Lower leaf harvesting options and leaf position effects on some agronomic, chemical, and mineral characteristics of flue...

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Title:
Lower leaf harvesting options and leaf position effects on some agronomic, chemical, and mineral characteristics of flue-cured tobacco
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viii, 106 leaves : ill. ; 29 cm.
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Stocks, Glenn Ralph, 1963-
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Tobacco -- Harvesting   ( lcsh )
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Thesis:
Thesis (Ph. D.)--University of Florida, 1991.
Bibliography:
Includes bibliographical references (leaves 99-103).
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by Glenn Ralph Stocks.
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Typescript.
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Vita.

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University of Florida
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LOWER LEAF HARVESTING OPTIONS AND LEAF POSITION EFFECTS ON SOME
AGRONOMIC, CHEMICAL, AND MINERAL CHARACTERISTICS
OF FLUE-CURED TOBACCO











By


GLENN RALPH STOCKS


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

1991


UNIVERSITY OF FLORIDA LIBRARIES














PRELUDE

"At the awful day of judgment, the discrimination of the good from the wicked, is

not made by the criterion of the sects or of dogmas, but by one which constitutes the

daily employment and the greatest end of agriculture. The judge upon this occasion has

by anticipation pronounced, that to feed the hungry, clothe the naked, and give drink to

the thirsty are the passports to future happiness; and the divine intelligence which

selected an Agricultural state as a paradise for its first favorite, has here again

prescribed the Agricultural virtues as the means for the admission of their posterity

into heaven."

John Taylor, 1813














ACKNOWLEDGMENTS


In this study, 30,024 tobacco leaves were used to generate 40,224 data points.

Of the 30,024 leaves used, 10,592 had the midribs removed manually. There were

3636 samples that were ground for either mineral and chemical, or total non-

structural carbohydrate analysis. To accomplish all of this work required the assistance

of many people and the author intends to duly recognize all involved who made this Ph.D

research program possible.

Gerald Durden, David Durden, and Shannon Brown's technical assistance in

harvesting and measurement of the agronomic parameters is greatly appreciated. Ernest

Terry's assistance in grinding, and processing of samples for TNC and mineral analysis

was invaluable. Chief Tobacco Technician Rick Hill's assistance in the total management

of this program is gratefully appreciated.

Much appreciation is given to Dr. D.G. Shilling and Dr. J.M. Bennett for their

advice and counsel all through the conducting of this study. Also, gratitude is given to Dr.

R.N. Gallaher for the use of his dryer for desiccating the samples used in the TNC study

and the use of his laboratory for sample preparation for mineral analysis. The use of

Dr. Bennett's lab space for the TNC analysis is much appreciated. Unfortunately, Dr.

F.M. Rhoads was stationed outside of Gainesville and interaction with him was limited by

the miles; however, his willingness to serve on the supervisory committee is

appreciated.

To all committee members, the author is grateful for the challenges presented to

him in the qualifying examinations. The diversity of this committee challenged the

author on the written and oral exams. Each and every member posed probing questions

that caused the student to question his reasoning for being in graduate school. The









successful completion of such a diverse qualifying examination process no doubt was the

turning point of this student's program.

The author's parents lifelong devotion to him is greatly appreciated. Had his

father, a farmer, not involved the author with the farming operation, a career in

agriculture might not have become a reality. The author's mom basically did all the

paper work for the author to be enrolled at N.C. State. Had she not done this, the author

might well have ended up in the armed forces.

By the time the author has completed this dissertation, he will will have taken a

bride. The loving devotion of Kathleen Best has carried the author through the inevitable

low points of this graduate program. Kathleen has shared the good times and bad times

with equal vigor and picked the author up when he was feeling down. Many years of

happy marriage are looked forward to by the author.

Dr. E.B. Whitty has been a godsend to the author. As a major professor, Dr.

Whitty has provided leadership to the author. As a person, Dr. Whitty is one of the

author's best friends. As a scientist, Dr. Whitty is clearly a leader in his field, as his

recognition as 1991 Florida Extension Specialist of the Year clearly demonstrates. In

the author's humble opinion (although some might say there is nothing humble about the

author), there is not a better major professor than Dr. Whitty. Academics aside, Dr.

Whitty has allowed the author to develop professionally by including him on extension

programs and sending him to numerous professional meetings. From a research stand-

point, whenever the author needed something for his studies, Dr. Whitty willingly

supplied the needed equipment. The author is forever indebted to Dr. Whitty for having

served as his major professor for both the M.S. and Ph.D programs. Without question,

whatever success the author experiences over his career will be directly due to his

interaction with Dr. Whitty.

The funding of the research assistantship by R.J. Reynolds Tobacco Company made

it possible for the author to obtain his Ph.D degree. R.J. Reynolds has been an intricate









part of the author's academic career since he was an RJR research apprentice in 1984.

This program got the author interested in agriculture research and led directly to him

attending graduate school at the University of Florida. The interaction with Mr. A.R.

Mitchum, Mr. R.C. Reich, Dr. D.L. Davis, and Dr. C.R. Miller, all of RJR, has allowed the

author to develop friendships with industry that hopefully will continue throughout the

author's career.

Without the cooperation of R.J. Reynolds and Philip Morris Tobacco Companies in

running the leaf chemical analyses, these studies would not have been complete. The

author is indebted to both companies for their willingness to run these analyses. The

cooperation of Mr. A.R. Mitchum of RJR and Mr. D.L Connor of PM is gratefully

appreciated in arranging to have these samples processed and analyzed.

In summary, anyone who has read this dissertation recognizes the immense

amount of data that was generated. This was clearly a team project and all involved

deserve and are accorded a gargantuan thank you from the author, Glenn R. Stocks.














TABLE OF CONTENTS


amBS
ACKNOW LEDGMENTS ................................................................................................. iii

ABSTRACT ............................................................... ...................... ........................... vii

CHAPTERS

1 INTRODUCTION ............................................................. 1

2 CHARACTERIZATION OF FLUE-CURED TOBACCO BY LEAF
POSITION PRODUCED UNDER NORMAL HARVESTING
METHODS OR MONITORED OVER TIME AFTER TOPPING 7

Introduction ................................................ .................. 7
Materials and Methods .................................................. 11
Results and Discussion ........................................ ........ 18
Conclusions .................................................................. 49

3 LOWER LEAF HARVESTING EFFECTS ON AGRONOMIC
CHARACTERISTICS OF FLUE-CURED TOBACCO ............. 50

Introduction ................................................................. 50
Materials and Methods ....................................... .......... 52
Results and Discussion ...................................... .......... 54
Conclusions ................................................. ................. 65

4 LOWER LEAF HARVESTING: THE INFLUENCE OF TIME ON
THE CHEMICAL AND MINERAL CHARACTERISTICS OF
FLUE-CURED TOBACCO LEAF POSITIONS .................... 66

Introduction ................................................. ................ 66
Materials and Methods ....................................... .......... 68
Results and Discussion ................................................ 73
Conclusions .................................................. ................ 89

5 SUMMARY AND CONCLUSIONS ....................................... 93

REFERENCES ......... ........... .. ...................................................................................... 99

BIOGRAPHICAL SKETCH ........................................................................................... 104













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

LOWER LEAF HARVESTING OPTIONS AND LEAF POSITION EFFECTS ON SOME AGRONOMIC,
CHEMICAL, AND MINERAL CHARACTERISTICS OF FLUE-CURED TOBACCO

By

Glenn Ralph Stocks

December, 1991
Chairman: Dr. E.B. Whitty
Major Department: Agronomy

Lower leaf harvesting options are management tools of flue-cured tobacco
(Niciana tabacum L.) farmers that are determined by economic considerations. The

lowest leaves of the flue-cured tobacco plant are the lowest in yield and value and some

farmers choose not to harvest them because of their relatively low economic return.

There are inherent differences in the agronomic, chemical, and mineral characteristics

of tobacco leaves depending on where the leaves are positioned on the stalk. Pruning and

discarding the lowest three or four leaves has been shown in some studies to not

adversely affect total yield. However, pruning lower leaves reduced yield in another

study. The objective of this study was to evaluate the effects of five lower leaf

harvesting options on leaf position characteristics of flue-cured tobacco plants having

exactly 21 leaves prior to pruning. The responses of leaf position parameters were

evaluated under normal harvesting methods and a time after pruning study involving

acquisition of all leaves that remained on plants at a given period of time. Not harvesting

the lower leaves reduced yield as total leaf number harvested declined, but average value

per kilogram was increased because the lowest leaves were the lowest valued No

differences in leaf position yield or total non-structural carbohydrate concentration

were found, leading to the conclusion that lower leaf harvesting treatments had no effect









on net photosynthesis of the leaves above them. Not pruning and not harvesting the lower

leaves resulted in lower lamina nicotine and P concentrations and higher concentration

ratios of N to nicotine and reducing sugars to nicotine. The reduced concentrations of

nicotine and P were associated with a dilution effect due to not pruning and not

harvesting the lowest leaves on the tobacco plants. Results from these studies describe

the influence that the position of tobacco leaves on the stalk had on leaf agronomic,

chemical, and mineral parameters and the influence of leaf maturity on the development

of those same leaf parameters.


viii













CHAPTER 1
INTRODUCTION

Lower leaf harvesting options of flue-cured tobacco (Nicotiana tabacum L)

provide practical means of managing the yield and value per unit of the crop. Lower leaf

harvesting options have become management tools for two reasons: 1) the federal

program for flue-cured tobacco controls the weight of tobacco that can be sold from a

farm, and 2) there are inherent differences in value per kilogram of tobacco based on

the position of the leaves on the stalk. The value per kilogram of flue-cured tobacco

generally is lowest for the lowest leaves and increases progressively to the middle- and

upper-leaf positions on the stalk.

Because the lowest leaves on the tobacco plant are the lowest in yield and value

and improved cultivars produce high yields, some growers choose not to harvest the

lowest leaves. Data from previous studies dealing with lower leaf harvesting options of

flue-cured tobacco suggested that pruning and discarding the lowest three or four leaves

did not adversely affect total yield and improved the average value per kilogram when

compared to harvesting all leaves (Suggs, 1972; Currin and Pitner, 1980; Stocks,

1988). However, Court and Hendel (1989) found that if the number of leaves harvested

was reduced from 18 to 15 to 12, either by lower leaf pruning or topping to a lower leaf

number, total yield was progressively reduced.

In that lower leaf harvesting options reduce the total number of leaves harvested,

other methods may be used to achieve the same goal. Woltz and Mason (1966) found that

increasing the leaf number per hectare through higher plant populations increased yield

with the response fitting a quadratic model. When leaf quality was evaluated, 296,400

leaves per hectare were found to be the optimum. Collins et al. (1969) found a leaf









population of 444,600 per hectare increased yield and value when compared to

296,400 leaves per hectare, but value per kilogram decreased with the increase in leaf

population. They concluded that the production of the additional leaves was not

economically feasible. Kittrell et al. (1972) found a leaf population of 370,500 per

hectare increased yield and value when compared to 296,400 leaves per hectare, but

gross and net prices were reduced by the higher leaf population.

Topping height and within-row plant spacing are the most common methods used

to achieve a desired leaf population. Kittrell et al. (1972) found that with equal within-

row plant spacings a topping height of 20 leaves per plant yielded higher than that

observed when topping at 16 leaves per plant with equal gross prices for the respective

topping heights. Net price was higher for the 20 leaves per plant topping height. Elliot

(1976) evaluated the effects of several within-row plant spacings and topping heights

on flue-cured tobacco. He found that increasing topping heights from 12 to 15 to 18

leaves per plant generally increased total yield and value per hectare for each increase

in leaf number per plant, except for a lower value per kilogram for the 18 leaf topping

height treatment. Lower topping did not increase the specific leaf weight of the lowest

leaves or highest leaves, but increased the specific leaf weight of the middle leaves.

Increasing within-row plant spacings decreased yield and value per hectare, but

increased specific leaf weight of all leaves. Lower topping heights and wider within-row

plant spacings increased nicotine and total N concentration, with no response of topping

height or plant spacing on reducing sugar concentration.

The objective of lower leaf harvesting options is to maximize the use of quota by

selling the highest quality leaves. By not harvesting lower leaf tobacco, the lowest

quality and value leaves are not marketed. Information in the literature suggests that

the weight, and perhaps the quality, of the lower leaves can be improved by wider

spacing of plants within the row. Lower topping did not increase the weight of the lowest

leaves, but lamina weight increases were found in the middle leaves on the stalk.









However, lower yields resulted from both options. Increasing the within-row plant

spacing or topping to a lower leaf number limits the farmer's lower leaf harvesting

options in that he probably will need to harvest all the leaves produced on the plant to

make full use of his allotted quota. Within-row spacings and topping heights that

produce good quality tobacco and that may allow a farmer to exceed his allotted quota

under favorable growing conditions would enable the farmer to decide whether or not to

harvest the lower leaves. Those decisions would be based on the expected quality and

price per kilogram for the lower leaves, as well as the expected total yield of the crop.

All the studies dealing with lower leaf harvesting options have reported total

yield and explanations for the yield responses were not fully elucidated. Yield of each

leaf position would be a useful parameter when evaluating the effects of lower leaf

harvesting options. If the studies reported by Suggs (1972), Currin and Pitner

(1980), and Stocks (1988) had included yield by leaf position, the distribution of the

total yield on the plant could have been evaluated to determine where the yield

redistribution occurred due to lower leaf pruning.

Physiologically, lower leaf harvesting options of tobacco can be viewed as a

manipulation of the source-sink relationship of the plant. In flue-cured tobacco

production, the reproductive sink is removed (topping) to increase the yield and quality

of the leaves that remain on the stalk. The topping process has been found to eliminate

the traditional source-sink relationship of the tobacco plant causing the leaves to

function as alternate sinks (Humg et al., 1989). Some effects of topping are increased

specific leaf weight and leaf carbohydrate concentration (Hurng et al., 1989), and

increased leaf nicotine concentration (Woltz, 1955). Given equivalent environmental

and nutritional conditions, the timing of topping has significant effects on the agronomic

and chemical properties of the tobacco crop. Early topping increased leaf yield and

nicotine concentration more than late topping (Woltz, 1955; Steinberg and Jeffery,

1957; Marshall and Seltmann, 1964; Elliot, 1966; Stocks and Whitty, 1992). Woltz









(1955), Marshall and Seltmann (1964), and Elliot (1966) found N and reducing sugar

concentrations were not affected by topping delays. As topping has such a dramatic

influence on some of the agronomic and chemical qualities of the crop, further

manipulation of the plant's source-sink relationship, i.e., lower leaf harvesting options,

might be expected to influence similar parameters. Suggs (1972) found that when the

lowest nine leaves were pruned and discarded from flue-cured tobacco plants nicotine

concentration was increased in the remaining leaves.

Numerous studies have dealt with source-sink manipulations in other plant
species. Removal of the grain sink of corn (Zea mavs L) caused dramatic increases in

the carbohydrate concentration of both upper and lower leaves (Allison and Weinmann,

1970). Pod removal from soybean (Glycine max L Merr.) increased leaf carbohydrate

concentration (McAlister and Krober,1958; Kollmann et al., 1974; Ciha and Brun,

1978; Mondal et al., 1978; Streeter and Jeffers, 1979; Crafts-Brandner et al.,

1984), and also increased N and P concentrations in the leaves (Kollman et al., 1974;

Crafts-Brandner et al., 1984). Kollmann et al. (1974) further reported that leaf Ca

and K concentrations were decreased due to depodding of soybean. Lawn and Brun

(1974) found soybean depodding decreased net photosynthesis. The photosynthetic

decline was linked to an accumulation of assimilate in the leaves. Depodding of soybean

also has been found to delay leaf senescence (Hicks and Pendleton, 1969; Mondal et al.,

1978; Crafts-Brandner et al., 1984). The effects of reproductive sink removal can be

summarized to include increased leaf carbohydrate levels with a subsequent decline in

net photosynthesis, increase in leaf N and P concentrations, decreases in leaf Ca and K

concentrations, and a delay in the onset of senescence.

Results from leaf removal studies with wheat (Triticum aestivum L) and oats

(Avena sativa L.) demonstrate that leaf area losses greater than 10% reduced grain yield

(Womack and Thurman, 1962). Grain yield of sorghum (Sorghum bicolor L. Moench)

was reduced as defoliation increased (Stickler and Pauli, 1961). Pauli and Stickler









(1961) found that increases in the percentage of leaves pruned from grain sorghum

plants decreased the total carbohydrates in the vegetative tissue and grain. Vegetative

tissue N concentration decreased and grain N concentration increased as the percentage of

defoliation increased. Weber (1955) and McAlister and Krober (1958) found soybean

seed yield and size decreased in response to defoliation. The causal mechanism of lower

grain yield with defoliation was the loss the total photosynthetic capacity of the plant.

Flue-cured tobacco is managed differently than most other agronomic crops in

that the leaves, and not the seed, are harvested. Also, leaves are removed as they mature

on the plant. Most source-sink manipulation studies with other crops have dealt with

the effects of either the removal of the reproductive sink on leaf and seed characters or

the removal of leaves on seed characters. Lower leaf harvesting option studies with

flue-cured tobacco have reported total yield and chemical effects only. To ascertain the

specific effects of lower leaf harvesting options, a study was designed and implemented to

evaluate leaf position responses to five lower leaf harvesting options of flue-cured

tobacco plants having exactly 21 leaves. The five lower leaf harvesting options were:

1) Harvest all 21 leaves in a normal manner (control).

2) Prune and discard the lowest 3 leaves, harvest remaining 18 leaves in a
normal manner.

3) Do not prune and do not harvest the lowest 3 leaves (leaves were left on the
stalk), harvest remaining 18 leaves in a normal manner.

4) Prune and discard the lowest 6 leaves, harvest remaining 15 leaves in a
normal manner.

5) Do not prune and do not harvest the lowest 6 leaves (leaves were left on the
stalk), harvest remaining 15 leaves in a normal manner.

To maintain the integrity of the leaf positions, the 21 leaves of the plants were

partitioned into seven, 3-leaf stalk positions with the lowest three leaves designated

1-3 and progressing up the stalk with the upper-most three leaves designated 19-21.

The objective of the present study was to evaluate the effects of pruning and

discarding, not pruning or harvesting, or harvesting in a normal manner, the lowest









three or six leaves on flue-cured tobacco plants on the leaves above those involved in the

the treatments. Leaf number per plant and by harvest was controlled so that any effects

due to the treatments could be reported by the position or node on the stalk where leaves

were formed. Previous work on this topic indicated positive effects on yield and value by

pruning and discarding lower leaves, but negative effects by not pruning or harvesting

the same leaves.

The present study involved the measurement of numerous properties of tobacco

leaves based on the position or node on the stalk at which leaves were formed. Data are

lacking that characterize the agronomic, chemical, and mineral qualities by leaf position

of currently-grown flue-cured tobacco cultivars. These data are reported by leaf

position for normally-harvested mature leaves and for the same leaf positions over time

after topping to maturity to contribute data on the characteristics and the development of

the leaves comprising a leaf position.














CHAPTER 2
CHARACTERIZATION OF FLUE-CURED TOBACCO BY LEAF POSITION PRODUCED UNDER
NORMAL HARVESTING METHODS OR MONITORED OVER TIME AFTER TOPPING

Introduction

Tobacco (Nicotiana tabacum L) production, unlike that of most other agronomic

crops, involves harvest of the vegetative tissue (leaves), not the reproductive tissue

(seeds). Flue-cured tobacco leaves, unlike most other tobacco types, are harvested

manually or mechanically as the leaves mature on the plant. Harvest of the flue-cured

tobacco leaves progresses with maturity from the lowest to the highest leaves on the

plant stalk. This harvest method allows for all leaves to become fully mature if proper

production practices are followed. With most other tobacco types, the entire plant is

severed in the field at a stage when a majority of the leaves on the plant are judged to be

mature. Although the progressive harvest of flue-cured tobacco results in mature

leaves, there are considerable physical and chemical differences depending on the

position or nodes at which leaves are formed on the stalk (leaf position).

Leaf position (LP) is of considerable importance to the tobacco industry. Tobacco

is marketed, graded, and sold according to LP regardless of type. Tobacco companies

purchase tobacco based on their needs for a given characteristic most often derived from

the inherent differences between LPs. The tobacco is processed and stored based on LP,

as well. Tobacco products are manufactured based on certain characteristic properties

of the cured leaf. Weybrew et al. (1984) determined that the leaf chemical components

of tobacco are influenced most by LP. In general, the LP characteristics of tobacco

dictate what type product can be manufactured.








Because LP is such an important factor in tobacco production and manufacturing,

numerous studies over the years have evaluated a wide array of LP properties for

tobacco types. Various burley tobacco LP properties have been described by Bowman and

Nichols (1968), and Williamson and Chaplin (1981). Flue-cured tobacco LP

properties have been described over the years. Agronomic, chemical, and mineral

composition data on LPs of flue-cured tobacco can be garnered from reports by Darkis et

al. (1936, 1952), Askew et al. (1947), Walker (1968), Brown and Terrill (1972,

1973), Bowman et al. (1973), Nel et al. (1974), Neas et al. (1978), and Campbell et

al. (1980). Raper and McCants (1966), Srivastava et al. (1984), and Bruns and

Mclntosh (1988) has reported dry matter accumulation data for tobacco, but these

studies were based on whole plant sampling with no segregation of the leaves.

One of the most important production and management practices of flue-cured
tobacco (Nicotian tabacum L.) is the removal of the apical meristem (topping). Tobacco

is topped to improve yield and quality of the upper leaves. The topping process breaks

apical dominance resulting in rapid axilary bud development and generally corresponds

to the onset of floral initiation. The axilary buds (suckers), if allowed to develop will

reduce yield and quality of the tobacco leaves. Consequently, farmers remove suckers by

hand or use growth regulators to suppress their development. The process of topping is

also associated with many important developments of the tobacco plant. Topping, for

practical purposes, eliminates the traditional source-sink relationship for a tobacco

plant. Hurng et al. (1989) found that topping tobacco plants increased leaf dry weight,

specific leaf weight, and P concentration indicating that in the absence of the

reproductive sink, the leaves acted as an alternate sink. Wolf and Gross (1937) detailed

anatomical changes associated with tobacco topping and found larger, thicker leaves

resulted due to an increase in cell size in response to topping. Steinberg and Jeffery

(1957) found topping increased root development and nicotine concentration. Nicotine

is synthesized in the roots of the tobacco plant and translocated to the leaf (Dawson and









Solt, 1959). Wolf and Bates (1964) showed that a more extensive tobacco root system

resulted in higher leaf nicotine concentrations.

Nicotine is the most unique of all the chemical components of a tobacco leaf, and

its accumulation in the leaf as a response to topping is an important event, but, other

important processes are associated with topping. Elliot (1975) reported that topping

increased, not only nicotine concentration, but also lamina weight, leaf N and reducing

sugar concentration, and quality of the cured leaf.

At topping, there is a clear and distinct gradient in leaf age on a tobacco plant.

The oldest and most mature leaves will be the lowest on the stalk, and leaf maturity will

decline as LP increases to the top of the plant. Wolf and Gross (1937) reported that

leaves that were the most mature at topping were least modified, while those that were

least mature were profoundly modified. In Florida, topping generally occurs 80 to 90

days after transplantation of seedlings. The topping process often coincides with the

first harvest of the mature lowest leaves.

Because the production of flue-cured tobacco involves the harvest of leaves as

they mature on the stalk, management of the crop becomes a science of leaf senescence.

Weybrew et al. (1984) state that "physiological maturity (of a tobacco leaf) marks the

transition from growth to senescence and is usually identified as the point of maximum

dry weight attainment." They estimated that a leaf becomes ripe approximately 12 days

after physiological maturity. Also, because the leaves emerge sequentially, they will

ripen progressively from the lowest leaf, earliest emerged, to the uppermost leaf, latest

emerged. Weybrew et al. (1984) proposed that weekly harvests of three leaves per

plant would be the optimum way to produce quality tobacco because one leaf should be

"ideally" ripe, and the other two leaves would be only two days before or past the

"ideally" ripe stage.

Moseley et al. (1963) stated that "as a tobacco leaf approaches maturity, it loses

much of its tackiness and acquires a velvety feel. It develops "grain" or mounds between









the small veins. It becomes more turgid, does not wilt readily, and will snap crisply

from the stalk. Both the yellow and green pigments decrease, but the green ones at a

faster rate, thus the leaf becomes less green and more yellow in appearance." Moseley et

al. (1963) studied the maturity of tobacco leaves at harvest and found chlorophyll

decreased at a faster rate than did carotene or xanthophyll. Nicotine concentration

increased with maturity. Reducing sugar, N, and K concentrations, and the N to nicotine

and reducing sugar to nicotine concentration ratios declined as the tobacco leaves

matured. Walker (1968) found that as tobacco leaves matured the Ca, Mg, and K

concentrations declined, and noted inherent concentration differences between leaf

positions.

On a more general nature, Leopold (1961) described two positive effects of leaf

senescence. As leaves become shaded, senescence allows for organic and inorganic

compounds, which had been committed to those leaves during growth and development, to

be remobilized to leaves that are actively growing in a more favorable environment.

Also, with age and shading, the photosynthetic activity of a leaf declines sharply. The

shaded leaves are not parasitic to the plant, rather they are removed through senescence.

Thomas and Stoddart (1980) suggested leaf senescence is: 1) controlled genetically, 2) a

result of competition for light, space, and nutrients, or 3) a response to environmental

factors such as light, temperature, water relations, mineral relations, and diseases.

Sinclair and deWit's (1975) "self-destruct" hypothesis suggested soybean leaf

senescence to be a result of N being transported out of the leaves to supply the developing

seed to such an extent the leaves senesce. Flue-cured tobacco leaves ripen or partially

senesce as result of N starvation. In Sinclair and deWit's (1975) hypothesis the N

demand of the developing soybean seed depleted leaf N. However, in flue-cured tobacco

production N is managed such that the plant gradually depletes the soil N supply, thereby

limiting the N availability to the leaves, progressively from the lowest leaves to the top

leaves.









In flue-cured tobacco production, proper N management is the key to quality

cured leaf. Ideally, the N management scheme would cause the soil N supply to be

depleted when all but the upper-most leaves have fully developed allowing for the

remobilization of N from mature to maturing leaves. This scheme is the objective of

every flue-cured tobacco farmer each year, however, environmental factors nearly

always make this scheme difficult to achieve.

Numerous studies have detailed nutrient and dry matter accumulation for various

tobacco types over time (Grizzard et al., 1942; Raper and McCants, 1966; Sims and

Atkinson, 1971,1973, and 1974; Atkinson and Sims, 1977; Raper et al., 1977;

Srivatava et al., 1984; Bruns and Mclntosh, 1988). In all the above mentioned studies,

composite samples were acquired and data were reported on a whole plant basis. Studies

are lacking that detail the effects of time on the development of individual LPs.

Data are lacking that describe the characteristics of leaves by position on the

stalk of currently grown flue-cured tobacco cultivars. In the present study, agronomic,

chemical, and mineral data were collected from two flue-cured tobacco cultivars

('NC37NF' and 'K-358') topped to 21 leaves. The purpose of this study was to provide

information on tobacco leaf characteristics based upon the position or node on the stalk at

which leaves were formed. Data on LP characteristics over time are presented because

changes in the characteristics of individual LPs have been described in few previous

studies.

Materials and Methods

Field experiments were conducted in 1989 and 1990 at the University of

Florida's Green Acres Agronomy Farm near Gainesville, FL. Two flue-cured tobacco

cultivars, 'NC37NF' and 'K-358', were grown in an Arredondo fine sand (fine-sandy

siliceous, Hyperthermic Grossarenic Paleudult) (Carlisle et al., 1989) for both years.









Plants were spaced 41 cm apart in rows spaced 121 cm apart. Transplanting dates were

10 March 1989 and 2 April 1990.

Prior to transplanting in 1989, weed and soil-borne pest management consisted

of 6.73 kg (a.i.) ha-1 fenamifos (nematicide) {Ethyl 3-methyl-4-(methylthio)

phenyl(1-methylethyl)phosphoramidate}, 2.26 kg (a.i.) ha-' chlorpyrifos

(insecticide) {O,0-Diethyl 0-(3,5,6-trichloro-2-pyridinyl)-phosphorothioate),

4.46 kg (a.i.) ha-' pebulate (herbicide) {S-Propyl butylethylthiocarbamate}, and 0.58

kg (a.i.) ha-1 pendimethalin (herbicide) {N-(1-ethylpropyl)-3,4-dimethyl-2,6-

dintro-benzenamine}. These pesticides were broadcast and then incorporated into the

soil by disking. The same pre-transplant pest management treatments were used in

1990, except that 56.0 L ha-1 of 1,3 dichloropene (nematicide) was used rather than

fenamifos. Acephate (0.83 kg (a.i.) ha-') {O,S-Dimethyl acetylphosphoramidothiate}

was used as needed to control foliage-feeding insects.

Fertilization for both years consisted of 448 kg ha-' of a 6-6-18 (N, K20, P20s)

fertilizer formulated for tobacco and 168 kg ha-' 15-0-14 (sodium-potassium nitrate)

at transplanting, 448 kg ha-' 6-6-18 at first cultivation, and 448 kg ha-1 6-6-18 at

last cultivation. All fertilizer was banded to the sides of the plants. Total application of

the primary and secondary nutrients was 106 kg ha-' N, 81 kg ha-1 P205, 266 kg ha-1

K20, 54 kg ha-' Ca, 54 kg ha-1 Mg, and 148 kg ha-' S. In 1990, due to leaching of N and

K, 112 kg ha-' of 15-0-14 was hand-applied to plots three weeks after the last

cultivation.

Leaf position (LP) will be defined as the leaves associated with a consecutive

grouping of nodes along the stem. To maintain the integrity of a designated LP, all plants

were topped to 21 leaves (each plant's leaves were counted). Suckers (axilary buds)

were chemically controlled by pouring 25 ml of a solution consisting of 20 ml L-1

flumetralin (2-chloro-N-[2,6-dinitro-4-(trifluoro-methyl)phenyll-N-ethyl-6-

flouro-benzene-methanamine) and 40 ml L-1 fatty alcohol (hexanol 0.5%, octonol 42%,









decanol 56%, dodecanol 1.5%) down the stalk at topping. The 21 leaves were

partitioned into seven, 3-leaf stalk positions for harvesting purposes.

A normal harvest study, so called because it most closely approximated the

normal production practices of flue-cured tobacco, was used to evaluate the responses of

the leaves when they were mature. A whole plant harvesting study was used to evaluate

the responses of individual LPs over time. These two studies will be described

separately because of the differences in design and layout in the field and the methods by

which the data were managed.

Normal Harvest Study

Ten plants comprised a plot and each plot was replicated four times each year.

Seven weekly leaf harvests were taken from 3-leaf stalk positions, beginning one week

after topping. Weybrew et al. (1984) suggested that flue-cured tobacco leaves should

mature at a rate of one leaf every two days. Based on their statement, harvesting three

leaves per week should ensure that mature tobacco was harvested each week. The lowest

three leaves were taken first, and subsequent harvests progressed up the plant until the

final harvest of the last three leaves that remained on the plants. The lowest three

leaves were designated LP 1-3, and uppermost three leaves designated LP 19-21, with

appropriate designations for the intermediate leaves.

Thirty leaves constituted a harvest on a given date (three leaves from each of 10

plants). On harvest days, 15 of the 30 leaves were measured for leaf area using a Li-

Cor 3100 leaf area meter. The 15 leaves for leaf area measurement always came from

the same five plants to minimize variation between plants and to ensure that the 15

leaves were composed of an equal number of leaves from each of the three leaf groups per

plant. To facilitate accurate leaf area measurement, individual leaves were severed

down the midrib and each leaf half was passed through the leaf area meter. Once leaf area

measurement was completed on the 15 leaves, fresh weight was determined using an

Ohaus 8000 digital scale. Fresh weight was also taken on the intact 15 leaves. Once leaf









area and fresh weight measurements were completed, the leaves were cured in the

normal flue-curing manner. The cut and intact samples were kept separate in the

curing barn so that the cured weights could be obtained on the leaf area samples for

specific cured leaf weight calculation. Once the leaves had been cured, the cut and intact

leaves were measured for cured weight. These weights were composite for the total plot

yield of cured tobacco.

Cured (intact) leaves from all samples were evaluated by United States

Department of Agriculture tobacco graders. Each sample was assigned a grade. These

grades were used to determine the value per hectare or kilogram, using the 1989 or

1990 Georgia-Florida flue-cured tobacco (Type 14) market average for the respective

grades. The numerical index for each grade (Bowman et al., 1988) was used to

determine a grade index for each LP. The grade index reported in this study for each LP

is the mean of the grade indices for the tobacco that was in the designated LP. Eight cured

leaves were selected from each sample of each LP for chemical or mineral analyses. The

midribs were removed from the lamina. The lamina was ground to 1 mm using a Wiley

mill. Total N, nicotine, and reducing sugar analyses were performed by R.J. Reynolds

Tobacco Company in Winston-Salem, N.C. For Ca, Mg, K, and P analyses, samples were

prepared by the ashing and acid digestion method described by Walsh (1971). The

resulting solutions were analyzed by Inductively Coupled Argon Plasma (ICAP) in the

University of Florida's Institute of Food and Agricultural Science Extension Soil Testing

Laboratory by methods described by Hanlon and Devore (1989).

A split-split-plot design was used for statistical analysis with years being main

plots, cultivars being sub-plots, and leaf positions being sub-sub-plots. Analysis of

variance and Fisher's Least Significant Difference (LSD) were carried out by methods

described by Gomez and Gomez (1984). For most variables, interactions were either

not significant, or were determined not to be of practical importance. No differences

between cultivars or years were found. The data reported were averaged across









cultivars and years. For clarity, only the LP effects are discussed because they were of

the greatest significance for each variable.

Whole Plant Harvest Study

This study was designed to monitor changes in LP characteristics for the duration

that each leaf position remained on the plant prior to normal harvest. Seven weekly

harvests were taken of all LPs that remained on the stalk, beginning one week after

topping. A set of samples was taken at topping to establish a baseline for all LPs.

Because this study was designed to monitor LP changes over time, LPs that were

harvested in the normal harvest study were removed from all plots that were to be used

for future harvest date analysis. The progression of the LPs as they were harvested is

illustrated in Table 2-1.


Table 2-1. The leaf positions removed per plant by harvest date in the days after
topping study.


--.-------------------

0 7
---------------------
19-21 19-21
16-18 16-18
13-15 13-15
10-12 10-12
7-9 7-9
4-6 4-6
1-3 1-3x


days
14
leaf
19-21
16-18
13-15
10-12
7-9
4-6x


after
21
positions
19-21
16-18
13-15
10-12
7-9x


topping
28 35
harvested
19-21 19-21
16-18 16-18
13-15 13-15x
10-12x


---------------------

42 49
----------------------

19-21 19-21x
16-18x


***Bold type indicates that leaf area, leaf mineral and chemical components, and
total non-structural carbohydrates (TNC) were measured on those leaf positions.
x Indicates leaf positions which were harvested normally on this date.

The data discussed herein were extracted from a larger study dealing with the

effects of lower leaf harvesting options on the same characters described in this paper.

Analysis of variance revealed that three treatments: 1) harvest all 21 leaves, 2) prune









the lowest 3 leaves, and 3) do not prune or harvest the lowest three leaves, had few if

any differences (discussed in Chapters 3 and 4). Most flue-cured tobacco farmers

practice one of these three treatments. Because of the lack of differences between

treatments, the data from the three treatments were combined, thereby increasing

sample size from 12 to 36 observations.

Each plot consisted of two plants occupying an area of 1 square meter. All LPs

were removed from the designated plants for a given harvest date as illustrated in Table

2-1. Each LP harvested consisted of six leaves (2 plants X a 3-leaf position). Fresh

weight was taken on all LPs using an Ohaus 8000 digital scale. Leaf area was measured

using a Li-Cor 3100 leaf area meter on LPs 1-3, 7-9, 13-15, and 19-21. To

facilitate accurate leaf area measures, each leaf was severed down the midrib and each

leaf half passed through the leaf area meter. All leaf samples were cured in the normal

flue-curing manner. After curing, the dried leaf samples were rehydrated for handling

purposes, and cured weights were taken on all LPs.

The six leaves from LPs 1-3, 7-9, 13-15, 19-21 of each plot were used for

leaf chemical or mineral analyses. The samples for chemical or mineral analyses were

processed the same as described in the normal harvest study above.

Total non-structural carbohydrate (TNC) data reported herein were derived

from separate plants than those used for leaf mineral and chemical analyses, but the

progression of harvesting LPs was identical to that indicated in Table 2-1. The TNC

samples were obtained from a single plant that was adjacent to those used for the

agronomic and mineral analyses. Because a single plant was used, total sample size was

three leaves per LP harvested. The lamina was severed from the midrib and placed in a

paper bag. The sampling and cutting of leaves was done in the field. When the samples

were obtained, bags containing samples were placed in a large plastic bag in a cooler

containing ice. This procedure was followed so that the leaf metabolism would be slowed,

conserving as much of the leaf carbohydrate as possible. Once all samples had been









taken, samples were transported to drying facilities. A Blue M forced air dryer was used

to desiccate samples. Prior to sampling, the dryer was set to 100 C, so that the

temperature would be at the desired level upon arrival at the drying facility. The paper

bags containing the leaf lamina were quickly placed in the dryer. The temperature was

maintained at 100 C for 1 hr to arrest lamina metabolism. After 1 hr at 100 C, the

temperature was reduced to 70 C for the duration of lamina drying. This drying method

was the best alternative to freeze-drying (Heberer et al., 1985). The elapsed time

between sampling initiation and arrival at the drying facilities was generally between 1

and 2 hr, depending on the number of samples that were to be harvested. Sample

acquisition always progressed from replication 1 to replication 4, so that any effects due

to sampling time could be accounted for. There were no appreciable differences in

replications indicating samples were processed in a timely manner. The TNC was

analyzed by a modified procedure of the methods described by Smith (1974).

Leaf chemical and mineral data are reported for leaf lamina only. Midribs were

not evaluated, and previous studies indicated that their chemical and mineral

constituents were different from the lamina (Darkis et al., 1952).

The treatments were arranged in a split-split-split-plot design in the field with

cultivars being main plots, lower leaf harvesting options being sub-plots, leaf position

being sub-sub-plots, and harvest dates being sub-sub-sub-plots. Analysis of the entire

data set was not possible due to lack of balance. With each harvest date there was a loss

of one LP (Table 2-1). Consequently, for harvest date evaluation, each harvest date was

analyzed individually. The statistical model was also a split-split-split-plot design

with years being main plots, cultivars being sub-plots, treatments being sub-sub-

plots, and leaf positions being sub-sub-sub-plots. The LSD values for LP comparison

given in the tables were calculated from the individual harvest date analysis of variance.

Data from each LP were regressed separately over the time period during which the

given LP remained on the plant. The model used was the same as that used for the








individual harvest date analysis except that harvest dates (days after topping) were

sub-sub-sub-plots. Analysis of variance, Fisher's Least Significant Difference (LSD)

and regression analysis of LP means were carried out by methods described by Gomez and

Gomez (1984). All regression equations reported were significant at the five-percent

level or greater (p>0.05). For most variables, interactions (except for leaf position X

harvest dates) were either not significant, or were determined not to be of practical

importance. For clarity, only the leaf position X harvest date effects are discussed

because they were of the greatest significance for each variable.

Results and Discussion

Each 3-leaf LP comprises 14.3% of the 21 total number of leaves that were

evaluated per plant. Fresh weights (FW) observed at the seven LPs during the normal

harvesting scheme were similar except for those at LPs 1-3 and 19-21 (Fig. 2-1).

Fresh weight accumulation of a given LP over time was dramatically influenced by leaf

age at topping (Fig. 2-2a and Fig. 2-2b.) At topping, the oldest leaves are those nearest

the bottom of the plant (i.e. LP 1-3), and leaf age declines progressively toward the the

top of the plant. The youngest and least mature LP at topping, 19-21, was initially the

lowest in FW, but increased in FW the most over time. The more mature LPs increased

the least in FW over time. The FW of the uppermost three LPs (or nine leaves), being

the least mature at topping, increased over time (Fig. 2-2a). However, the FW of the

lowermost four LPs (or 12 leaves) did not increase over time (Fig. 2-2b). This

observation quantified the suggestion by Wolf and Gross (1937) that more mature

leaves at topping would have fewer changes over time.

The cured weight (CW) of leaves harvested when they were mature was

influenced more by LP than was FW (Fig. 2-3). The highest two LPs, 16-18 and 19-

21, contributed 37.1% of the total cured weight of 4141 kg ha-1, while the lowest two

LPs, 1-3 and 4-6, contributed only 19.9% of the total CW. Under good growing

conditions like those encountered in the present study, the upper leaves typically will












19-21

16-18
.-
13-15
CL
0
10-12

7-9

4-6

1-3 LSD

0 500 1000 1500 2000 2500 3000 3500 4000 4500 5000
Fresh Leaf Weight (kg/ha)
Fig. 2-1. Fresh weight of flue-cured tobacco as influenced by leaf position (LP) (LSD
0.05 = 193). Vertical line represents mean across all LPs. Numerical values
within bars are means (n=16) of each LP and (%) is that LPs contribution to
the total yield of 26 317 kg/ha.










"V


400




300-


100 I I I I I I i I
0 7 14 21 28 35 42 49

Days After Topping (DAT)
Fig. 2-2a. Fresh weight of flue-cured tobacco upper stalk leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
21, 28, 35, and 42 DAT are 30, 16, 14, 14, 15, 10, and 15, respectively.



,nf-


340-


320-




300-


.. -.--o



i-----------------cI'- -


-*'-"- LP 1-3 Means (n=36)
--*- LP 4-6 Means (n=36)
--- -- LP 7-9 Means (n=36)
- -4* LP 10-12 Means (n=36)


4.
s
r


lzU 1 I I 1 1
0 7 14 21 28

Days After Topping (DAT)
Fig. 2-2b. Fresh weight of flue-cured tobacco lower stalk leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
and 21 DAT are 30, 16, 14, and 14, respectively.


* LP 13-15 Means (n-36)
0 LP 16-18 Means (n=36)
* LP 19-21 Means (n=36)

C .-***" """"" **










S"- LP13-15 y = 272.3 + 1.86x R^2 0.87
LP 16-18 y = 239.1 + 3.55x R^2 = 0.94

---- LP 19-21 y = 148.5 + 12.2x 0.14x^2 R^2= 0.96


r-










19-21

16-18

0
13-15
0
S10-12

7-9


4-6

1-3

0 100 200 300 400 500 600 700 800 900 1000
Cured Leaf Weight (kg/ha)
Fig. 2-3. Cured weight of flue-cured tobacco as influenced by leaf position (LP) (LSD
0.05 = 40). Vertical line represents mean across all LPs. Numerical values
within bars are means (n-16) of each LP and (%) is that LP's contribution to
the total yield of 4141 kg/ha.









contribute the highest CW of all leaves (Brown and Terrill, 1972; Darkis et al., 1936,

1952). Over time after topping, the CW increases observed for individual LPs were a

result of the Inherent differences in age of the leaves which comprised that LP (Fig. 2-

4a and 2-4b). The influence of leaf age on CW increases over time was similar to that

previously discussed for FW, except that the CW of LP 10-12 also increased with time

after topping (Fig. 2-4b). Only the lowest three LPs (or nine leaves) did not

significantly increase in CW over time.

Cured weight yield (CWY) is the percentage cured leaf that results from a given

amount of freshly harvested leaf after the curing process. Cured weight yield generally

increased from the lowest LPs to the highest LPs (Fig. 2-5). The CWY of other tobacco

crops was also shown to increase as LP increased (Suggs et al., 1987). It is desirable to

have a high CWY because the costs per unit of curing and handling are reduced due to the

higher return of salable product. The CWY for an individual LP over time was driven by

the differential in leaf age of the LPs at topping. The CWY response to time after topping

was similar to that observed with CW in that the upper four LPs (or 12 leaves)

increased in CWY over time (Fig. 2-6a), while the the lowest three LPs (or nine

leaves) did not (Fig. 2-6b). Handling costs per LP would be similar on a FW basis.

However, because CWY and CW increased progressively with higher LPs, the net returns

per LP likely would increase as LP advanced up the plant.

Leaf area was highest at the lower LPs and lowest at the highest LPs (Fig. 2-7).

This leaf area distribution is expected since the lowest leaves are formed prior to or

shortly after the seedlings are transplanted. The first leaves formed in the field after

transplanting are normally the largest with the upper leaves being smaller giving most

flue-cured tobacco cultivars their characteristic conical shape. Lower topping heights

than those used in this study can result in larger upper leaves, and some cultivars

produce large upper leaves. Turner and Incoll (1971) reported 80% of the leaf area of

a tobacco plant was contained in the 20 cm to 80 cm sections on the stalk. The stalk











bU


E 55-


50


450


40


S35------ LP 1-3 means (n=36)
*-* LP 4-6 means (n=36)
---- LP 7-9 means (n=36)
30- ,-,
0 7 14 21
Days After Topping (DAT)
Fig. 2-4a. Cured weight of flue-cured tobacco lower stalk leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, and
14 DAT are 3, 2, and 2, respectively.


II I I I I I I I
0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-4b. Cured weight of flue-cured tobacco upper stalk leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs 0, 7, 14,
21, 28, 35, and 42 DAT are 3, 3, 2, 3, 3, 3, and 4, respectively.


A LP 10-12 Means (n=36)
* LP 13-15 Means (n=36) toU0*0 t"" i
o LP 16-18 Means (n=36) ,,t* t 00"
* LP 19-21 Means (n=36) 1,* 0
O








........ LP 10-12 y =44.8 + 0.50x RA2 =0.86
S-""""" LP13-15 y = 40.9 + 0.60x RA2=0.79
LP16-18y=33.1 +1.01x RA2=0.91
LP 19-21 y = 17.8 + 2.60x 0.05x2 R^2 = 0.97


!


I












19-21

S16-18

13-15
O.
10-12

-J 7-9


4-6

1-3

0 2 4 6 8 10 12 14 16 18 20
Cured Weight Yield (g cured leaf/100 g green leaf)
Fig. 2-5. Cured weight yield of flue-cured tobacco as influenced by leaf position (LP)
(LSD 0.05 = 0.7). Vertical line represents mean across all LPs. Numerical
values within bars are means (n=16) of each LP.
















0





CO
.a



00


14 ..--............ -.. ....
aT a -*"*ieelu


11 1 I I I
0 7 14 21
Days After Topping
Fig. 2-6a. Cured weight yield of flue-cured tobacco lower stalk leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, and
14 DAT are 1.0, 0.4, and 0.5, respectively.


a LP 10-12 Means (n.
* LP 13-15 Means (n
o LP 16-18 Means (n.
* LP 19-21 Means (n


111


----


.36)
Ws) 0
=36) a






L 1y 1401



LP 10-12 y = 14.16+0.11x R^2 = 0.78
LP 13-15 y = 15.09 + 0.09x R2 = 0.59
LP 118 y = 14.35 + 0.13x R2 = 0.80
LP 19-21 y = 13.09 + 0.27X 00002xA2 R^2 0.93


Ir I r I I 1 I
0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-6b. Cured weight yield of flue-cured tobacco upper stalk leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
21, 28, 35, and 42 DAT are 1.0, 0.4, 0.5, 0.5, 0.8, 0.5 and 0.5, respectively.


-W* -- LP 1-3 means (n=36)


- LP 4-6 means (n36)
-"*- LP 7-9 means (n=36)


.YiS




; a



03
U
bO
60
8-


r-
.r~cl










19-21

16-18
----------

= 13-15 LSD
0
10-12
CD
.j 7-9

4-6

1-3

0 1000 2000 3000 4000 5000 6000 7000
Leaf Area (sq. m/ha)
Fig. 2-7. Leaf area of flue-cured tobacco as influenced by leaf position (LP) (LSD 0.05
= 322). Vertical line represents mean across all LPs. Numerical values within
bars are means (n=16) of each LP.









heights for the plants used in the present study were about 100 cm. Each leaf should

occupy about 5 cm on the stalk. The middle five LPs, which contributed 74% of the total

leaf area, would fall in the 20 cm to 80 cm sections on the stalks of these plants. A true

measure of leaf area index (LAI) was not possible because the leaves were harvested as

they matured, however, the composite LAI was 4.32 for the data given here. Only LPs

1-3, 7-9, 13-15, and 19-21 were measured over time, but the leaf area development

was similar to the responses for the other parameters previously discussed. The leaf

areas for those LPs that were measured were influenced by leaf age and maturity over

time (Fig. 2-8).

Specific cured leaf weight (SCLW) is a measure of CW per unit of leaf area.

Specific cured leaf weight increased with each LP (Fig. 2-9), ranging from a low of

6.13 mg of CW cm-2 leaf area at LP 1-3 to a high of 13.69 mg of CW cm-2 leaf area at LP

19-21. The SCLW may be a useful measure for interpreting differences in CW between

LPs. Cured weight yield is simply a ratio of the CW from a given amount of FW. The

CWY is largely the difference in water concentration. However, SCLW measures the CW

per unit of leaf area thereby negating to some extent the influence of water concentration

between a group of leaves. At LP 19-21, the CW was 2.36 times higher than that at LP

1-3, while the SCLW at LP 19-21 was 2.23 times higher than that at LP 1-3. Based on

the SCLW measures observed in this study, there may be a higher proportion of

structural tissue at higher LPs. Specific cured leaf weight of the individual LPs

responded to time (Fig. 2-10) in a similar manner as did the parameters previously

discussed.

Value per kilogram (Fig. 2-11) increased as LP progressed from the lowest to

the highest leaves on the plant. Value per hectare (Fig. 2-12), being a function of CW

and value per kilogram, also increased at higher LPs. The uppermost LP (leaves 19-

21) contributed 19.9% of the total value ($15 315 ha-1) of the tobacco crop, while the

lowest LP (leaves 1-3) contributed only 7.4% of the total value. Brown and Terrill






28




7000


6000


E 5000 -
E 50 ---- LP 1-3 Means (n=36)
A to^* --0-- LP 7-9 Means (n=36)
( 0' 0 LP 13-15 Means (n=36)
4000 LP 19-21 Means (n=36)


3000
LP 13-15 y = 5070+23x R^2 =0.64
LP 19-21 y= 2988 + 148x- 1.9x^2 RA2=0.94
2000 I I I II I
0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-8. Leaf area of flue-cured tobacco leaf positions (LP) in response to days after
topping. LSDs (0.05) for comparison of LPs at 0, 7, 14, and 21 DAT are 503,
316, 253, and 218, respectively. DAT 28 and 35 are not significant.











-I


19-21

16-18

13-15

10-12

7-9


--A
LSD


I I I I I I I I
0 2 4 6 8 10 12 14
Specific Cured Leaf Weight (mg/sq. cm)
Fig. 2-9. Specific cured leaf weight of flue-cured tobacco as influenced by leaf position
(LP) (LSD 0.05 = 0.4). Vertical line represents mean across all LPs.
Numerical values within bars are means (n=16) of each LP.


I I
14 21


I 5
28 35


4I
42 49


Days After Topping (DAT)
Fig. 2-10. Specific cured leaf weight of flue-cured tobacco leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
21, 28, and 35 DAT are 0.48, 0.28, 0.34, 0.38, 0.56, and 0.47, respectively.


* LP 13-15 y = 8.02 + 0.08x R2 = 0.83
S LP 19-21 y = 6.61 + 0.28x 0.003x^2 R2 = 0.98










.^^V*.* -- 1- LP 1-3 Means (n=36)
-.--'O-- LP 7-9 Means (n=36)
0 LP 13-15 Means (n=36)
......... *0 LP 19-21 Means (n=36)


-..<


3x












19-21

16-18

13-15

10-12

7-9


4-6

1-3

3.00


4.00


Dollars per Kilogram
Fig. 2-11. Value per kilogram of flue-cured tobacco as influenced by leaf position (LP)
(LSD 0.05 = 0.06). Vertical line represents mean across all LPs. Numerical
values within bars are means (n=16) of each LP.


19-21

16-18

13-15

10-12

7-9


0 500 1000 1500 2000 2500 3000 3500
Dollars per Hectare
Fig. 2-12. Value per hectare of flue-cured tobacco as influenced by leaf position (LP)
(LSD 0.05 = 152). Vertical line represents mean across all leaf positions.
Numerical values within bars are means (n=16) of each LP and (%) is that LP's
contribution to the total value of 15 315 $/ha.









(1972) found the middle and upper leaves of flue-cured tobacco contributed the most

value to the total value of the crop. The higher value of the upper LPs would support the

suggestion that management practices should strive to protect the upper leaves from

pests late in the harvesting season so as to derive the maximum yield and value from

those leaves.

The numerical grade index of flue-cured tobacco (Bowman et al., 1988) is based

on the USDA flue-cured tobacco grading system which considers several leaf

characteristics including position on the stalk, maturity, color and quality of the cured

leaf. Based on the grades received for the tobacco evaluated in this study, the lowest five

LPs were equal in grade index with the upper two LPs being lower in grade index (Fig.

2-13). The grade index values reported for the lower five LPs indicated ripe and

mature tobacco, while the upper leaf grade index values were indicative of unripe and/or

immature leaf. Suggs (1986) reported a high correlation of grade index to unit price of

tobacco crops. However, based on the data for grade index and value per kilogram by LP

reported in this study there was a poor relationship between price and grade index

across LPs. The grade indices typically reported are a weighted average across all LPs.

Higher quality tobacco, and subsequently higher value tobacco will have a higher grade

index. However, the prices paid per grade of tobacco from a given LP are determined by

market demand, and may not correlate as well with the grade indices from that LP.
Leaf Chemical Characteristics

Lamina N concentration (Fig. 2-14) generally increased as LP progressed up the

stalk. Brown and Terrill (1973), Nel et al. (1974), Neas et al. (1978), Darkis et al.

(1936, 1952) also found N concentration increased as LP advanced up the stalk. The age

of the leaves contained within a given LP influenced the N concentration of that LP over

time (Fig. 2-15). The least mature LPs at topping were the highest in N concentration,

but as those immature LPs increased in CW and leaf area, the N concentration declined

rapidly. Raper and McCants (1966) reported that flue-cured tobacco plants had












19-21

16-18 LS

0. 13-15

L. 10-12

-1 7-9

4-6

1-3 &

0 10 20 30 40 50 60
Grade Index (not weighted)
Fig. 2-13. Grade index of flue-cured tobacco as influenced by leaf position (LP) (LSD
0.05 = 7.7). Vertical line represents mean across all LPs. Numerical values
within bars are means (n=16) of each LP.











19-21

16-18

13-15

10-12

7-9


4-6

1-3

0 2 4 6 8 10 12 14 16 18 20 22 24 26
Lamina Nitrogen Concentration (g/kg)
Fig. 2-14. Lamina N concentration of flue-cured tobacco as influenced by leaf position
(LP) (LSD 0.05 = 1.8). Vertical line represents mean across all LPs.
Numerical values within bars are means (n=16) of each LP.

50 I


10 I I I I I 1
0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-15. Lamina N concentration of flue-cured tobacco leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
21, 28, and 35 DAT are 3.1, 2.6, 1.0, 1.7, 1.4, and 0.6, respectively.


" '" LP 7-9 y = 23.95 0.77x + 0.023x^2 R^2 = 0.99
LP 13-15 y= 33.10-1.10x + 0.019x^2 R2 = 0.97
LP 19-21 y = 47.07 1.32x + 0.017x^2 R2 = 0.99

----I-- LP 1-3 Means (n=36)
O LP 7-9 Means (n=36)
,, a LP 13-15 Means (n=36)
S0* LP 19-21 Means (n=36)

r 4.-- A=s ups..o- s- sW








accumulated 89.5% of the total N uptake 77 days after the seedlings were transplanted.

This period of time approximates the topping time in the present study. The total N

uptake was not measured in the present study. However, the rapid decline in N

concentration at the immature LPs may indicate that most of the N had been taken up by

the plants at topping and was diluted as the leaves expanded and accumulated dry matter.

Nicotine concentration (Fig. 2-16) was influenced by LP. Nicotine concentration

declined from the lowest LP to the middle LPs, then increased again to the upper LPs.

This nicotine concentration by LP pattern was similar to that reported by Walker

(1968) and Nel et al. (1974), however, other studies dealing with flue-cured tobacco

found nicotine concentration increased with higher LPs (Brown and Terrill, 1973; Neas

et al., 1978; and Darkis et al., 1936, 1952). Nicotine concentration increased with

leaf maturity (Fig. 2-17). Srivastava et al. (1984) reported that nicotine

accumulation of Dixie Shade Wrapper tobacco was only 18.4% of the total accumulation

70 days after seedlings were transplanted. Nicotine is synthesized in the roots (Dawson

and Solt, 1959), and the topping process is believed to stimulate root growth (Steinberg

and Jeffery, 1957). Nicotine concentration increase in the tobacco leaf is a most

important response to topping and leaf maturity (Steinberg and Jeffery, 1957; Moseley

et al., 1963; Walker, 1968; Elliot, 1975). Topping established a set leaf number per

plant and assuming root growth was stimulated, nicotine concentration should increase

as the leaves mature.

Total non-structural carbohydrates (TNC) are the energy reserves the plant has

accumulated which can be used for new plant growth (Smith, 1981). The concentration

of TNC in leaves changed over time after topping (Fig. 2-18). The rapid increases in

TNC concentration at LPs 13-15 and 19-21 were probably due to both topping and leaf

maturity. The decline in the TNC concentration at the LPs as the final harvest neared

was likely due to a reduction in the photosynthetic capacity of the ageing leaves in

combination with some respiration of the TNC fraction due to maturity, ripening, and













-H


19-21

16-18

13-15

10-12

7-9


LSD
LSD


0 2 4 6 8 10 12 14 16 18 20 22 24
Lamina Nicotine Concentration (g/kg)
Fig. 2-16. Lamina nicotine concentration of flue-cured tobacco as influenced by leaf
position (LP) (LSD 0.05 = 1.4). Vertical line represents mean across all LPs.
Numerical values within bars are means (n=16) of each LP.

I


--- -- LP 1-3 Means (r
0 LP 7-9 Means (n
0 LP 13-15 Means
LP 19-21 Means
...--*e~l


-=36)
1=36) ,-e, ---- -
(n=36) ,- w '
(n=36) ,










LP 7-9 y= 9.17 + 0.22x R2 = 0.94
LP13-15 y =7.11 +0.26x R^2=0.97
S LP 19-21 y = 4.00 + 0.60x 0.006xA2 R2 = 0.98


U l I I I I I I i
0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-17. Lamina nicotine concentration of flue-cured tobacco leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
21, 28, and 35 DAT are 0.9, 0.7, 0.4, 0.5, 0.6, and 0.7, respectively.


J


















131





to -- -- LP 1-3 Means (n=36)

*--0-- LP 7-9 Means (n=36)
S/ LP 13-15 Means (n=36)
S* LP 19-21 Means (n=36)


SI- LP 13-15 y 415.0 + 9.8x 0.19xA2 R^2 0.84
LP 19-21 y = 283.1 + 16.2x 0.30xA2 R2 = 0.90


I I


I I I I


0 7 14 21 28 35 42 45
Days After Topping (DAT)
Fig. 2-18. Lamina total non-structural carbohydrate concentration of flue-cured
tobacco leaf positions (LP) in response to days after topping. LSDs (0.05) for
comparison of LPs at 0, 7, 14, 21, 28, and 35 DAT are 70.3, 54.4, 49.8, 50.3,
37.5, and 24.3, respectively.


CO
,-



2 -
0


( D
00
z 0

0


0 0
C5o

CO
0d


J












senescence of the leaves. The leaves taken for the final harvests in the present study

were judged to be mature to ripe. Kakie and Sugizaki (1970) and Kakie (1972)

reported that flue-cured tobacco leaves declined in total carbohydrate concentration as

the leaves progressed from maturity to over-maturity. The total carbohydrate decline

was associated with a decrease in starch concentration and an increase in the soluble

sugar fraction. But, there was some respiration of starch because the soluble sugar

increase was not equal to the starch decrease.

Reducing sugar concentration is important to the quality of flue-cured tobacco.

The reducing sugars are a product of starch (mostly TNC) metabolism induced during the

curing process (Askew and Blick, 1947). Kakie (1972) reported that total

carbohydrate concentration of mature green tobacco leaves ranged from 340 g kg-' to

416 g kg-1 with a starch concentration ranging from 190 g kg-' to 370 g kg-1. The ratio

of starch to total carbohydrates was 56% to 89%. The higher proportion of starch

occurred when the leaves were progressing from immaturity to maturity, while the

lower proportion of starch occurred as the leaves were progressing from maturity to

over-maturity. Reducing sugar concentration of flue-cured tobacco leaves will vary

greatly depending on the location, cultivar, and ultimately seasonal climatic conditions.

However, a concentration range generally from 150 g kg-1 to 280 g kg-' has been

reported. The starch concentration of cured leaves is usually less than 30 g kg-1 (Neas

et al., 1978; Nel et al., 1974) if mature leaves are harvested and cured properly.

Reducing sugar concentration was highest in the leaves from the middle LPs (Fig.

2-19). The middle leaves of flue-cured tobacco are nearly always the highest in

reducing sugar concentration (Brown and Terrill, 1973; Nel et al., 1974; Neas et al.,

1978; Darkis et al., 1936, 1952; Walker, 1968). Over time after topping, the

reducing sugar concentration (Fig. 2-20) followed a similar pattern to that of TNC

concentration. This result would be expected because the TNC is metabolized to reducing

sugars in the curing process. The reducing sugar concentrations at LPs 13-15 and 19-












19-21

16-18

13-15

10-12

7-9


4-8

1-3 LSD

0 50 100 150 200 250 300
Lamina Reducing Sugar Concentration (g/kg)
Fig. 2-19. Lamina reducing sugar concentration of flue-cured tobacco as influenced by
leaf position (LP) (LSD 0.05 = 13.5). Vertical line represents mean across all
LPs. Numerical values within bars are means (n=16) of each LP.

S300- ."----.


.2 250





S150
---- ------ -- LP 1-3 Means (n=36)
e --.0-" LP 7-9 Means (n=36) &
100 LP 13-15 Means (n=36)
S* LP 19-21 Means (n=36)
cc 50 *
S'ca LP 13-15 y= 132.3 + 7.5x 0.14x2 R^2=0.88
E LP 19-21 y = 37.6 + 9.6x 0.16x^2 RA2 = 0.85

0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-20 Lamina reducing sugar concentration of flue-cured tobacco leaf positions
(LP) in response to days after topping. LSDs (0.05) for comparison of LPs at 0,
7, 14, 21, 28, and 35 DAT are 33.3, 10.6, 10.4, 15.2, 11.3, and 6.0,
respectively.









21 changed proportionally with the TNC concentration of the same LPs (Fig. 2-21). A

poor relationship of reducing sugar to TNC concentration was found at LPs 1-3 and 7-9.

The N to nicotine (N:Nic) and reducing sugars to nicotine (Sug:Nic) concentration

ratios have been used to evaluate the potential smoking quality of flue-cured tobacco

(Moseley et al., 1963; Tso, 1972). The N:Nic (Fig. 2-22) was influenced by LP. The

lower N:Nic ratio values at LPs 1-3, 4-6, 13-15, 16-18, and 19-21 were within the

ranges found of other tobaccos for similar LPs (Brown and Terrill, 1973; Darkis et al.,

1952; Walker, 1968), however, the higher N:Nic values for LPs 7-9 and 10-12

exceeded published values for similar LPs. Tso (1972) determined that an N:Nic

approximating 1:1 was the most desirable. The N:Nics of the LPs evaluated over time

after topping were different because of the differences in leaf age at topping (Fig. 2-

23). Maturation of tobacco leaves results in lower concentrations of N and higher

concentrations of nicotine (see Figs. 2-15 and 2-17). Moseley et al. (1963) reported

maturation of tobacco leaves resulted in lower N:Nic. The Sug:Nic ratio (Fig. 2-24) also

varied by LP. The Sug:Nic was highest at the middle LPs with lower Sug:Nic observed at

the upper and lower LPs. Brown and Terrill (1973) reported increasing Sug:Nic up to

the middle LPs, then a decline to the upper LPs, but their values for the middle LPs were

lower than the ones reported in the present study. Florida flue-cured tobacco is

typically higher in reducing sugar concentration and lower in nicotine than tobacco from

other states. Over time, each LP declined in the Sug:Nic (Fig. 2-25) indicating that

nicotine concentration was increasing proportionally higher than the reducing sugar

concentration. Moseley et al. (1963) found a reduction of the Sug:Nic was characteristic

of maturing tobacco leaves.










300
SReducing Sugar Concentration = -137.2 + 0.658 (TNC)
25 RA2 = 0.820
250-
I*C
200


8 150


CO 100
c *
50

S0 III I1
200 250 300 350 400 450 500 550 600
Total Non-Structural Carbohydrate Concentration (g/kg)
Fig. 2-21. Relationship (p=0.001) between the total non-structural carbohydrate
(TNC) concentration of rapidly dried leaf lamina and the reducing sugar
concentration of cured leaf lamina from flue-cured tobacco leaf positions 13-15
and 19-21.












19-21

16-18

13-15

10-12

7-9


Fig. 2-22.
by
all

20


10


0.00 0.25 0.50 0.75 1.00 1.25 1.50
Nitrogen to Nicotine Concentration Ratio
Lamina N to nicotine concentration ratio of flue-cured tobacco as influenced
leaf position (LP) (LSD 0.05 = 0.13). Vertical line represents mean across
LPs. Numerical values within bars are means (n=16) of each LP.


--*- LP 1-3 Means (n=36)
O LP 7-9 Means (n=36)
n LP 13-15 Means (n=36)
S LP 19-21 Means (n=36)


%m LP 7-9 y = 2.52 0.065x RA2 = 0.85
S .- LP 13-15 y = 5.14 0.29x + 0.005x^2 R^2= 0.97
4, ---- LP 19-21 y =12.60 0.73x + 0.011x^2 R^2 = 0.86
%-

sg


0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-23. Lamina N to nicotine concentration ratio of flue-cured tobacco leaf positions
(LP) in response to days after topping. LSDs (0.05) for comparison of LPs at 0,
7, 14, 21, 28, and 35 DAT are 2.35, 1.18, 0.18, 0.12, 0.08, and 0.07,
respectively.


v


%






42




19-21

16-18

o 13-15

CL 10-12

7-9

4-6

1-3 LSD

0 2 4 6 8 10 12 14 16 18 20 22
Reducing Sugar to Nicotine Concentration Ratio
Fig. 2-24. Lamina reducing sugar to nicotine concentration ratio of flue-cured tobacco
as influenced by leaf position (LP) (LSD 0.05 = 1.5). Vertical line represents
mean across all LPs. Numerical values within bars are means (n=16) of each
LP.


30
I-** LP 1-3 Means (n=36)
O LP 7-9 Means (n=36)
25- a LP 13-15 Means (n=36)
SLP 19-21 Means (n=36)
20


CO
| 15 =""" --. = ,,


10..........---,


5 """"" LP 7-9 y = 22.07 0.25x R^2 = 0.98
n" """" LP 13-15 y= 22.30-0.17x R"2=0.78
LP 19-21 y = 15.53 0.17x R^2 = 0.49
I I I I I I I

0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-25. Lamina reducing sugar to nicotine concentration ratio of flue-cured tobacco
leaf positions (LP) in response to days after topping. LSDs (0.05) for
comparison of LPs at 0, 7, 14, 21, 28, and 35 DAT are 3.6, 1.7, 1.4, 1.5, 1.3,
and 1.3, respectively.









Leaf Mineral Characteristics

Lamina Ca concentration (Fig. 2-26) was highest in the lowest LPs. This result

was expected because Ca is considered to be non-mobile in plants (Tisdale and Nelson,

1975). Calcium concentrations have been found to be highest in the lower leaves of

tobacco in previous studies as well (Darkis et al., 1936, 1952; Askew et al., 1947;

Walker, 1968; Neas et al.,1978). Calcium concentrations changed little over time after

topping for all LPs (Fig. 2-27). Raper and McCants (1966) found that flue-cured

tobacco plants had accumulated 81.8% of the total Ca uptake 77 days after seedling

transplantation. One possible interpretation of these data is that Ca uptake continued at a

fairly constant rate after topping because for the leaves within the given LPs to maintain

the Ca concentration in response to CW increases a considerable amount of Ca must be

taken up by the roots and translocated to the leaves. Raper and McCants (1966) based

their total uptake percentages on the composition of plants harvested 91 days after

seedlings were transplanted. Data collection for the present study continued until 130

days after the seedlings had been transplanted.

Lamina Mg concentration by LP (Fig. 2-28) followed an almost identical pattern

as that of Ca. A similar distribution of Mg concentration in leaves has been reported by

Askew et al. (1947), Darkis et al. (1952), and Walker (1968). The Mg concentration

by LP over time after topping was constant (Fig. 2-29), much like that observed for Ca.

Raper and McCants (1966) reported that 77 days after the seedlings were transplanted

tobacco plants had accumulated 90.0% of the total Mg that was accumulated after 91 days

in the field. Because Mg and Ca concentration patterns, not the actual concentrations,

were similar over time, the same argument previously made about Ca concentration over

time should be valid for Mg. It also could be argued, based on the observations in the

present study, that had Raper and McCants (1966) continued to take samples after 91

days after transplanting the total uptake of Mg and Ca may have been different. This

suggestion is supported by observations by Srivastava et al. (1984) who found that





44




19-21

16-18

_o 13-15

n. 10-12

S 7-9 LSD

4-6

1-3

0 5 10 15 20 25 30 35
Lamina Calcium Concentration (g/kg)
Fig. 2-26. Lamina Ca concentration of flue-cured tobacco as influenced by leaf position
(LP) (LSD 0.05 = 1.4). Vertical line represents mean across all LPs.
Numerical values within bars are means (n=16) of each LP.

35
---U-- LP1-3Means(n=36)
"O" -- LP 7-9 Means (n=36)
o 30 --O- LP 13-15 Means (n=36)
c LP 19-21 Means (n=36)

25

8
o -. LP19-21 y=11.43=0.078x R-2= 0.54
w 20
20
--O
.. 15 .....-.........o.....




10
0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-27. Lamina Ca concentration of flue-cured tobacco leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
21, 28, and 35 DAT are 2.5, 0.6, 0.8, 0.8, 0.5, and 0.7, respectively.












19-21

16-18

13-15

C. 10-12


4 7-9

4-6

1-3

0 2 4 6 8 10 12 14 16
Lamina Magnesium Concentration (g/kg)
Fig. 2-28. Lamina Mg concentration of flue-cured tobacco as influenced by leaf position
(LP) (LSD 0.05 0.7). Vertical line represents mean across all LPs.
Numerical values within bars are means (n=16) of each LP.



----- LP 1-3 Means (n=36)
..., -0- LP 7-9 Means (n=36)
S**.0 LP 13-15 Means (n=36)
O* LP 19-21 Means (n=36)
C 12


S10-
Sio
O -- LP 13-15 y= 6.04 0.07x +0.0001x2 R2 =0.96
8 ---- LP 19-21 y 6.08-0.06x +0.0001x^2 R^2 0.48


E 6 a ,r,,



0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-29. Lamina Mg concentration of flue-cured tobacco leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
21, 28, and 35 DAT are 1.3, 0.3, 0.3, 0.4, 0.2, and 0.2, respectively.









Dixie Shade Wrapper tobacco had taken up only 54.2% and 42.7% of the total Ca and Mg,

respectively, 80 days after seedling transplantation. They compared the percentage

uptake to final harvest values taken 120 days after transplanting.

Lamina K concentration (Fig. 2-30) was highest at the lower LPs and lowest at

the upper LPs. Potassium is generally considered to be a mobile element in plants

(Tisdale and Nelson, 1975). However, the fact that lower K concentrations in upper

leaves than lower leaves have been reported by Darkis et al. (1936, 1952), Askew et

al. (1947), Walker (1968), and Neas et al. (1978) may indicate that K is not as

mobile in the tobacco plant as in other plants. The change in K concentration over time

was influenced by the age of the leaves within an LP (Fig. 2-31). Raper and McCants

(1966) reported that flue-cured tobacco plants had accumulated 97.4% of all K 77 days

after transplanting. Srivastava et al. (1984) found Dixie Shade Wrapper tobacco had

taken up only 55.4% of all K 80 days after transplanting. Atkinson et al. (1977) found

Burley tobacco plants had accumulated 95.1% of all K 73 days after seedling

transplantation. The tobacco plants in the present study may have accumulated most of

the K at topping because the K concentrations at the least mature LPs declined after

topping, while K concentrations at the more mature LPs remained about the same.

Lamina P concentration (Fig. 2-32) differed little at LPs. Differences existed

between some LPs, but unlike other leaf mineral components previously discussed, these

P concentration data differed from other published reports. Askew et al. (1947),

Darkis et al. (1952), and Nel et al. (1974) found P concentrations to be higher at

higher LPs. Of all the leaf parameters evaluated over time, the most dynamic

relationship existed for P concentration by LP (Fig. 2-33). These differences between

LPs existed due to the inherent differences in leaf age at topping. The least mature LP at

topping, 19-21, had the highest P concentration and most likely had the highest

metabolic activity. The most interesting aspect of the P concentration changes over time

was that the P concentrations at LPs 7-9, 13-15, and 19-21 increased over the 21












19-21

16-18

13-15

10-12

7-9


5 10 15 20 25


Lamina Potassium Concentration (g/kg)
Fig. 2-30. Lamina K concentration of flue-cured tobacco as influenced by leaf position
(LP) (LSD 0.05 = 1.6). Vertical line represents mean across all LPs.
Numerical values within bars are means (n=16) of each LP.


32
--- *-- LP 1-3 Means (n=36)
30 --0- LP 7-9 Means (n=36)
Ca D LP 13-15 Means (n=36)
28 LP 19-21 Means (n=36)
28-

26

24

S22-
cc
E
E 20 """*---
E.20 LP 13-15 y=27.0-0.19x R2 =0.86
LP 19-21 y = 30.97 0.45x + 0.005x^2 R2 = 0.95
18 ,,,,,,
0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-31. Lamina K concentration of flue-cured tobacco leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
21, 28, and 35 DAT are 3.1, 1.5, 1.5, 1.5, 0.8, and 1.3, respectively.


LSD
LSD



-I



-H


30
30












19-21


16-18


13-15


10-12


7-9


4-6


1-3

0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0
Lamina Phosphorus Concentration (g/kg)
Fig. 2-32. Lamina P concentration of flue-cured tobacco as influenced by leaf position
(LP) (LSD 0.05 = 0.3). Vertical line represents mean across all LPs.
Numerical values within bars are means (n=16) of each LP.


O 0
i


O
1


LP 1-3 Means (n=36)
LP 7-9 Means (n-36)
LP 13-15 Means (n=36)
LP 19-21 Means (n=36)
".""- LP 7-9 y = 2.75 + 0.029 x RA2 = 0.93
LP 13-15 y = 3.09 0.05x + 0.001xA2 R^2= 0.92
LP 19-21 y= 4.50 -0.14x 0.002x^2 RA2=0.98

%* %o o
*" -


0 7 14 21 28 35 42 49
Days After Topping (DAT)
Fig. 2-33. Lamina P concentration of flue-cured tobacco leaf positions (LP) in
response to days after topping. LSDs (0.05) for comparison of LPs at 0, 7, 14,
21, 28, and 35 DAT are 0.4, 0.2, 0.2, 0.2, 0.2, and 0.2, respectively.


5.0


4.5


4.0


3.5


3.0


2.5









days prior to final harvest. Because the leaves comprising LP 19-21 were the most

recently formed and were on the plant for the longest period of time, the most

information was accumulated for this LP. However, when viewing the concentration

curves for LPs 7-9 and 13-15, it may be suggested that had these LPs been sampled

over the same stages of development as was LP 19-21, their respective relationships

would look similar. This theory may be supported by the lowest P concentration points,

which were about 2.6 g kg-1, on the respective curves that were similar at all three LPs.

The final P concentrations also were nearly equal for each LP which agreed with the data

reported for the individual LPs in Fig. 2-32.
Conclusions

Flue-cured tobacco is harvested by LP as the leaves mature. Differences existed

for agronomic, chemical, and mineral characteristics based on the position or node on

the tobacco stalk at which leaves were formed. The highest yield and value were found at

the highest LPs. At topping, differences in the agronomic, chemical, and mineral

characteristics at LPs were a result of leaf age and previous development. The yield and

chemical and mineral concentrations of the lowest nine leaves did not change over time

after topping indicating that those leaves were essentially mature at topping. However,

the yield, chemical, and mineral characteristics of the upper 12 leaves (or 4 LPs) did

change over time, with the largest changes evident in the uppermost leaves. The

observation that characteristics of flue-cured tobacco leaves are different based on the

position or node at which the leaves are formed would support continuation of the

current practices by which leaves are harvested and marketed based on LP.













CHAPTER 3
LOWER LEAF HARVESTING EFFECTS ON AGRONOMIC
CHARACTERISTICS OF FLUE-CURED TOBACCO


Introduction

Federal programs dictate how much flue-cured tobacco (Nicotiana tabacum L)

can be marketed. A combination of area allotments and weight quotas is assigned to each

farm. New tobacco cultivars and improved cultural practices enable many tobacco

growers to exceed their weight quotas on less than the area allotted to produce the

tobacco. Consequently, some farmers choose not to plant their entire area allotment,

since they cannot sell the tobacco produced in excess of the weight quota. This

management decision is sound under good growing conditions, where yields can be

predicted with reasonable accuracy.

Lower leaf harvesting options are available that may allow farmers opportunities

to increase their net income from tobacco. If the upper leaves are substantially more

valuable in price than the lower leaves, the farmer may wish to produce his weight quota

with the upper leaves. Consequently, the lower leaves could be discarded.

Lower leaf pruning of flue-cured tobacco became an issue in the mid 1970's

when an over-supply of lower stalk tobacco pushed prices downward causing receipts of

this type tobacco to be increased by the farmer-owned Flue-Cured Tobacco Cooperative

Stabilization Corporation (FCTCS) (Wemsman and Matzinger, 1980). The FCTCS

purchases tobacco that does not receive bids of more than one cent above the designated

United States Department of Agriculture price support. Because of the large receipts of

this lower leaf tobacco, the federal tobacco program was amended in 1978 to allow









farmers to plant 10% more acreage if they would prune or not harvest the lowest four

leaves of their tobacco crop. This program was known as the "four leaf program."

Studies dealing with lower leaf harvesting options have resulted in mixed

conclusions with respect to effects on yield. Currin and Pitner (1980) reported that

pruning and discarding the lowest four leaves did not lower yields or affect the leaf

chemical component balance of the remaining leaves, when compared to harvesting all

the leaves. Leaving the lowest four leaves on the stalk and not harvesting them, reduced

yield compared to the treatments involving either pruning and discarding the lowest four

leaves or harvesting all leaves. Stocks (1988) found that pruning and discarding the

lowest three or four leaves resulted in equal yields and an increase in the price per

kilogram when compared to harvesting all leaves. But, not removing and not harvesting

the same lower leaves reduced yields. Currin and Stanton (1989) and Stocks (1988)

reported that pruning more than four leaves had a negative impact on yield and value of

flue-cured tobacco.

Court and Hendel (1989) evaluated leaf number management by several topping

height and lower leaf pruning regimes. Eighteen leaves per plant was the control.

Reduction in leaf number harvested from 18 to 15 to 12 resulted in a reduction in yield

and value of the crop for each reduction in leaf number. Pruning the lower leaves or

topping down to the desired leaf number produced similar reductions in yield and value.

Suggs (1972) evaluated a wide array of leaf pruning treatments and concluded that yield

and value decreased with an increase in the number of leaves pruned. However, the

lowest three leaves could be pruned with little decrease in yield. Crop value declined as

the number of lower leaves pruned exceeded three. Wernsman and Matzinger (1980)

found that yield reductions due to lower leaf pruning of mammoth cultivars could be

compensated for by topping to a higher leaf number.

In the above-mentioned studies, there are discrepancies regarding the effects of

lower leaf pruning. Results from studies conducted in the Southeastern U.S. (Suggs,









1972; Currin and Pitner, 1980; Stocks, 1988; Currin and Stanton, 1989) suggested

that pruning the lowest three or four leaves of flue-cured tobacco did not reduce yield or

value appreciably. However, in a study conducted in Southern Canada, Court and Hendel

(1989) did find yield losses associated with lower leaf pruning. The present study

evaluated the influence of five lower leaf harvesting options on tobacco plants topped to

exactly 21 leaves. Because results from some previous work suggests that the tobacco

plant compensates in some manner for the cured weight lost due to lower leaf pruning,

the 21 leaves on each plant were partitioned into seven, 3-leaf stalk positions so that

yield by leaf position could be evaluated to determine where any "yield compensation"

may occur.

Materials and Methods

The location, cultivars, and management practices for the present study were

identical to those previously discussed in Chapter 2. Only methods used in the present

study that differ from those discussed in Chapter 2 are discussed below.

Five lower leaf harvesting options were evaluated in this study. These

treatments were as follows: 1) harvesting all 21 leaves (LPs 1-3 to 19-21)(control),

2) prune and discard the lowest three leaves on the stalk, harvest 18 leaves (LPs 4-6 to

19-21), 3) do not prune or harvest the lowest three leaves, harvest 18 leaves (LPs 4-

6 to 19-21), 4) prune and discard the lowest six leaves on the stalk, harvest 15 leaves

(LPs 7-9 to 19-21), and 5) do not prune or harvest the lowest six leaves, 15 leaves

harvested (LPs 7-9 to 19-21). Leaves were removed from plants of the pruning

treatments at topping when leaf number was established. Currin and Pitner (1980)

found pruning the lower leaves at early topping produced the highest yields. The topping

time used in the present study would be considered early topping. The pruned leaves

were cured in the normal manner to evaluate what their contributions to yield and value

would have been.









A split-plot arrangement of treatments was utilized in the field with cultivars

being main plots and lower leaf harvesting treatments being sub-plots. But, the addition

of the leaf position variable resulted in a split-split-plot design with leaf position being

the sub-sub-plot. Four replications of the cultivars and lower leaf harvesting

treatments were evaluated each year.

A split-split-split-plot design was used for statistical analysis with years being

main plots, cultivars being sub-plots, lower leaf harvesting options being sub-sub-

plots, and leaf positions being sub-sub-sub-plots. Analysis of variance and Fisher's

Least Significant Difference (LSD) were carried out by methods described by Gomez and

Gomez (1984). The factors of primary interest in this study were lower leaf

harvesting options and leaf positions, and their interaction. There were no significant

interactions between harvesting options and leaf position, year, or cultivar for the

agronomic variables, so the following discussion focuses on the effects of harvesting

options effects across years and cultivars. Total yields were computed by summing

yields of all leaf positions and analysis of variance was performed on all data to

determine significant effects due to harvesting options. A split-split-plot design with

years being main plots, cultivars being sub-plots, and lower leaf harvesting options

being sub-sub-plots was used for the analysis. A second analysis was also performed on

yield data which included weights of pruned leaves in order to evaluate the contributions

of the pruned leaves. The LSD values computed from the two statistical analyses were

very similar. Therefore, another analysis including the two treatments with the pruned

leaves added was carried out. No differences were found in how the harvesting options

were segregated by means comparison from the individual analyses. For simplification,

the LSD values given in the figures were computed from the combined analysis of

treatments, otherwise two LSD values of numerical value would have been necessary.









Results and Discussion

Lower leaf harvesting treatments did not influence the fresh weight, cured

weight, leaf area, or value at the LPs above those that were either pruned or were left on

the plant and not harvested (data not shown). Suggs (1972), Currin and Pitner

(1980), and Stocks (1988) found equivalent yields and economic values from

treatments involving pruning of the lower three or four leaves or harvesting all of the

leaves. Furthermore, Currin and Pitner (1980) and Stocks (1988) found that leaving

the lowest three or four leaves on the plant and not harvesting them reduced yields and

values when compared to pruning and discarding those leaves or harvesting them

normally. From those studies it may be inferred that pruning treatments increased

yield and economic value in the leaves above those pruned, while not pruning or

harvesting the lower leaves did not influence the yield or values at the higher LPs.

Fresh weight (FW) (Fig. 3-1), cured weight (CW) (Fig. 3-2), and leaf area

index (LAI) (Fig. 3-3) were reduced as a result of the lower leaf harvesting treatments.

Pruning or not pruning per se had no effect on these parameters; rather, the number of

harvested leaves per treatment was responsible for the observed results. Court and

Hendel (1989) also noted that reduced leaf number per plant reduced yield parameters,

either by lower leaf pruning or topping to a lower leaf number.

In the present study, the pruned leaves were saved and cured normally to

evaluate the effects of the early harvesting by pruning compared to normal harvesting of

the same leaves. When the FW, CW, and LA of the pruned leaves were added back to the

yields of the corresponding pruning treatments (Figs. 3-1, 3-2, 3-3), no differences

in FW, CW, or LAI were found between the pruning treatments (computed with 21 total

leaves) and the control treatment (21 leaves harvested). This observation supports the

view that at topping the lower leaves which were pruned had reached maturity. In

Chapter 2, it was also noted that the lowest nine leaves had reached maturity at topping.










No Prune Low| Actual Treatment Means [ Pruned Leaves Added
6 Leaves 19190


Prune Low
6 Leaves

No Prune Low
3 Leaves

Prune Low
3 Leaves

Hanrvat All


19350


23600


23440


Leaves 26320
I I I
10000 15000 20000 25000
Fresh Weight (kg/ha)
Fig. 3-1. The effects of lower leaf harvesting on fresh weight of flue-cured tobacco
and the effects of the pruned leaves on the total yield of the pruning treatments.
LSD (0.05) = 1197 for all comparisons. Values outside bars are the treatment
means.


No Prune Low I Actual Treatment Means El Pruned Leaves Added
6 Leaves 3350


Prune Low
6 Leaves


No Prune Low
3 Leaves

Prune Low
3 Leaves

Harvest All


3360


1


3810


3820


4


4180






1138


Leaves 4150
I I I I I
2000 2500 3000 3500 4000 4500
Cured Weight (kg/ha)
Fig. 3-2. The effects of lower leaf harvesting on cured weight of flue-cured tobacco
and the effects of the pruned leaves on the total yield of the pruning treatments.
LSD (0.05) = 148 for all comparisons. Values outside bars are the treatment
means.


5910





26304


2


I










No Prune L Acual Treatment Means 0T Pruned Leaves Added
6 Leaves 3.11


Prune L.w
6 Leaves

No Prune Low
3 Leaves

Prune Low
3 Leaves

Harvest All
Leaves


3.13


3.88


3.82


14.34






S4.37



4.32
I


2.0 2.5 3.0 3.5 4.0 4.5 5.0
Leaf Area Index (sq. m leaf/sq. m land area)
Fig. 3-3. The effects of lower leaf harvesting on the leaf area index of flue-cured
tobacco and the effects of the pruned leaves on the leaf area index of the pruning
treatments. LSD (0.05) = 0.17 for all comparisons. Values outside bars are the
treatment means.









No previous studies dealing with the effects of lower leaf removal evaluated the leaves

that were pruned.

Harvest index (HI) is a ratio of the weight of the economic portion of a crop to the

total shoot biomass of the above-ground portion of the crop. The HI is typically used

with grain crops to evaluate the grain yield efficiency of a cultivar. A reduction in the

number of leaves harvested resulted in a decline in HI (Fig. 3-4). However, a higher HI

was found when the lower six leaves were pruned compared to not pruning or harvesting

the same six leaves. This result was due to the lower stalk cured weight when six leaves

were pruned (Fig. 3-5) since CWs were equal for the two treatments (Fig 3-2). The

difference in stalk weight was unexpected. Literature relating tobacco stalk weight to

such treatments does not exist. Kollman et al. (1974) found reduced sink size of

soybeans (Glycine max L. Merr.) increased stem and petiole weight. A maximum HI of

71 was found when no leaves were pruned and all leaves were harvested. As with other

agronomic parameters previously discussed, the addition of the CW of pruned leaves

resulted in His which were almost identical to the treatment where all leaves were

harvested (Fig. 3-4). A HI for tobacco by the definition given above has not been

reported, but Suggs (1984) found leaf weight to total leaf plus stalk weight ranged from

47% to 64%.

Cured weight yield (CWY) (Fig. 3-6) and specific cured leaf weight (SCLW)

(Fig. 3-7) increased as more lower leaves were pruned and discarded or were not

harvested. The average CWY and SCLW of the pruning treatments were not different

from the control when the corresponding CWY and SCLW of the pruned leaves were

included in the totals. In Chapter 2, the CWY and SCLW were found to be lowest in the

lowest LPs. Consequently, the average CWY and SCLW of the total crop would be expected

to increase as the number of lower leaves harvested decreased. Suggs et al.(1987) found

the CWY of the first harvest of a tobacco crop was the lowest of all harvests. This first

harvest also required one of the longest curing times. More importantly, the ratio of












No Prune Low
6 Leaves

Prune Low
6 Leaves

No Prune Low
3 Leaves


Prune Low
3 Leaves


U Actual Treatment Means U] Pruned Loaves Added


65.1


66.5


68.7


68.7


Harvest AJI 1
Leaves 71.0

40 50 60 70
Harvest Index (g cured leaf/100 g stalk + leaves)
Fig. 3-4. The effects of lower leaf harvesting on the harvest index of flue-cured
tobacco and the effects of the pruned leaves on the harvest index of the pruning
treatments. LSD (0.05) = 1.0 for all comparisons. Values outside bars are the
treatment means.


No Prune 6 Low Leaves


Prune 6 Low Leaves


No Prune 3 Low Leaves


Prune 3 Low Leaves


Harvest all Leaves

1000


1802


1696


1738


I1742


1702


1200


1400


1600


1800


Stalk Cured Weight (kg/ha)
Fig. 3-5. The effects of lower leaf harvesting on the stalk weight of flue-cured
tobacco. LSD (0.05) = 88 for all comparisons. Values outside bars are the
treatment means.


- --


i


* Actual Treatment Means


I] Pruned Leaves Added












No Prune Low
6 Leaves


Prune Lo.
6 Leaves

No Prune Low
3 Leaves

Prune Low
3 Leaves

Harvest All
Leaves


U Actual Treatment Means[II Pruned Leaves Added


17.4


.I


16.2


5.8


15.8


Cured Weight Yield (g cured leaf/100 g fresh leaf)
Fig. 3-6. The effects of lower leaf harvesting on the cured weight yield of flue-cured
tobacco and the effects of the pruned leaves on the cured weight yield of the
pruning treatments. LSD (0.05) = 0.5 for all comparisons. Values outside bars
are the treatment means.


No Prune Low
6 Leaves

Prune Low
6 Leaves

No Prune Low
3 Leaves


Prune Low
3 Leaves

Harvest AI
Leaves


E Actual Treatment Means


0I Pruned Leaved Added


lI fllll l hl l llTTIT 9.66


10.77


10.76


9.82


9.48


9.99


9.61

.0 8.0 9.0 10.0 11.0
Specific Cured Leaf Weight (mg cured leaf/sq cm leaf)


Fig. 3-7. The effects of lower leaf harvesting on the specific cured leaf weight of flue-
cured tobacco and the effects of the pruned leaves on the specific cured leaf weight
of the pruning treatments. LSD (0.05) 0.31 for all comparisons. Values
outside bars are the treatment means.


I


I


E Actual Treatment Means


0I Pruned Leaves Added


18.3


I









curing cost to CW was highest for the first harvest. Not harvesting the lower leaves may

result in more efficient handling and curing of the tobacco crop due to the higher CW per

unit of FW harvested and reduced curing cost per unit of CW.

Value per hectare of the tobacco crop declined as the total number of leaves

harvested decreased (Fig. 3-8). As with FW and CW, the values of the pruned LPs were

equal to the values of the those same LPs when normally harvested. Value per kilogram

improved when the lowest three or more leaves were not harvested (Fig. 3-9). An

increase in the average value per kilogram as a result of lower leaf pruning has been

observed in other studies (Suggs, 1972; Currin and Pitner, 1980; Stocks, 1988; Court

and Hendel, 1989; Currin and Stanton, 1989). The increase in average price resulted

because the lowest LPs were of the lowest value per kilogram (Table 3-1).

The potential benefits associated with not harvesting lower leaves include: 1) a

higher average price per unit of the crop, 2) higher returns of CW per unit of FW

handled, 3) a lower curing cost per unit of CW, and 4) a higher value returned to quota.

Although there was a statistically significant increase in value per kilogram in the

present study, there also was a significant reduction in yield and value per hectare when

the lower leaves were not harvested.

The critical value used to determine if a given lower leaf harvesting treatment

significantly influenced value per hectare was $627 ha-1 (Fig. 3-10). The difference

between the control and harvest 18 leaves treatments was $1060 ha-'. The difference

between the critical value and the actual difference was $433 ha-1. For there to be no

statistical difference in the values of the treatments, the harvest 18 leaves treatments

must increase in value by at least $433 ha-1. This difference can be made up by

increased yield of the remaining leaves as found by Currin and Pitner (1980), Stocks

(1988), and Currin and Stanton (1989). However, in the present study and that by

Court and Hendel (1989) yield and value were lowered by reducing the total number of









No Prune Low
6 Leaves

Prune LQw
6 Leaves

No Prune Low
3 Leaves


1


E Actual Treatment Means


m Pruned Leaves Added


I


14410


1443(


5000


7500


15490


10000 12500 15000
Dollars per Hectare ($/ha)


15459
0


Fig. 3-8. The effects of lower leaf harvesting on the value per hectare of flue-cured
tobacco and the effects of the pruned leaves on the value per hectare of the
pruning treatments. LSD (0.05) = 573 for all comparisons. Values outside bars
are the treatment means.


No Prune Low.
6 Leaves


Prune Low
6 Leaves


E Actual Treatment means


[1 Pruned leaves added


aim 3.73


No Prune Low
3 Leaves


3.79

3.79

3.79


3.73


Prune Low
3 Leaves

Harvest All
Leaves


3.78


3.73


3.50 3.55 3.60
Dollars


3.65 3.70
per Kilograms ($/kg)


3.75 3.80


Fig. 3-9. The effects of lower leaf harvesting on the value per kilogram of flue-cured
tobacco and the effects of the pruned leaves on the value per kilogram of the
pruning treatments. LSD (0.05) = 0.04 for all comparisons. Values outside
bars are the treatment means.


15616


Prune Low
3 Leaves

Harvest All
Leaves


I


12710

12740


I


t









Table 3-1. Value per kilogram by leaf position of the lower leaf harvesting treatments.


------------ Lower Leaf Harvesting Treatments ---------------
Leaf harvest all prune low no prune prune low no prune
Position leaves 3 leaves low 3 leaves 6 leaves low 6 leaves
----------------------- $ kg-1 --------------------
19-21 3.77 3.76 3.73 3.75 3.73
16-18 3.79 3.79 3.81 3.74 3.74
13-15 3.82 3.82 3.84 3.84 3.86
10-12 3.71 3.72 3.77 3.74 3.75
7-9 3.67 3.69 3.69 3.70 3.70
4-6 3.57 3.63 3.61 3.49x NHY
1-3 3.29 3.41x NHY 3.40x NHY


these leaves were pruned at topping.
these leaves from these treatments were


not harvested.


No Prune 6 Low Leaves 12710


Prune 6 Low Leaves 12740


No Prune 3 Low Leaves 14410


Prune 3 Low Leaves 14430


Harvest all Leaves 15490

10000 12000 14000 16000
Dollars per Hectare ($/ha)
Fig. 3-10. The effects of lower leaf harvesting on the value per hectare of flue-cured
tobacco. LSD (0.05) 627 for all comparisons. Values outside bars are the
treatment means.


X/indicates
Y/indicates









leaves harvested. Consequently, in these cases the value differences must be

compensated for by either increasing the value per kilogram or planting more area.

The harvest 18 leaves treatments yielded 3815 kg ha-1. Based on this yield and

the needed minimum value increase ($433 ha-1 ) not to be different from the control,

value per kilogram would need to increase by $0.11 kg-1 ($433 ha-1 /3815 kg ha-').

An increased value of $0.11 kg-' is not unrealistic, but the same value per kilogram to

make up the value loss for the harvest 15 leaves option was $0.64 kg-' ($2138 ha-'

/3355 kg ha-'). Pruning or otherwise not harvesting the lowest six leaves was not

feasible based on the results from the present study.

The other method of compensating for value lost by not harvesting lower leaves is

planting more area. The value per hectare when all 21 leaves were harvested was

$15,490 ha-1. The average value per hectare when only 18 or 15 leaves were harvested

was $14,420 ha-1 or $12,730 ha-1, respectively. The additional hectareage required to

compensate for the reduced value when harvesting 18 leaves was 1.07, whereas 1.22

additional ha would be required for the 15-leaf harvest. Pruning or otherwise not

harvesting the lowest three leaves appeared to be the more viable option since value per

kilogram was equal and yield and value per hectare were higher when compared to the

pruning or otherwise not harvesting the lower six leaves.

In the middle 1980's, lower leaf harvesting decisions were based on economics.

The average value per kilogram for the lower leaf (P group) grades of flue-cured

tobacco were well below all others. A comparison of leaf position grade groups for 1986

and 1990 revealed the dramatic changes that have occurred on the U.S. flue-cured

tobacco market floor (Table 3-2). Beginning in 1989, increased demand for lower leaf

U.S. flue-cured tobacco drove prices for that leaf to levels approaching that of middle

leaves. The lower leaf grades (P group) increased 22 to 31% in value depending on

maturity classification, while the middle and upper stalk grades (X, C, and B groups)

increased very little in value over the same period (Table 3-2). The concept of not









Table 3-2. Changes in the Georgia-Florida (Type 14) flue-cured tobacco market
average value per kilogram from 1986 to 1990 of all ripe and mature or unripe
and immature grades common to both years.


-- Ripe & Mature Grades ---

Leaf Position ---- year ----
Grade Group 1986 1990
--- $ kg-1 ---- Change
B Group 3.85 3.97 +3%

C Group 3.62 3.66 +1%

X Group 3.44 3.51 +2%

P Group 2.78 3.51 +22%

Source: 1986 and 1990 Georgia-Florida (Type 14)
the Market News Reporter, Doug Hendrix.


-- Unripe & Immature Grades --

---- year ----
1986 1990
--- $ kg-1 ---- Change
3.64 3.75 +3%

3.53 3.62 +3%

3.18 3.42 +8%

2.51 3.29 +31%

tobacco market grade reports from









harvesting the lowest leaves at the present price structure is not economically sound,

unless potential yield exceeds quota. Still, if the present price structure were similar

to that of 1986, not harvesting the lower leaves would be a realistic option.

Conclusions

Harvesting methods for lower leaves of tobacco did not affect yield parameters of

individual leaves. Pruning and discarding lower leaves was no more or less beneficial to

total yield and value than leaving the lower leaves on the stalk and not harvesting or

discarding them. Total yield and value was reduced as the total number of leaves

harvested was reduced by lower leaf harvesting treatments. Average value per kilogram

was increased by not harvesting three or more lower leaves because these leaves were

the lowest in value per kilogram and CW. At the time of pruning, the lower leaves had

reached their maximum yield and value because returning their yields and values to the

respective pruning treatments resulted in no differences between the total yield or value

of the control treatment, which involved normal harvesting of the same leaves that were

pruned.

If the potential yield of the tobacco crop is such that the yield lost by not

harvesting lower leaves can be tolerated, this practice is advantageous because the

average price per kilogram is enhanced. Total curing cost likely will be lower by not

having to handle the lowest yielding leaves. Yield and value loss must be tolerated, but

not harvesting lower leaves of flue-cured tobacco will likely result in a higher per

kilogram profit margin.













CHAPTER 4
LOWER LEAF HARVESTING: THE INFLUENCE OF TIME ON THE CHEMICAL AND MINERAL
CHARACTERISTICS OF FLUE-CURED TOBACCO LEAF POSITIONS.


Introduction

Lower leaf harvesting options of flue-cured tobacco (Nicotiana tabacum L) have

been recommended based on the total crop response to removal of lower leaves. Studies

by Suggs (1972), Currin and Pitner (1980), and Stocks (1988) found pruning and

discarding the lowest three or four leaves had no impact on the total yield or chemical

balance of tobacco leaves. Court and Hendel (1989) reported yield reductions associated

with reduced leaf number harvested by either lower leaf pruning or topping to a lower

leaf number, but no effects on the leaf chemical components.

In tobacco production, the leaves to be harvested are in a constant state of flux as

they progress from young, very immature leaves to mature leaves which are ready for

harvest. Turgeon (1989) stated that "leaves of dicotylendous plants stop importing and

begin to export photoassimilate when they are 30-60% fully expanded. Developing

leaves continue to import photoassimilate from source leaves for a period after they

have begun to export their own products of photosynthesis." Prior to the topping

process, a definitive source-sink relationship exists on the tobacco plant. There is an

apical meristem (sink) that may have terminated to an inflorescence, newly-unfolded

leaves (sink), leaves in a source-sink transition, and leaves that are mature or

maturing and are exclusively exporters of photosynthate. Hurng et al. (1989)

concluded that removal of the reproductive sink (topping) caused the leaves of tobacco

plants to act as alternate sinks. Crafts-Brandner et al. (1984) found that depodding (or

sink removal) of soybean (Glycine max L Merr.) plants caused leaves and stems to act









as alternate sinks. After topping, based on changes in the area of the leaves at given LPs

over time (Chapter 2), only the leaves at the upper-most LP (19-21) probably would

be in the transition phase of source-sink development as described by Turgeon (1989).

All other leaves should be exclusive exporters of photoassimilate. After topping, the

leaves of a tobacco plant serve two purposes: 1) generation of substrates for dry matter

production via photosynthesis, and 2) storage organs for the products of photosynthesis

(starch) and other processes, i.e. nicotine synthesis of the roots.

Lower leaf harvesting options are essentially manipulations of the source and

sink leaves. If one chooses to not prune or harvest lower leaves, it is possible they may

act as sinks. If one chooses to prune the lower leaves, potential sinks are removed and

the remaining leaf area (sinks) remaining will be smaller. Normal harvesting of flue-

cured tobacco is essentially a leaf pruning procedure in that leaves are removed

methodically throughout the harvest season as they mature on the stalk.

Numerous studies have quantitified the effects of source-sink manipulations of

other plants. Removal of the grain sink of com (Z~a mays L.) caused increases in the

carbohydrate concentration of both upper and lower leaves (Allison and Weinmann,

1970). Soybean pod pruning increased leaf carbohydrate concentration (McAlister and

Krober, 1958; Kollmann et al., 1974; Ciha and Brun, 1978; Mondal et al., 1978;

Streeter and Jeffers, 1979; Crafts-Brandner et al., 1984), and also increased leaf N

and P concentrations (Kollman et al. 1974; Crafts-Brandner et al., 1984). Kollmann et

al. (1974) further reported that leaf Ca and K concentrations were decreased due to

depodding of soybean. Lawn and Brun (1974) found soybean depodding decreased net

photosynthesis, and that the photosynthetic decline was linked to an accumulation of

assimilates in the leaves that would have been translocated to the developing pods.

Depodding of soybean also delayed leaf senescence (Hicks and Pendleton, 1969; Mondal et

al., 1978; Crafts-Brandner et al., 1984). The effects of reproductive sink removal can

be summarized to include increased leaf carbohydrate levels with a subsequent decline in









net photosynthesis, increase in leaf N and P concentrations, decreases in leaf Ca and K

concentrations, and a delay in the onset of senescence.

Removal of more than 10% of the leaf area of wheat (ticum aestivum L.) and

oat (Avna sativa L.) plants reduced grain yield (Womack and Thurman, 1962). Pauli

and Stickler (1961) found that increasing the percentage of leaves pruned from grain

sorghum (Sorhum bicolor L. Moench) plants decreased the total carbohydrate

concentration in the vegetative tissue and grain. Grain N concentration increased, but

vegetative tissue N concentration and the grain yield of sorghum were reduced as

defoliation increased (Stickler and Pauli, 1961). Weber (1955) and McAlister and

Krober (1958) found soybean seed yield and size decreased in response to increasing

defoliation. Grain yield reductions by defoliation were induced by a decline in the total

photosynthetic capacity of the plant caused by a reduction in leaf area.

Flue-cured tobacco is managed differently than most other agronomic crops in

that the leaves, and not the seed, are harvested. Also, leaves are removed as they mature

on the plant. Most source-sink manipulation studies have dealt with either the effects of

removal of the reproductive sink on leaf and seed characters or the influence of

defoliation on seed characters.

The objective of the present study was to evaluate the effects of pruning, not

pruning or harvesting, or harvesting normally, the lower three or six leaves on flue-

cured tobacco plants. The effects were evaluated on leaves harvested as they matured on

the stalk, or monitored over time to maturity. Because previous studies have not been

designed to examine effects relative to leaves at specific stalk positions, the present

study focused on changes by leaf position.

Materials and Methods

The location, cultivars, and management practices for the present study were

identical to those previously discussed in Chapter 2. The lower leaf harvesting

treatments used in the present study were the same as those previously described in









Chapter 3. Methods that differ from those discussed in Chapters 2 or 3 are described

below.

Two separate studies were included to evaluate the effects of lower leaf

harvesting options. A normal harvest study was conducted to evaluate the effects of

lower leaf harvesting options under typical management. A time after pruning and

topping study was designed to monitor temporal changes in leaves at a given LP to

determine when differences appeared.

Normal Harvest Study

Plots consisted of ten plants spaced 41 cm apart, planted in rows spaced 121 cm

apart. Four replications of lower leaf harvesting treatments and cultivars were

evaluated each year in a split-split-plot design. Cultivars were main plots, harvesting

options were sub-plots, and leaf positions were sub-sub-plots. A split-split-split-

plot design was used for statistical analysis with years being main plots, cultivars being

sub-plots, harvesting options being sub-sub-plots, and leaf positions being sub-sub-

sub-plots. Analysis of variance and single degree of freedom contrast procedures were

carried out by methods described by Gomez and Gomez (1984).

Eight cured leaves were subsampled for chemical or mineral analyses from

leaves at LPs 7-9, 13-15, and 19-21. These LPs were selected because they were

common to all treatments and inherently different in many characteristics. The midribs

were removed from the lamina. The lamina was ground using a Wiley mill with a 1 mm

screen. Total N, nicotine, and reducing sugar analyses were performed by R.J. Reynolds

Tobacco Company in Winston-Salem, NC, and Philip Morris Tobacco Company in

Richmond, VA. For Ca, Mg, K, and P analyses, samples were prepared by the ashing and

acid digestion method described by Walsh (1971). The resulting solutions were

analyzed by Inductively Coupled Argon Plasma (ICAP) in the University of Florida's

Institute of Food and Agricultural Science Extension Soil Testing Laboratory by methods

described by Hanlon and Devore (1989).









The leaf chemical and mineral data are reported for leaf lamina only. Midribs

were not evaluated, and previous work indicated that their chemical and mineral

constituents are different from those of the lamina (Darkis et al., 1952).
Time After Pruning Study

Plots consisted of two plants spaced 41 cm apart, planted in rows spaced 121 cm

apart (1 m2 of total area). Three replicates of cultivars and lower leaf harvesting

treatments were evaluated each year and were arranged in the field in a split-split-

split-plot design with cultivars being main plots, harvesting options being sub-plots,

leaf position being sub-sub-plots, and harvest dates being sub-sub-sub-plots. The 21

leaves were partitioned into seven, 3-leaf stalk positions for harvesting purposes.

Weekly harvests were taken of all 3-leaf stalk positions that remained on the stalk,

beginning one week after topping. A set of samples was taken at topping, when the

pruning treatments were imposed, to establish a baseline of the initial levels for the

components at all LPs. Because this study was designed to monitor LP changes over time,

LPs that were harvested in the normal harvest study were removed from all plants that

were to be used for future harvest date analysis. The progression of the LPs as they

were harvested is indicated in Table 4-1.

Because the lower leaf harvesting options resulted in reductions in the number of

leaves harvested either by pruning, or not pruning or harvesting of the lower leaves, all

LPs were not harvested for all treatments. When the lower three leaves were pruned

and discarded at topping or were otherwise not pruned or harvested, only LPs 4-6

through 19-21 were harvested. When the lower six leaves were pruned and discarded at

topping or were otherwise not pruned or harvested, only LPs 7-9 through 19-21 were

harvested. The control treatment (harvest all 21 leaves) resulted in harvesting of all

LPs. For the harvest dates from 21 to 49 days after pruning or topping, all treatments

had an equal number of LPs since LPs 7-9 through 19-21 were common to all

treatments.










Table 4-1. The leaf positions removed per plant by harvest date in the days after
pruning and topping study.


-------------------- days after pruning and topping ------------------
0 7 14 21 28 35 42 49


leaf positions


harvested


19-21 19-21 19-21 19-21 19-21 19-21 19-21 19-21x


16-18 16-18 16-18


10-12 10-12


7-9
4-6
1-3


16-18 16-18 16-18 16-18x


10-12 10-12 10-12X


7-9 7-9 7-9x


4-6
1-3x


4-6x


***Bold type indicates that leaf area, leaf mineral and chemical components, and
total non-structural carbohydrates (TNC) were measured on those leaf positions.
x Indicates leaf positions which were harvested normally on this date.


13-15 13-15 13-15 13-15 13-15 13-15x









Each LP sample consisted of six leaves (2 plants X a 3-leaf position). All leaf

samples were cured in the normal flue-curing manner. After curing, the leaves were

rehydrated for handling purposes. The six leaves at each LP were used for leaf chemical

or mineral analyses. The midribs were removed and the lamina was ground to 1 mm

using a Wiley mill. Total N, nicotine, reducing sugar, Ca, Mg, K, and P analyses were

performed by the same methods described above.

Analysis of the entire data set was not possible due to lack of balance. With each

successive harvest date there was a loss of one LP that was harvested in the normal

harvest study (Table 4-1). As a result, each harvest date was analyzed individually.

Leaf position was an overwhelming factor across harvest dates. For analysis of the

influence of harvesting effects at LP over time, each LP was analyzed for each harvest

date. The statistical design used then was a split-split-plot design with years being

main plots, cultivars being sub-plots, and harvesting options being sub-sub-plots.

Analysis of variance and single degree of freedom contrasts procedures were carried out

by methods described by Gomez and Gomez (1984).

Lower Leaf Harvesting Options Statistical Comparisons

Single-degree of freedom contrasts were chosen to determine differences between

harvesting schemes. With contrasts the possible comparisons are limited to the number

of degrees of freedom for the effect analyzed. There were a total of five harvesting

treatments, so there were four possible contrasts. The comparisons chosen were derived

from a desire to determine combined and individual effects of the harvesting schemes. To

determine the effects of pruning lower leaves versus not pruning and not harvesting

lower leaves, a comparison was made between the combined effects of pruning the lower

three and six leaves and the combined effects of not pruning and not harvesting the lower

three and six leaves. This contrast was designated "prune vs no prune". There were few

effects associated with the lower three leaf harvesting options, but strong effects

associated with the lower six leaf harvesting options. Consequently, a comparison was









made between pruning of the lower six leaves and not pruning or harvesting the lower

six leaves. This contrast was designated "prune 6 vs no prune 6". The harvest all leaves

treatment was considered the control. To evaluate deviations from the norm of the lower

leaf harvesting regimes, comparisons were made between the control versus the

combined pruning treatments, designated "harv all vs prune", and the control versus the

combined not pruning and not harvesting lower leaves treatments, designated "harv all

vs no prune".
Results and Discussion

Leaf Chemical Characters

Nicotine concentration was lower in the leaves that comprised 13-15 due to not

pruning or harvesting the lower six leaves in the normal harvest study (Table 4-2).

Suggs (1972) found higher nicotine concentrations when nine lower leaves were

pruned, but no differences when six or less leaves were pruned. Currin and Pitner

(1980), Stocks (1988), and Court and Hendel (1989) reported no differences in

nicotine concentration due to lower leaf pruning. Over time, nicotine concentrations in

leaves at LPs 7-9, 13-15, and 19-21 were lower as a result of not pruning or

harvesting the lower leaves (Fig. 4-1 & Table 4-3, Fig. 4-2 & Table 4-4, Fig. 4-3 &

Table 4-5). Nicotine concentration for the control was not different from that of the

pruning treatments in the normal harvest study. This result was also observed over

time, however, over the first 14 days after pruning the nicotine concentrations at all

LPs were similar to those at the LPs in the no pruning treatments. Leaves were

discarded from the pruning treatments at topping. Normal harvesting is technically a

pruning process, but since only three leaves were removed per harvest, all lower six

leaves of the control treatment had not been harvested until 14 days after the pruning

treatments were imposed. Nicotine concentrations under normal harvesting were

similar to the no pruning treatments initially, but as more leaves were harvested from

the control treatment the nicotine concentration approached that observed in the pruning










Table 4-2. The effects of lower leaf harvesting options on nicotine concentration of
flue-cured tobacco by leaf position (normal harvest study).


Lower Leaf
Harv. Options


Harv all 21 leaves

Prune low 3 leaves

No prune low 3 leaves

Prune low 6 leaves

No prune low 6 leaves


Prune Treats mean

No Prune Treats mean


-------------------
7 9
7-9
----------- nicotine

12.5

12.7

12.9

12.6

11.4


12.7

12.1


leaf position
13 15
concentration (

15.8

15.6

15.9

15.4

13.2


15.5

14.6


-----------------
19 21
g kg-) -----

22.0

20.8

21.7

20.5

19.6


20.7

20.7


CCNTRASS ----------------- P > F ---------------------

Prune vs No Prune 0.376 0.142 0.989

Prune 6 vs No Prune 6 0.125 0.019 0.473

Harv all vs Prune 0.858 0.641 0.226

Harv all vs No Prune 0.585 0.098 0.222

CV (%) (reps = 4) 17.3 16.9 16.5

































Days After Pruning and Topping

Fig. 4-1. Influence of lower leaf harvesting options on the lamina nicotine
concentration of flue-cured tobacco leaf position 7-9. Appropriate contrasts are
given below.

Table 4-3. Single degree of freedom contrasts for the data presented in Fig. 4-1.

--------------- days after pruning ----------------
7 14 21

ONTRASTS -------------------- P > F ---------------------

Prune vs No Prune 0.084 0.009 0.042

Prune 6 vs No Prune 6 0.187 0.034 0.012

Harv all vs Prune 0.711 0.002 0.244

Harv all vs No Prune 0.077 0.284 0.006

CV (%) (reps = 3) 17.3 13.6 13.2


- Hawrvst A Leaves
---- Pune Low 6 Leaves
-0-- No Prune Low 6 Leave
---A-- Puning Treatments Mean
- -f No Pruning Treatments Mean


.1



























8- ?


7 14 21 28 35
Days After Pruning and Topping

Fig. 4-2. Influence of lower leaf harvesting options on the lamina nicotine
concentration of flue-cured tobacco leaf position 13-15. Appropriate contrasts
are given below.

Table 4-4. Single degree of freedom contrasts for the data presented in Fig. 4-2.

---------------- days after pruning ---------------
7 14 21 28 35

CONTRASTS ------- --------- P > F ---------------------

Prune vs No Prune 0.191 0.001 0.006 0.005 0.219

Prune 6 vs No Prune 6 0.583 0.048 0.018 0.005 0.679

Harv all vs Prune 0.522 0.002 0.118 0.855 0.084

Harv all vs No Prune 0.092 0.926 0.001 0.028 0.008

CV (%) (reps = 3) 21.9 15.1 16.6 21.0 22.2


--"-- Harvest Al Leaves
* Prune Low 6 Leaves
--0" No Prune Low 6 Leaves
-*-*"- Pruning Treatments Mean
- No Pruning Treatments Mean











22.5-


20.0-


17.5-


15.0-


12.5-


10.0-


7.5-


b.U I I I I I I I
0 7 14 21 28 35 42 49
Days After Pruning and Topping
Fig. 4-3. Influence of lower leaf harvesting options on the lamina nicotine
concentration of flue-cured tobacco leaf position 19-21. Appropriate contrasts
are given below.

Table 4-5. Single degree of freedom contrasts for the data presented in Fig. 4-3.

---------------- days after pruning ---------------
7 14 21 28 35 42 49

CONTRASTS ---------.-------- P > F ---------------------

Prune vs No Prune 0.035 0.001 0.007 0.004 0.171 0.019 0.009

Prune 6 vs No Prune 6 0.199 0.031 0.013 0.002 0.496 0.009 0.004

Harv all vs Prune 0.710 0.020 0.461 0.856 0.241 0.345 0.197

Harv all vs No Prune 0.203 0.525 0.004 0.100 0.026 0.006 0.001

CV(%) Reps =3 28.8 18.6 19.8 22.8 24.8 19.4 12.5


-*0- Harvest Al Leaves
-0*- Prune Low 6 Leaves
----- No Prune Low 6 Leaves
-.-A Pruning Treatments Mean
S-f No Pruning Treatment Mean









treatments. Nicotine is synthesized in the roots and translocated to the shoot where it

accumulates predominantly in the leaf (Wolf and Bates, 1964). Based on the nicotine

concentration data over time, it seems likely that the lower leaves that remained on the

plants acted as sinks for nicotine.

Nitrogen concentration was not affected by lower leaf harvesting. This response

was surprising since the concentration of nicotine, a N-containing compound, was

decreased by not removing lower leaves. Leaf N concentration was found to increase in

response to reproductive sink removal in soybean (Glycine max L. Merr.) (Kollman et

al., 1974; Crafts-Brandner et al., 1984). The N concentration of composite leaf and

stem tissue decreased as a result of leaf removal of grain sorghum (Sorghum bicolor L.

Moench) (Pauli and Stickler, 1961). However, delayed topping of flue-cured tobacco

decreased the nicotine concentration in leaves with no affect on the N concentration

(Woltz, 1955; Marshall and Seltmann, 1964; Elliot, 1966).

In the normal harvest study, the N to nicotine concentration ratio (N:Nic) at LP

13-15 was higher as a result of not pruning the lowest six leaves (Table 4-6). The

N:Nic at LPs 13-15 and 19-21 were also higher for the no pruning treatments over

time (Fig. 4-4 & Table 4-7, Fig. 4-5 & Table 4-8). The responses for N:Nic largely

paralleled those observed for nicotine concentration. Since N concentration was not

affected by pruning or not pruning lower leaves, the changes in the N:Nic were due

primarily to changes in the nicotine concentration. Tso (1972) reported that an N:Nic

approximating 1:1 was desirable for the best smoking quality of flue-cured tobacco

leaves. While small differences in N:Nic were found among treatments and the N:Nic was

always higher than 1:1, it is questionable whether the smoking quality would differ

significantly among any of the imposed treatments. Weybrew et al. (1984) found that

smoker preference, being a qualitative variable, and quantitative chemical measures

were not always closely correlated.










Table 4-6. Effects of lower leaf harvesting options on the N to nicotine concentration
ratio of flue-cured tobacco by leaf position (normal harvest study).


Lower Leaf
Harv. Options


Harv all 21 leaves

Prune low 3 leaves

No prune low 3 leaves

Prune low 6 leaves

No prune low 6 leaves


Prune Treats mean

No Prune Treats mean


----------------
7 9
------------ N
7-9
N
1.29

1.27

1.27

1.25

1.34


1.26

1.31


leaf position
13 15
to nicotine concentration
1.14

1.17

1.16

1.16

1.29


1.17

1.23


-----------------
19 21
ratio --------
1.10

1.10

1.09

1.10

1.15


1.10

1.12


CONTRASTS ---------- ---- P > F -------------------

Prune vs No Prune 0.246 0.120 0.555

Prune 6 vs No Prune 6 0.094 0.016 0.332

Harv all vs Prune 0.563 0.515 0.967

Harv all vs No Prune 0.708 0.057 0.659

CV (%) (reps= 4) 11.5 11.7 13.1













3.00-




2.50-



2.00-




1.50-


1.00 1 I II I
7 14 21 28 35
Days After Pruning and Topping
Fig. 4-4. Influence of lower leaf harvesting options on lamina N to nicotine
concentration ratio of flue-cured tobacco leaf position 13-15. Appropriate
contrasts are given below.

Table 4-7. Single degree of freedom contrasts for the data presented in Fig. 4-4.

-------------- days after pruning ----------------
7 14 21 28 35

CONTRASTS --------------- P > F ---------------------

Prune vs No Prune 0.464 0.007 0.045 0.001 0.148

Prune 6 vs No Prune 6 0.802 0.097 0.132 0.006 0.524

Harv all vs Prune 0.983 0.065 0.589 0.547 0.138

Harv all vs No Prune 0.563 0.652 0.256 0.029 0.015

CV (%) (reps = 3) 27.7 16.9 15.4 10.8 9.7


Harvest All Leaves
--- Prune Low 6 Leaves
---0 No Prune Low 6 Leaves
---A- ~Pruning Treatments Mean
- --- No Pruning Treatments Mean













Z 6.0-


.-0- Harvest All Leaves
E 5--0- Prune Low 6 Leaves
S--0- No Prune Low 6 Leave
0 --A.- Pruning TreatmentsMeen
0 No Pruning Treatments Mean
S4.0- \


z
9. 3.0
Z

E 2.0



1.0
0 7 14 21 28 35 42 49
Days After Pruning and Topping

Fig. 4-5. Influence of lower leaf harvesting options on lamina N to nicotine
concentration ratio of flue-cured tobacco leaf position 19-21. Appropriate
contrasts are given below.

Table 4-8. Single degree of freedom contrasts for the data presented in Fig. 4-5.

--------------- days after pruning ---------------
7 14 21 28 35 42 49

CONTRASTS ------------------ P > F ---------------------

Prune vs No Prune 0.049 0.002 0.002 0.003 0.038 0.002 0.017

Prune 6 vs No Prune 6 0.543 0.039 0.030 0.003 0.891 0.001 0.023

Harv all vs Prune 0.045 0.061 0.962 0.463 0.507 0.816 0.913

Harv all vs No Prune 0.013 0.390 0.009 0.065 0.021 0.016 0.038

CV(%) Reps =3 54.5 21.2 17.1 18.8 13.2 9.8 9.6









The reducing sugar concentration of leaves from LP 7-9 were higher when lower

leaves were not pruned or harvested, both over time and in the normal harvest study

(Table 4-9, Fig. 4-6 & Table 4-10). Higher leaf carbohydrate levels resulted when

reproductive sinks were removed from soybean and corn (McAlister and Krober, 1958;

Allison and Weinmann, 1970; Kollmann et al., 1974; Ciha and Brun, 1978; Mondal et

al., 1978; Streeter and Jeffers, 1979; Crafts-Brandner et al.,1984). Pauli and

Stickler (1961) reported lower plant tissue carbohydrate concentration in grain

sorghum resulted from defoliation. Total non-structural carbohydrate (TNC)

concentration at LP 7-9 was not affected by harvesting treatments. Intuitively, a higher

reducing sugar concentration should be the result of a higher TNC concentration. As

discussed in Chapter 2, the reducing sugar concentration changed proportionally with

the TNC concentration at LPs 13-15 and 19-21, however, reducing sugar concentration

did not change proportionally with TNC concentration at LP 7-9.

The reducing sugar to nicotine concentration ratio (Sug:Nic) was higher at LP 7-

9 and LP 13-15 from the normal harvest study in response to not pruning or discarding

lower leaves (Table 4-11). The Sug:Nic of the leaves from LP 7-9 was higher because

of a combination of higher sugar concentration and lower nicotine concentration when the

lower leaves were not pruned or harvested. However, a higher Sug:Nic at LP 13-15

solely resulted from a higher nicotine concentration. Over time, the Sug:Nic of leaves

within LPs 7-9, 13-15, and 19-21 varied due to the effects of either pruning, not

pruning or harvesting, or harvesting the lower leaves in the normal manner (Fig. 4-7

& Table 4-12, Fig. 4-8 & Table 4-13, Fig. 4-9 & Table 4-14).
Mineral Characteristics

Phosphorus was the only mineral whose concentration was responsive to the

harvesting treatments. In the normal harvest study, the P concentrations in leaves from

LPs 7-9 and 13-15 were lower as a result of not pruning or harvesting the lower









Table 4-9. Effects of lower leaf harvesting options on the reducing sugar concentration
of flue-cured tobacco by leaf position (normal harvest study).


Lower Leaf
Harv. Options


Harv all 21 leaves

Prune low 3 leaves

No prune low 3 leaves

Prune low 6 leaves

No prune low 6 leaves


Prune Treats mean

No Prune Treats mean


------------------- leaf
7-9
7 9
----- reducing sugar

234.5

232.3

241.8

231.9

255.0


232.1

248.4


position
13 15
concentration

210.6

217.8

214.0

217.6

224.0


217.7

219.0


-----------------
19 21
(g kg) --------

148.3

143.9

147.7

141.9

151.3


142.9

149.5


CONTRASTS ----------------- P > F --------------------

Prune vs No Prune 0.006 0.791 0.206

Prune 6 vs No Prune 6 0.006 0.361 0.205

Harv all vs Prune 0.728 0.248 0.399

Harv all vs No Prune 0.052 0.172 0.847

CV (%) (reps = 4) 9.5 9.1 14.0











250


240-


230-


220-


210-


200-


190-


Days After Pruning and Topping

Fig. 4-6. Influence of lower leaf harvesting options on reducing sugar concentration of
flue-cured tobacco leaf position 7-9. Appropriate contrasts are given below.

Table 4-10. Single degree of freedom contrasts for the data presented in Fig. 4-6.

--------------- days after pruning ---------------
7 14 21

ONTRASS -------------------- P > F ---------------------

Prune vs No Prune 0.104 0.057 0.025

Prune 6 vs No Prune 6 0.259 0.234 0.032

Harv all vs Prune 0.543 0.274 0.225

Harv all vs No Prune 0.457 0.623 0.004

CV (%) (reps = 3) 19.4 17.9 14.4


-- Harvest Al Leaves
- Prune Low 6 Leaves
0----- No Prune Low 6 Leaves
-.-A*- Pruning Treatments Mean
- No Pruning Treatments Mean


r 1











Table 4-11. Effects of lower leaf harvesting options on the reducing sugar to nicotine
concentration ratio of flue-cured tobacco by leaf position (normal harvest
study).


Lower Leaf
Harv. Options


Harv all 21 leaves

Prune low 3 leaves

No prune low 3 leaves

Prune low 6 leaves

No prune low 6 leaves


Prune Treats mean

No Prune Treats mean


------.--..---.-- lea
7-9
7 9
------ reducing sugar to

19.3

19.0

20.0

19.8

23.3


19.4

21.7


f position
13 15
nicotine conce

14.2

14.8

14.3

15.0

17.9


ntr


-----------------
19 21
ation ratio -------

7.0

7.3

7.2

7.1

8.2


14.9

16.1


CONRASTS ------- -------- P > F ---------------------

Prune vs No Prune 0.060 0.138 0.342

Prune 6 vs No Prune 6 0.041 0.011 0.134

Harv all vs Prune 0.961 0.495 0.812

Harv all vs No Prune 0.111 0.061 0.311

CV (%) (reps = 4) 23.2 20.8 27.0
















n-o






z
S 3

5|


16-1


lb I I I
7 14 21
Days After Pruning and Topping
Fig. 4-7. Influence of lower leaf harvesting options on reducing sugar to nicotine
concentration ratio of flue-cured tobacco leaf position 7-9. Appropriate
contrasts are given below.

Table 4-12. Single degree of freedom contrasts for the data presented in Fig. 4-7.

---------------- days after pruning ---------------
7 14 21

CONTRASS -------------------- P > F ---------------------

Prune vs No Prune 0.101 0.007 0.033

Prune 6 vs No Prune 6 0.208 0.055 0.016

Harv all vs Prune 0.517 0.040 0.081

Harv all vs No Prune 0.474 0.837 0.001

CV (%) (reps = 3) 31.4 26.4 24.5


-D- Harvest All Leaves
---1- Prune Low 6 Leaves
---0-- No Prune Low 6 Leaves
-.- *.- Pruning Treatments Mean
- No Pruning Treatments Mean

































7 14 21 28 35
Days After Pruning and Topping
Fig. 4-8. Influence of lower leaf harvesting options on reducing sugar to nicotine
concentration ratio of flue-cured tobacco leaf position 13-15. Appropriate
contrasts are given below.

Table 4-13. Single degree of freedom contrasts for the data presented in Fig. 4-8.

--------------- days after pruning ----------------
7 14 21 28 35

CONTRASTS -------.--------- P > F ---------------------

Prune vs No Prune 0.187 0.009 0.001 0.007 0.067

Prune 6 vs No Prune 6 0.186 0.131 0.011 0.004 0.572

Harv all vs Prune 0.596 0.070 0.081 0.516 0.711

Harv all vs No Prune 0.575 0.692 0.001 0.103 0.064

CV (%) (reps = 3) 35.7 27.2 22.8 24.8 22.1


-- Harvest All Leaves
-- Prune Low 6 Leaves
----- No Prune Low 6 Leaves
-.-A- Pruning Treatments Mean
-, No Prne Treatments Mean


r... -....

%rr. %- \



A---.... ^^^^^0 1.....













0

C
0 .
0o-

5.2


zE
z


0 7 14 21 28 35 42 49

Days After Pruning and Topping
Fig. 4-9. Influence of lower leaf harvesting options on reducing sugar to nicotine
concentration ratio of flue-cured tobacco leaf position 19-21. Appropriate
contrasts are given below.

Table 4-14. Single degree of freedom contrasts for the data presented in Fig. 4-9.

---------------- days after pruning ---------------
7 14 21 28 35 42 49

CONTRASTS -------- ------- P > F ---------------------

Prune vs No Prune 0.032 0.038 0.010 0.021 0.124 0.004 0.001

Prune 6 vs No Prune 6 0.105 0.277 0.005 0.010 0.212 0.001 0.001

Harv all vs Prune 0.254 0.349 0.444 0.290 0.890 0.628 0.182

Harv all vs No Prune 0.655 0.420 0.005 0.374 0.259 0.005 0.001

CV(%) Reps = 3 54.0 38.3 32.2 33.7 28.7 26.4 18.8


- Harvest All Leaves O
- -" Prune Low 6 Leaves
--- No Prune Low 6 Leaves "
--A".- Pruning Treatments Mean
- No Pruning Treatments Mean









leaves (Table 4-15). Differences in the P concentrations at LPs 7-9 and 13-15 due to

not removing lower leaves were also found over time (Fig. 4-10 & Table 4-16, Fig. 4-

11 & Table 4-17). Soybean pod pruning increased P concentration in the leaves

(Kollman et al., 1974; Crafts-Brandner et al., 1984). Whereas soybean depodding

involves removal of a strong sink for P, the P concentrations of leaves likely differed in

the present study because the lower leaves that were not pruned or harvested acted as a

sink for P.
Conclusions

The quality of the cured leaf is affected by the ratios of the chemical components

of the leaf. If the chemical characteristics of the control (harvest all leaves) treatment

were used as standards, some generalizations can be made about the effects of the lower

leaf harvesting. The leaf chemical balance of the lower leaf pruning treatments more

closely approximated that of the control. When lower leaves were left on the plant,

nicotine dilution in the remaining leaves was the most obvious response. The lower

nicotine concentration resulted in higher than normal N to nicotine and reducing sugar to

nicotine concentration ratios. Phosphorous concentration of leaves from LPs 7-9 and

13-15 were lowered in response to not pruning or harvesting the lower leaves. The

concentration of other minerals were not affected by lower leaf harvesting.

Interesting responses occurred as a result of pruning, not pruning, or normal

harvesting, of lower leaves over time. Over the first 14 days after pruning, the

chemical and mineral characteristics were similar for the leaves above those which

were either not pruned or were harvested normally. However, as normal harvesting of

lower leaves progressed, the chemical and mineral characteristics of leaves above those

normally harvested approached those observed for the same leaves when lower leaves

were pruned and discarded.










Table 4-15. Effects of lower leaf harvesting options on the P concentration of flue-
cured tobacco by leaf position (normal harvest study).


Lower Leaf
Harv. Options


Harv all 21 leaves

Prune low 3 leaves

No prune low 3 leaves

Prune low 6 leaves

No prune low 6 leaves


Prune Treats mean

No Prune Treats mean


-------------------
7-9
7 9
-------------- P

3.23

3.31

3.28

3.43

3.05


3.37

3.14


leaf position
13 15
concentration (g
3.05

3.12

2.96

3.03

2.81


3.08

2.87


-----------------
19 21
kg-1) --------
3.32

3.17

3.18

3.10

3.42


3.14

3.33


CONTRASTS ----------- ----- P > F --------------------

Prune vs No Prune 0.009 0.011 0.203

Prune 6 vs No Prune 6 0.001 0.032 0.078

Harv all vs Prune 0.133 0.752 0.220

Harv all vs No Prune 0.482 0.070 0.852

CV (%) (reps = 4) 9.1 9.7 15.5


m






















3.0 -


2.8 -4


2.6 -- I I
7 14 21
Days After Pruning and Topping
Fig. 4-10. Influence of lower leaf harvesting options on the P concentration of flue-
cured tobacco leaf position 7-9. Appropriate contrasts are given below.


Table 4-16. Single degree of freedom contrasts for the data presented in Fig. 4-10.

---------------- days after pruning ----------------
7 14 21

CONTRASTS -------------------- P > F ---------------------

Prune vs No Prune 0.601 0.079 0.242

Prune 6 vs No Prune 6 0.293 0.289 0.589

Harv all vs Prune 0.999 0.552 0.162

Harv all vs No Prune 0.669 0.045 0.022

CV (%) (reps = 3) 13.4 18.7 12.4


-
.. --
, ..o. -* "--



--- Harvest All Leaves
-- Prune Low 6 Leaves
----- No Prune Low 6 Leaves
-.-A.- Pruning Treatments Mean
- -Af No Pruning Treatments Mean































2.4 1 I I I
7 14 21 28 35

Days After Pruning and Topping

Fig. 4-11. Influence of lower leaf harvesting options on the P concentration of flue-
cured tobacco leaf position 13-15. Appropriate contrasts are given below.


Table 4-17. Single degree of contrasts for the data presented in Fig. 4-11.

---------------- days after pruning ---------------
7 14 21 28 35

NTRASS -------------------- P > F ---------------------

Prune vs No Prune 0.047 0.045 0.092 0.005 0.278

Prune 6 vs No Prune 6 0.009 0.284 0.104 0.024 0.743

Harv all vs Prune 0.999 0.130 0.357 0.890 0.403

Harv all vs No Prune 0.101 0.883 0.025 0.014 0.090

CV (%) (reps 3) 10.7 11.9 13.5 12.7 14.7


-- Harvest A Leaves
-**)- Prune Low 6 Leaves
---O-- No Prune Low 6 Leaves
---A- Pruning Treatments Mean
-' No Pruning Treatments Mean