This may have been due to the fact that sulfometuron has both foliar and soil activity (DuPont, 1982). By removing a large portion of the foliage prior to application, more herbicide may have directly .contacted the soil and was affecting emerging cogongrass shoots at that site. Though sulfometuron does exhibit post- emergence foliar activity in many species, it did not appear to affect cogongrass in this manner. Subsequent mowing had less of an affect than did prior mowing in conjunction with sulfometuron. Mowing in conjunction with glyphosate increased cogongrass control only when done both prior and subsequent to herbicide treatment. Mowing did not have a significant effect on the control of foliage regrowth by imazapyr. Control with imazapyr was excellent regardless of mowing treatment. Imazapyr, like sulfometuron, has both foliar and soil activity (American Cyanamide, 1983). How- ever, unlike sulfometuron, imazapyr exhibited significant foliar activity on cogongrass. Mowing had less impact on cogongrass rhizomes than foliage. Two mowings were required to significantly impact rhizome dry weight and then only when no herbicides were applied or dalapon or sulfometuron were applied. The use of mowing prior to and subsequent to imazapyr treatment signif- icantly reduced control as compared to mowing prior to application only. This may have occurred due to the fact that the major cause of imazapyr degradation is photolysis (American Cyanamide, 1983). By removing the cogongrass foliage, more light reached the soil increasing the degradation of imazapyr. In the second experiment (Table 4.3), herbicides and mowings significantly interacted to affect cogongrass foliage regrowth. When used without chemical treatments, mowing only inhibited foliage regrowth when done twice. This may have been caused by a depletion of carbohydrates that were needed to re-establish above-ground biomass. Inhibition of foliage dry weight with sulfometuron increased from -24% (growth stimulation) with no mowings to 77% with two mowings. The stimulation of foliage regrowth found when sulfometuron was used with no prior mowing may have been caused not by direct chemical stimulation of cogongrass, but by decreased competition from broadleaf species which sulfo- meturon controlled. This hypothesis is supported by the fact that sulfometuron with one or two mowings was not signifi- cantly different from mowing treatments alone. The use of mowing with dalapon gave greater control than dalapon alone or two mowings alone. When cogongrass was mowed prior and subsequent to dalapon treatments control was 88%, however this level of control probably would not be adequate consid- ering cogongrass' ability to survive and spread (Dickens, 1973). As opposed to the first experiment, the use of mowing with glyphosate and imazapyr treatments did improve cogon- grass control over the use of these herbicides alone, with glyphosate efficacy being improved the most. Control of foliage regrowth with glyphosate was increased from 27% with Table 4.3. The effect of mowing and herbicides on cogon- grass foliage and rhizome dry weight (Experiment 2 1986 through 1988). Mowings1 0 1 2 Rate -ah1 2 3 Herbicide kg ai'ha FDW RDW FDW RDW FDW RDW --- % Inhibition Imazapyr 0.8 64 53 61 52 94 82 Glyphosate 3.4 27 33 61 73 95 73 Dalapon 16.8 -8 3 55 50 88 76 Sulfometuron 1.1 -24 -11 16 49 77 70 Untreated 0 0 5 31 65 51 LSD(0.1)Mowings 16 18 LSD(0.1)Herbicides 21 23 0 mowings indicates herbicides applied to undisturbed cogongrass, 1 mowing indicates mowing prior to herbicide application, and 2 mowings indicates mowing prior and subsequent to herbicide application. 2Cogongrass regrowth foliage dry weight harvested from 0.25 square m quadrants at 12 months after second mowing operation. 3Cogongrass rhizome dry weight harvested from 4170 cubic cm soil cores at 12 months after second mowing operation. Inhibition values computed using unmowed and untreated plots which contained 15.7 and 5.2 g of foliage and rhizome, respectively. no mowing to 61 and 95% when cogongrass was mowed once or twice, respectively. Imazapyr efficacy was increased over imazapyr alone only when cogongrass was mowed both prior and subsequent to herbicide treatment, but was overall .the most efficacious herbicide when used without mowing. The inhibition of rhizome growth in the second experiment followed a pattern similar to that found for foliage regrowth except that the effect of mowing alone increased with each mowing (0 to 31 to 51% inhibition with 0, 1, and 2 mowings, respectively). This indicated that although mowing cannot provide long-term control, it can be utitlized to reduce the potential spread of cogongrass. Overall, from these experiments it was apparent that imazapyr and glyphosate were the most active herbicides tested. Disking-herbicide studies. There was an interaction between experiments and treat- ments in the disking-herbicide studies. Differences in cogon- grass density and environmental conditions may have differ- entially impacted the mechanical and herbicide treatments between experiments. Tables 4.4 and 4.5 present the data collected for the 1985-1987 and the 1986-1988 experiments, respectively. In the first experiment (Table 4.4), the use of a single disking operation, either alone or prior to herbicide treatment, did not control cogongrass regrowth more than the herbicides applied to undisturbed cogongrass. Detachment and decapitation (removal of the apex) of rhizomes caused by Table 4.4. The effect of disking and herbicides on cogon- grass foliage and rhizome dry weight (Experiment 1 1985 through 1987). Diskings1 0 1 2 Rate Rate 1 2 3 Herbicide kg ai'ha FDW RDW FDW RDW FDW RDW --% Inhibition - - Imazapyr 0.8 91 68 85 51 94 85 Glyphosate 3.4 69 74 76 60 85 85 Dalapon 16.8 42 40 56 34 94 85 Sulfometuron 1.1 19 15 4 33 64 51 Untreated 0 0 4 29 57 59 LSD(0.1)Diskings 14 12 LSD(01)Herbicides 17 16 (o.1) 0 diskings indicates herbicides applied to undisturbed cogongrass, 1 disking indicates disking prior to herbicide application, and 2 diskings indicates disking prior and subsequent to herbicide application. Cogongrass regrowth foliage dry weight harvested from 0.25 square m quadrants at 14 months following second disking operation. Cogongrass rhizome dry weight harvested from 4170 cubic cm soil cores at 14 months following second disking operation. 4Inhibition values computed using undisked and untreated plots which contained 66.9 and 17.6 g of foliage and rhizome, respectively. disking may have released dormant nodes from the inhibiting influence of endogenous growth regulator or nutrient gradients (Chancellor, 1974; Leaky, et al., 1975; McIntyre, 1965; and Nyahoza, et al., 1971). These activated nodes may .have produced new shoots in a quantity equivalent to or greater than the amount of shoot tissue destroyed by disking, herbicide treatment, or a combination of the two. However, a second disking reduced foliage regrowth 57% indicating that depletion of reserves and subsequent regrowth potential had been reduced. When dalapon or glyphosate were applied between two diskings the control was 94 and 85%, respectively. Control of cogongrass regrowth by imazapyr was unaffected by disking treatments. Disking alone significantly reduced cogongrass rhizome biomass. Rhizome dry weight was inhibited by 29 and 59% by one and two diskings, respectively. The effects of disking varied with the herbicide evaluated. With sulfometuron, disking caused the greatest enhancement of herbicide activity. When used with dalapon, disking increased control only when implemented prior and subsequent to herbicide treatment. When imazapyr or glyphosate were used with disking, control was significantly reduced by a single pretreatment disking. In the case of imazapyr this apparent reduction in activity may have been caused by increased photolysis which would have occurred in a more open canopy caused previously by disking. This reduction in control may have also been caused by leaf tissue production at the expense of rhizome growth. By changing the source-to-sink pattern from leaf-to-rhizome to rhizome-to-leaf less imazapyr and glyphosate may have been translocated to the rhizomes. This apparent effect of source-sink pattern on glyphosate and other translocatable herbicides has been proposed for several perennial grasses (Atkinson, 1985; Kivlin and Doll, 1988). When disking was implemented prior and subsequent to imazapyr treatment rhizome biomass was reduced 85% which was greater than imazapyr alone. However, the use of glyphosate with two diskings did not inhibit rhizome growth more than glyphosate alone. This may indicate that rhizomes being produced had dormant nodes that were activated by the subsequent disking treatment. In the second experiment (Table 4.5), disking alone had a greater effect on both foliage regrowth and rhizome prod- uction than in the first experiment. Two diskings provided 89 and 72% control of foliage and rhizomes, respectively. This may have occurred due to increased rhizome mortality caused by the dry periods in 1987 following the sequential disking. As opposed to the first experiment, the efficacy of herbicide treatments were increased with each disking performed. This may have been due to the the density of the cogongrass stand in the area in which the second experiment was conducted. This area contained one-fourth the foliage biomass and one-fifth the rhizome biomass of the first experiment area. Table 4.5. The effect of disking and herbicides on cogon- grass foliage and rhizome dry weight (Experiment 2 1986 through 1988). Diskings1 0 1 2 Rate -1 2 3 Herbicide kg ai'ha FDW RDW FDW RDW FDW RDW ---- % Inhibition4 - - Imazapyr 0.8 63 40 71 77 95 95 Glyphosate 3.4 72 30 75 76 99 93 Dalapon 16.8 20 -20 74 79 98 85 Sulfometuron 1.1 -65 -41 61 58 96 81 Untreated 0 0 -9 25 89 72 LSD(0.1)Diskings 20 15 LSD(0.1Herbicides 26 19 0 diskings indicates herbicides applied to undisturbed cogongrass, 1 disking indicates disking prior to herbicide application, and 2 diskings indicates disking prior and subsequent to herbicide application. Cogongrass regrowth foliage dry weight harvested from 0.25 square m quadrants at 12 months following second disking operation. Cogongrass rhizome dry weight harvested from 4170 cubic cm soil cores at 12 months following second disking operation. 4 Inhibition values computed using undisked and untreated plots which contained 17.4 and 3.3 g of foliage and rhizome, respectively. Cogongrass control with the less efficacious herbicides sulfometuron and dalapon were the most affected by the addition of disking. A single disking prior to treatment with sulfometuron increased foliage inhibition from 65% .stimulation (-65% inhibition) to 61% inhibition. Control attained with dalapon increased 270% with the addition of prior disking and 390% with prior and subsequent disking as compared to dalapon alone. The use of prior disking in conjunction with imazapyr or glyphosate did not improve control of foliage regrowth over that attained with either herbicide alone. However, rhizome inhibition did increase with each disking. This indicated that shoot production caused by a single disking was sufficient to overcome rhizome mortality. When imazapyr or glyphosate were used with prior and subsequent disking, control of foliage regrowth and rhizomes was near 100%. Although disking may not be feasible in some situations (i.e., steep terrain or heavily-wooded areas), in pasture renovation and silvicul- tural site preparation tillage is an integral part, there- fore the integration of herbicides (preferably imazapyr or glyphosate) with disking could provide long-term cogongrass control. The influence of photoperiod on shoot initiation in cogongrass rhizomes. Three measurements of shoot initiation potential (total shoot number per productive rhizome segment, percent shoot initiation on productive segments, and percent shoot initation from total node number tested) were utilized to determine the effect of photoperiod. These data are presented in Table 4.6. Continuous dark or continuous light inhibited all three measures of shoot initiation in comparison to the three photoperiods. Leaky, et al., (1978) reported that nodes of rhizome sections of quackgrass [Agropyron repens (L.)Beauv. #AGRRE] were inhibited by light. The cogongrass rhizomes exposed to continuous light were green at the time of harvest which indicated that chlorophyll was present which may have at least in part, negated the need for shoot intiation. This is supported by the fact that rhizomes subjected to the various photoperiods did not visibly produce chlorphyll, but did initiate shoots more consistently (higher initation percentage) and in greater quantity (higher shoot percentage). There was no significant difference in shoot number or percent initiation between long-day, short-day, and interrupted photoperiods. The short-day photoperiod resulted in a higher shoot percentage than did the long-day photoperiod, but not when compared to the interrupted photoperiod. These results, taken in toto, seem to discount a classic photoperiodic and phytochrome response. The implications of this data to shoot initiation following disking seems to point to a minimum amount of daylength required to initiate shoot morphogenesis, but no absolute dark requirement. Rhizomes that become buried during the'disking operation would seem to be less likely to produce shoots than those closer to the Table 4.6. The influence of photoperiod on shoot initiation from cogongrass rhizomes at one week after planting. Light Dark Shoot #1 Initiation2 Shoots3 - hr - - Continuous Continuous 15 8 8 +1 1.7 B4 1.0 B 3.0 A 2.3 AB 2.2 AB 25 B 26 B 64 A 55 AB 67 A 13 C 9 C 45 A 25 BC 35 AB Shoot number per productive rhizome least one shoot). (segments producing at 2Number of segments with at least one shoot/total number of segments tested. Number of shoots/total number of nodes. Means within a column followed by the same letter are not significantly different according to Duncan's Multiple Range Test (o = 0.05). One hour of light during the dark cycle. -----$---- surface. This is even more likely to occur in cogongrass which has specialized rhizome anatomy that reduces the effects of dessication (Holm, 1977). The influence of cogongrass stage of development and defoliation on glyphosate efficacy. Table 4.7 presents data collected from greenhouse studies on the influence of time of defoliation on cogongrass growth. Plants were monitored for 14 weeks, however, after 9 weeks rhizomes were growing out of the pot and leaf area was difficult to determine as lower leaves senesced. Therefore, these data are not presented. Defoliation and subsequent regrowth of three-week-old cogongrass produced plants with a leaf area similar to three-week-old plants. In response to this defoliation, root-rhizome growth temporarily ceased as the plants produced leaf tissue. This is a common phenomenon in perennial grasses (Sturkie, 1930; Youngner and Nudge, 1976). The leaf and root-rhizome weight ratios for three- and six-week-old plants were similar, whereas the plants defoliated at three weeks had a higher root-rhizome weight ratio and a correspondingly lower leaf weight ratio. Plants older than six weeks, whether defoliated or not, had leaf areas that were judged to be too great to attain adequate spray coverage in the subsequent glyphosate studies. There- fore, three growth stages, three-week-old (Stage A), six- week-old (Stage B), and plants defoliated at three weeks after planting and allowed to regrow for three weeks (Stage Table 4.7. The influence of time of defoliation on cogon- grass growth. Weeks after Planting + Weeks Regrowth 3+0 3+3 6+0 6+3 9+0 Leaf Area 2 cm 15.8 (5.4)3 13.2 (5.3) 40.7 (19.4) 189.3 (21.0) 391.0 (91.6) Leaf Weight Ratio1 0.69 (0.06) 0.21 (0.04) 0.58 (0.05) 0.70 (0.04) 0.55 (0.04) Root-Rhizome Weight Ratio2 0.31 (0.04) 0.79 (0.04) 0.40 (0.03) 0.30 (0.03) 0.45 (0.03) 1Leaf dry weight (g)/total plant dry weight (g). 2Root-rhizome dry weight (g)/total plant dry weight (g). 3Means followed by 1 standard deviation. C), were selected for use in studies to determine the influ- ence of relative source-to-sink pattern and relative leaf to below-ground biomass ratio on glyphosate efficacy in cogongrass. The growth stages selected significantly influenced glyphosate efficacy (Table 4.8). Stage A cogongrass was extremely sensitive. A predictive model could not be developed for inhibition of regrowth leaf dry weight (IRLDW) because of a lack of differential response. The predictive model for inhibition of root-rhizome growth (IRRG) indicated that 0.02 kg ae'ha-1 would inhibit growth 50% (I50). Stages B and C were more tolerant of glyphosate than Stage A. There was little difference between the 150 for IRLDW between Stage B and C (0.14 and 0.15, respectively). However, the 150 for IRRG was 0.03 for Stage B and 0.09 for Stage C. This would seem to indicate that Stage B was in a leaf-to- root/rhizome source-sink pattern while Stage C was in a root/rhizome-to-leaf pattern. Glyphosate has been shown to be translocated to rhizomes more in plants that are actively producing rhizomes (therefore a metabolic sink) (Lolas and Coble, 1980). The Stage B cogongrass was known to be actively producing rhizome (Table 4.7) whereas the Stage C plants were producing leaf tissue to the detriment of root-rhizome growth. The fact that the 150 for IRLDW was the same for both these stages does not detract from this hypothesis because the majority of glyphosate recovered by Lolas and Coble was on or in the treated leaf while 2 to 8% Table 4.8. The influence of defoliation on glyphosate efficacy in cogongrass. Regrowth 2 2 Parameter Regression Equation R I50 - - - - Stage A3 -- - -- -- Leaf Dry Weight (RLDW) NS - Rhizome-Root Dry Weight (RRG) %I = -130 45*(Logl0Rate) - - - - Stage B3 - - - Leaf Dry Weight (RLDW) Rhizome-Root Dry Weight (RRG) %I = -212 189*(Log10Rate) 0.90 0.14 %I = -92 34*(Log10Rate) + 35*(Log10Rate)2 0.99 0.03 - - - - Stage C3 -------- Leaf Dry Weight (RLDW) Rhizome-Root Dry Weight (RRG) %I = -228 217*(Log10Rate) %I = -114 60*(Logl0Rate) Glyphosate efficacy based on percent inhibition (%I) of leaf regrowth (RLDW) and composite rhizome-root growth (RRG) over the treatment period. 2Glyphosate rate in kg ae'ha-1 required to cause a 50% reduction in the particular regrowth parameter. Stage A cogongrass was treated 3 weeks after planting, Stage B cogongrass was treated 6 weeks after planting, and Stage C cogongrass was treated 3 weeks after being defoliated at 3 weeks after planting. 0.86 0.02 0.92 0.88 0.15 0.09 was translocated to the rhizomes of johnsongrass. Similar 14 results have been reported using 14C-labelled glyphosate in other species (Wyrill III and Burnside, 1976; Gougler and Geiger, 1981). Stages B and C cogongrass utilized in this study are analagous to established cogongrass infestations in late summer to fall and spring, respectively. Willard (1988) found that cogongrass produced more rhizomes during the later part of the growing season than during the spring. Therefore, it could be assumed that glyphosate applications would be more efficacious when applied during the fall than during the spring. Increased efficacy of fall treatments of glyphosate has been reported for johnsongrass (Derting, et al., 1973) and bermudagrass [Cynodon dactylon (L.)Pers. #CYNDA] (Andrews, et al., 1974; Whitwell and Santelmann, 1978). The results of this research could also be applied to the proper timing of glyphosate following mowing. It would appear that glyphosate applied before sufficient leaf area has regrown to continue production of rhizomes would result in reduced efficacy. Baird, et al., (1973) indicated that mowing within a week of glyphosate treatment reduced control of quackgrass. Mowing outside this time range had no effect indicating that sufficient leaf area had been produced. From a practical standpoint these data indicated that disking or mowing in the spring to remove dead foliage and stimulate new shoot growth then waiting until early fall 78 before applying glyphosate would provide the greatest level of cogongrass control. CHAPTER 5 THE INFLUENCE OF STAGE OF DEVELOPMENT AND MOWING ON BAHIAGRASS [Paspalum notatum var. saurae Parodi 'Pensacola'] AND COGONGRASS [Imperata cylindrica (L.)Beauv.] INTERFERENCE. INTRODUCTION Cogongrass [Imperata cylindrica (L.)Beauv. # IMPCY] has been reported to negatively impact many annual and perennial crop species (Eussen, 1979; Eussen, et al., 1976; Eussen and Wirjahardja, 1973; Ivens, 1975; Jagoe; 1938). Competition for light, water, and nutrients, physical plant injury (caused by the rhizome apices penetrating crop roots), and allelopathy have been reported to be mechanisms of cogongrass interference (Boonitee and Ritdhit, 1984; Eussen and Soerjani, 1975; Eussen and Soerjani, 1976; Soerjani, 1970; Sunarwidi and Batugal, 1982; Zaenuddin, et al., 1986). The combination of these factors and man's inability to effectively control cogongrass has resulted in infestations of 200 million hectares in southeast Asia, several thousand hectares in the southern United States and over 500 million hectares worldwide (Falvey, 1981; Dickens, 1974). The impact of cogongrass on crop production (and Letters following this symbol are a WSSA-approved computer code from Composite List of Weeds, Weed Sci. 32,Suppl. 2. Available from WSSA, 309 West Clark Street, Champaign, IL 61820. subsequently the quality of human life) on a worldwide basis has earned cogongrass the distinction of being one of the ten worst weeds (Holm, et al., 1977). Bahiagrass [Paspalum notatum var. saurae Parodi 'Pensacola' # PASNS] is the predominant grass used on rights-of-way and in pastures in Florida due to its ease of establishment from seed, drought tolerance, and lack of insect and disease problems, while responding to additional inputs of fertilizer, etc. (Beard, 1973; Heath, et al., 1985; Turgeon, 1980). Establishment and mowing are the two major cultural practices used for bahiagrass management on highway rights-of-way (Lewis, 1986). In Florida, cogongrass has become a serious pest on these rights-of-way. It has been hypothesized that on highway rights-of-way cogongrass has become a problem in these areas due to use of rhizome- contaminated soil during bahiagrass establishment following roadway construction and the relatively short-term effect that mowing has on cogongrass growth and development (Willard, 1988). However, the spread of cogongrass into undisturbed, established bahiagrass appeared to be inhibited. Therefore, research was conducted to determine the influence of bahiagrass stage of development and mowing on its interaction with cogongrass. This information will help predict the most efficient route for cogongrass invasion into bahiagrass as well as lead to an understanding the influence that mowing has on the spread of cogongrass into bahiagrass. Materials and Methods Preliminary studies. Prior to conducting replacement studies to determine the influence of bahiagrass stage of development and mowing on its interaction with cogongrass, two greenhouse exper- iments were necessary. The carrying capacity and the response to fertility of each species was determined for the system utilized for these studies. A density study was conducted for each species to determine the carrying capacity of the experimental system utilized in the replacement studies. Cogongrass was propagated from cleaned, de-scaled rhizome segments collected from an infestation in Gainesville, Florida in 1988. Multi-node segments were planted horizontally in trays covered with vermiculite and placed in a growth chamber. The environmental conditions in the growth chamber were; 16-hr light/8-hr dark photoperiod with an intensity of 200)E'm 2"s' and 30 C 2 C. Two weeks after planting single nodes that had produced a two-leaf shoot were exised. Shoots were planted into 2.8 L pots filled with 2.4 L of soil mix containing 50% fumigated2 Arredondo fine sand (loamy, siliceous, hyperthermic, Grossarenic Paleudults) and 50% vermiculite on a volume basis. Densities of 2, 4, 8, 16, and 32 shoots per pot were planted. Commercially obtained 2Bromo-O-Gas (methyl bromide + chloropicrin), Great Lakes Chemical Corp., P.O. 2200, West Lafayette, IN 47906. Pensacola bahiagrass seed3 were planted 0.5 cm deep at 50, 100, 200, 400 per pot. These seeding rates produced plant densities of 29, 54, 104, and 222 plants. Pots were maintained in a greenhouse with the following environmental conditions; 16-hr light/8-hr dark photoperiod, day tempera- + + ture of 30 5 C, night temperature of 25 +5 C and a mean light intensity at noon of 900 pE'm2s -1. Pots were watered daily and fertilized weekly with nutirent solution (Hoagland and Aronon, 1950). Cogongrass and bahiagrass leaf dry weights were determined eight weeks after planting. Leaf dry weight per plant (LDWPP) was determined by dividing the leaf dry weight by the corresponding plant density (PD). The relationship between LDWPP and PD is described in the following equations: Bahiagrass LDWPP = 0.1 0.0003*(PD); r2 = 0.97 Cogongrass LDWPP = 12.2 0.9*(PD) + 0.02*(PD)2; r2 = 0.94 The LDWPP declined as PD for bahiagrass and cogongrass increased. From these studies a PD of 100 for bahiagrass (to be acheived by planting 200 seed) and sixteen for cogon- grass was utilized for the subsequent replacement studies. These densities insured that the replacement studies were conducted at a density that was independent of yield and would therefore maximize interspecific competition. B and G Seed Co., Williston, FL. The effect of added fertility was determined for each species by establishing a PD of 200 and 16 for bahiagrass and cogongrass, respectively, and on a weekly basis applying 0, 40, 200, 400, and 800 ml of nutrient solution per pot (Hoagland and Aronon, 1960). Bahiagrass and cogongrass propagation methodology and greenhouse environmental conditions were identical to those in the density studies. Leaf dry weight (LDW) was determined eight weeks after planting. Leaf dry weights were converted to relative yields (RY) based on the the LDW yield of the highest fertility input. Regression analysis (RY versus ml nutrient solution) was performed on these data. Bahiagrass RY = 0.007 + 0.004*(ml) 0.00003*(ml)2; r2 = 0.96 Cogongrass RY = 0.18 + 0.002*(ml) 0.000001*(ml)2 r2 = 0.97 From these models replacement-series studies were fertilized weekly with 500 ml nutrient solution. This fertility level was utilized to alleviate interspecific competition for nutrients. Seedling bahiagrass and emerging cogongrass interference studies. The interaction of seedling bahiagrass and emerging cogongrass was studied under greenhouse conditions using a replacement-series experimental design (Wit, 1960). Bahia- grass and cogongrass were planted in 2.8 L pots containing the same soil mix used in the preliminary studies. Five treatments consisting of varying ratios of bahiagrass PD to cogongrass PD were utilized; 100:0, 75:4, 50:8, 25:12, and 0:16. All pots were watered daily and fertilized weekly .with nutrient solution. The first experiment was conducted in a greenhouse with the same environmental conditions as those described previously. The second experiment was conducted in a greenhouse with the same environmental condi- tions except that the light intensity at noon averaged 1300 pE'm-2.s-1 (averaged 25% higher intensity at any given time during the day)4. Eight weeks after planting, leaf dry weight (LDW) and height of each species and shoot number and rhizome dry weight of cogongrass were determined. Relative yield (RY) of LDW for each species was obtained by dividing the absolute yield at each ratio by the yield of each species in monoculture. Relative yield totals (RYT) were determined by adding the RY of each species within each ratio mixture. Relative crowding coefficients (RCC) were determined for each species as described by Harper (1977). The influence of mowing on established bahiagrass and emerging cogongrass interference. To determine the influence of cogongrass on established bahiagrass as affected by mowing frequency, bahiagrass densities of 25, 50, 75, and 100 plants per pot (under environmental and soil conditions identical to those used in 4Li-Cor Model LI-188B Integrating Quantum/Radiometer/Photo- meter, Li-Cor, Inc., Lincoln, NE. the seedling study) were established. Eight weeks after planting bahiagrass foliage above 10 cm was removed using hand-held, horizontally-reciprocating shears5. Into these pots were planted single-node cogongrass rhizome segments that had produced a two-leaf shoot. A replacement-series design was utilized in which the following bahiagrass PD to cogongrass PD were established; 100:0, 75:4, 50:8, 25:12, and 0:16. These pots were watered daily and fertilized weekly with nutrient solution. Four weeks after estab- lishing these mixtures two-thirds of the pots were clipped at 10 cm above the soil surface. During this procedure the leaf weight (LDW) of each species in each mixture was determined. Four weeks later half of the previously clipped pots were clipped at 10 cm and species dry weights recorded. This produced three clipping regimes (0, 1, and 2) for each species ratio. Four weeks following the second clipping all pots were harvested. At this time, LDW and height of each species and shoot number and rhizome dry weight of cogongrass was determined. Total leaf dry weight (TLDW) of each species was determined by adding the LDW harvested at each clipping interval to the final LDW. Relative yield (RY) of each species was computed by dividing the TLDW of each species in competition by the TLDW produced by each species in monoculture. The relative yield total (RYT) was determined by adding the RY of each species within each Diston Model EGS-HD4, Danville, VA 24543 ratio mixture. The relative crowding coefficients (RCC) were also determined. Experimental design and analysis. All experiments were conducted twice using a randomized complete block design with four replications. Analysis of variance was utilized to determine if the influence of single factors (species, ratio, mowing, and experiment) and interactions between these factors significantly affected yield components (Helwig and Council, 1982). When experiment significantly interacted with species-ratio treatments (P < 0.05) the individual experiments are presented separately. When there was no experiment by species-ratio treatment interaction (P > 0.05) data were pooled. Results and Discussion Seedling bahiagrass and emerging cogongrass interference studies. The two experiments conducted to determine the interaction of seedling bahiagrass and emerging cogongrass differentially impacted the competitiveness of the two species. Table 5.1 presents the separate results of these two experiments. The first experiment was conducted in a greenhouse that transmitted, on average 25%, less light than the greenhouse in which the second experiment was conducted. The influence of reduced light intensity on cogongrass growth and development has been reported by Soerjani (1970) and Patterson (1980). Reduced light caused a greater Table 5.1. The interaction of seedling bahiagrass (B) and emerging cogongrass (C) as measured by relative yield (RY), relative yeild total (RYT) and relative crowding coefficient (RCC) of leaf dry weight and height and total pot dry weight at eight weeks after planting. Experiment 11 Experiment 2 --------------------- Leaf Dry Weight -------------------- 0.58 (0.04)3 0.41 (0.02) 0.99 1.57 0.64 0.20 (0.03) 0.54 (0.05) 0.74 0.48 2.08 -------------------------- Height ------------------------- - cm- - - HeightB (Monoculture) 35.6 (0.9) Heights (50:50% Mixture) 36.6 (3.6) HeightC (Monoculture) 59.0 (1.5) HeightC (50:50% Mixture) 51.8 (2.4) 41.5 (2.3) 28.8 (1.9) 71.5 (4.3) 67.5 (2.2) ------------------- Total Pot Dry Weight ------------------- - -- -g-- ---- Monoculture B 50:50% Mixture Monoculture C 33.3 (2.5) 31.6 (1.6) 33.9 (2.6) 29.4 (3.8) 28.1 (3.4) 45.6 (6.5) Experiment 1 conducted in a greenhouse with 25% less light intensity as compared to Experiment 2. 2 RY of total leaf dry weight of each species and RYT computed at the 50:50% ratio mixture. Where appropriate mean values are followed by standard errors in parenthesis. RCC values computed using mean dry weight per plant of each species at the 50:50% ratio mixture and in monoculture. 2 RY B 2 RY RYT2 RCCB4 4 RCCC C reduction in rhizomes than leaves. However, as rhizomes are a product of carbon fixation in the leaf, effects on leaf biomass must be considered paramount. Under the reduced light conditions in the first experi- -ment seedling bahiagrass was more competitive than emerging cogongrass, with RY's of 0.58 and 0.41, respectively. Using RCC, bahiagrass was 2.4 times more competitive than cogon- grass (RCC of 1.57 and 0.64, respectively). The RYT was 0.99 indicating that the two species in mixture were competing interspecifically. Since water and nutrient status was maintained at an optimal level, this indicated that light was the limiting factor. Though cogongrass was taller on an absolute basis, its erect leaf morphology may have allowed sufficient light penetration to maintain bahiagrass. Bahia- grass height was not affected by cogongrass competition, (as compared to bahiagrass in monoculture), but cogongrass height was reduced 12.2% when competing interspecifically (59.0 versus 51.8). Total biomass produced was equivalent for each species in monoculture and when the two species were competing interspecifically. Under the higher light intensity in the second experi- ment, cogongrass was more competitive than bahiagrass, with RCC of 2.08 and 0.48, respectively. This indicates that cogongrass became 4.3 times more competitive under the high light conditions. The RY values of 0.54 and 0.20 for cogon- grass and bahiagrass, respectively, indicated that cogongrass was contributing over half of the total yield in the system. |
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| 0 | sobekcm_page_globals.get_entire_collection_hierarchy | Retrieving hierarchy information |
| 0 | sobekcm_assistant.get_entire_collection_hierarchy | |
| 0 | cached_data_manager.retrieve_item_aggregation | |
| 0 | cached_data_manager.retrieve_item_aggregation | Found item aggregation on local cache |
| 0 | item_aggregation_builder.get_item_aggregation | Found 'all' item aggregation in cache |
| 0 | system.web.ui.page.page_load (ufdc.page_load) | |
| 0 | sobekcm_page_globals.constructor.on_page_load | |
| 0 | html_echo_mainwriter.add_style_references | Adding style references to HTML |
| 0 | html_echo_mainwriter.add_text_to_page | Reading the text from the file and echoing back to the output stream |
| 40 | html_echo_mainwriter.add_text_to_page | Finished reading and writing the file |