Citation
Effect of the vegetable leafminer, Liriomyza sativae Blanchard, and the associated plant pathogens on yield and quality of the tomato, Lycopersicon esculentum Mill. cv. Walter

Material Information

Title:
Effect of the vegetable leafminer, Liriomyza sativae Blanchard, and the associated plant pathogens on yield and quality of the tomato, Lycopersicon esculentum Mill. cv. Walter
Alternate Title:
Liriomyza sativae
Alternate Title:
Lycopersicon esculentum
Creator:
Keularts, Jozef Leo Willem, 1945- ( Dissertant )
Waddill, Van H. ( Thesis advisor )
Strayer, John R. ( Reviewer )
McMillan, Robert T. ( Reviewer )
Pohronezny, Kenneth L. ( Reviewer )
Place of Publication:
Gainesville, Fla.
Publisher:
University of Florida
Publication Date:
Copyright Date:
1980
Language:
English
Physical Description:
x, 154 leaves : graphs ; 28 cm.

Subjects

Subjects / Keywords:
Defoliation ( jstor )
Fruits ( jstor )
Infestation ( jstor )
Leafminers ( jstor )
Leaves ( jstor )
Parasites ( jstor )
Planting ( jstor )
Species ( jstor )
Tomatoes ( jstor )
Vegetables ( jstor )
Dissertations, Academic -- Entomology and Nematology -- UF
Entomology and Nematology thesis Ph. D
Leafminers -- Control ( lcsh )
Tomatoes -- Diseases and pests -- Control ( lcsh )
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Abstract:
In the period from 1977 to 1980 a number of field experiments were carried out at the University of Florida Agricultural Research and Education Center in Homestead to determine the effect of various levels of discrete or repeated, mechanical defoliation of 'Walter 1 tomato plants on components of marketable yield. Treatments consisted of 100% defoliation and separate 20%, 40%, 60%, and 80% defoliations of the lower or the upper part of the plants. A differential sensitivity to defoliation in the course of the plants' development was observed. The most sensitive times appeared to be early in the season and at mid-season. In most cases, however, at least 60% of the foliage had to be removed before total marketable fruit yields and yields in the largest, most profitable size categories were significantly reduced when compared to yields from the control plants. Tomato plants exhibited less tolerance of repeated defoliation with removal of 40% of the total leaf area often resulting in yield loss in the first harvest. However, the total yield of the first two harvests combined was not significantly reduced when compared to yields from nondefoliated plants. The total marketable yield of the tomato plants at any level of defoliation was significantly correlated with the gross revenue a grower would obtain from the harvested fruit based on different prices for the various size categories. Major defoliation associated with leafminer damage in commercial production plantings is the result of the adverse effect of pathogens inhabiting the leaf mines. In this study the pathogen most commonly associated with the leaf mines has been identified as Alternaria alternata (Fries) Keissler. It appears to be only weakly parasitic, its detrimental effect depending on the nutrient supply provided in the mine by mesophyll cells lacerated by the leafminer larvae. Additional damage to the leaf can also be done by other pathogens such as J<anthomonas vesicatoria (Doidge) Dows., which may enter mines when bacterial spot disease pressure is high. The actual damage to the tomato leaf by the leafminer larvae themselves seems to be restricted to the removal of photosynthetically active tissue. The main concern for growers, therefore, should lie in the occurrence of infection of the mines. Infection is probably less likely to occur when the nutrient supply available for the pathogen is too little for it to do harm to the leaf tissue. This is the case when the larvae are killed early in their development so that only a small amount, of leaf tissue has been consumed. The most effective way for ensuring their early death and, consequently, low leafminer populations is effective use of the numerous parasites of the fly. This can best be achieved by applying sound pest management practices. If there are too few parasites to control the leafminer population effectively then insecticide applications specifically to control the leafminer are necessary. If the defoliation level in prebloom plants reaches 30% or in postbloom plants reaches 50%, then insecticide treatments are recommended.
Thesis:
Thesis--University of Florida.
Bibliography:
Bibliography: leaves 143-153.
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Jozef Leo Willem Keularts.

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
Resource Identifier:
000014425 ( alephbibnum )
06800011 ( oclc )
AAB7649 ( notis )

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EFFECT OF THE VEGETABLE LEAFMINER,
Liriomyza sativae BLANCHARD, AND THE ASSOCIATED PLANT
PATHOGENS ON YIELD AND QUALITY OF THE TOMATO,
Lycopersicon esculentum MILL. CV. WALTER








By


JOZEF LEO WILLEM KEULARTS


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


UNIVERSITY OF FLORIDA































Voor mijn ouders, Leo en Maria Keularts,
in dankbaarheid en toewijding













ACKNOWLEDGEMENTS


I would like to thank many people for their cooperation and

support:

Dr. Van H. Waddill, my advisor and chairman of my committee, for

his constructive suggestions, his constant availability to help, and

his confidence in me and in this project.

Dr. Kenneth L. Pohronezny for his practical help, sharing ideas,

serving on my committee, and positive criticisms.

Drs. Robert T. McMillan, Jr., and John R. Strayer for their in-

terest and cooperation in serving on my committee.

Dr. Thomas L. Davenport for sharing his laboratory facilities and

his expertise in photosynthesis studies and gas chromatography.

Dr. Carol A. Musgrave for sharing her research experience and

exchanging ideas.

Dr. Stratton H. Kerr for his continuous sound advice and encourage-

ment both before my entrance to and throughout my progress in this

course of study.

The entire staff of the University of Florida Agricultural Research

and Education Center in Homestead for their cooperation, particularly

Whitaya Chaisit, Rodney Chambers, Wilbur Dankers, Phyllis Daum, Joyce

Francis, Charla Phillips, and Steven Williams, whose voluntary help

and enthusiastic support in the arduous task of grading tomatoes will

never be forgotten.







Jorge Pena, for his encouraging sense of humor and friendship as

well as his help in data collection.

The Florida Tomato Exchange and the Center for Environmental

Programs and Natural Resources whose financial support made this pro-

ject possible.

Mia, my 22-month-old daughter, whose birth and life brought sorely

needed moments of joy and laughter to the tedium of study.

My wife, Mary Jane Provost, who encouraged me to take up this

study, for her steadfast support and patience, but especially for being

my wife.














TABLE OF CONTENTS


ACKNOWLEDGEMENTS................................................ iii

ABSTRACT........................................ ............... viii

CHAPTER

I INTRODUCTION.......... ............................... 1

II A REVIEW OF THE LITERATURE ON THE IDENTIFICATION AND
CONTROL OF THE VEGETABLE LEAFMINER, Liriomyza sativae
BLANCHARD............................................ 5

Introduction.................................... 5

Identification of the Vegetable Leafminer....... 6

Control of the Vegetable Leafminer .............. 8

Control by Parasites...................... 8

Cultural Control.......................... 16

Control by Host Plant Resistance.......... 16

Chemical Control .......................... 17

Conclusions................................... 19

III MECHANICAL DEFOLIATION OF THE TOMATO, Lycopersicon
esculentum MILL. CV. WALTER, AND ITS EFFECT ON YIELD
AND FRUIT QUALITY. ................................... 20

Introduction................................... 20

Materials and Methods........................... 23

General................................... 23

One-Time Defoliation ...................... 24

Experiment 1........................ 26

Experiment 2........................ 27

Experiment 3........................ 27









Repeated Defoliation .................... 27

Experiment 4........................ 28

Experiment 5........................ 28

Gross Revenue Computation................ 28

Results ........................................ 28

Experiment 1.............................. 28

Experiment 2.............................. 32

Experiment 3.............................. 33

Experiment 4.............................. 35

Experiment 5.............................. 36

Discussion.................................... 36

Conclusion..................................... 113

IV MICRO-ORGANISMS ASSOCIATED WITH MINES OF
Liriomyza sativae BLANCHARD ........................... 115

Introduction.. ................................. 115

Materials and Methods........................... 116

Isolation from Leaves..................... 116

Pathogenicity Tests....................... 118

Fungal tests........................ 118

Bacterial tests..................... 119

Isolation from Flies...................... 119

Results........................................ 121

Symptoms ................................. 121

Isolations from Leaves and Flies.......... 121

Discussion ..................................... 127

Conclusions.. .................................. 131











V EFFECT OF THE VEGETABLE LEAFMINER, Liriomyza sativae
BLANCHARD, ON THE PHOTOSYNTHETIC ACTIVITY OF
INDIVIDUAL TOMATO LEAFLETS ............... ...... ...... 133

Introduction.................................... 133

Materials and Methods........................... 133

Results........................................ 134

Discussion.................................... 135

Conclusions..................................... 137

VI CONCLUSIONS........................................... 140

REFERENCES CITED............................................... 143

BIOGRAPHICAL SKETCH.............................................. 154













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


EFFECT OF THE VEGETABLE LEAFMINER,
Liriomyza sativae BLANCHARD, AND THE ASSOCIATED PLANT
PATHOGENS ON YIELD AND QUALITY OF THE TOMATO,
Lycopersicon esculentum MILL. CV. WALTER

By

JOZEF LEO WILLEM KEULARTS

August 1980


Chairman: Dr. Van H. Waddill
Major Department: Entomology and Nematology

In the period from 1977 to 1980 a number of field experiments

were carried out at the University of Florida Agricultural Research

and Education Center in Homestead to determine the effect of various

levels of discrete or repeated, mechanical defoliation of 'Walter'

tomato plants on components of marketable yield. Treatments

consisted of 100% defoliation and separate 20%, 40%, 60%, and 80%

defoliations of the lower or the upper part of the plants.

A differential sensitivity to defoliation in the course of the

plants' development was observed. The most sensitive times appeared to

be early in the season and at mid-season. In most cases, however, at

least 60% of the foliage had to be removed before total marketable

fruit yields and yields in the largest, most profitable size categories

were significantly reduced when compared to yields from the control

plants. Tomato plants exhibited less tolerance of repeated defoliation









with removal of 40% of the total leaf area often resulting in yield

loss in the first harvest. However, the total yield of the first two

harvests combined was not significantly reduced when compared to yields

from nondefoliated plants.

The total marketable yield of the tomato plants at any level of

defoliation was significantly correlated with the gross revenue a

grower would obtain from the harvested fruit based on different prices

for the various size categories.

Major defoliation associated with leafminer damage in commercial

production plantings is the result of the adverse effect of pathogens

inhabiting the leaf mines. In this study the pathogen most commonly

associated with the leaf mines has been identified as Alternaria

alternate (Fries) Keissler. It appears to be only weakly parasitic,

its detrimental effect depending on the nutrient supply provided in the

mine by mesophyll cells lacerated by the leafminer larvae. Additional

damage to the leaf can also be done by other pathogens such as

Xanthomonas vesicatoria (Doidge) Dows., which may enter mines when

bacterial spot disease pressure is high.

The actual damage to the tomato leaf by the leafminer larvae

themselves seems to be restricted to the removal of photosynthetically

active tissue. The main concern for growers, therefore, should lie in

the occurrence of infection of the mines. Infection is probably less

likely to occur when the nutrient supply available for the pathogen is

-oo little for it to do harm to the leaf tissue. This is the case when

the larvae are killed early in their development so that only a small

amount of leaf tissue has been consumed. The most effective way for

ensuring their early death and, consequently, low leafiiner populations








is effective use of the numerous parasites of the fly. This can best be

achieved by applying sound pest management practices. If there are too

few parasites to control the leafminer population effectively then

insecticide applications specifically to control the leafminer are

necessary. If the defoliation level in prebloom plants reaches 30% or

in postbloom plants reaches 50%, then insecticide treatments are

recommended.













CHAPTER I

INTRODUCTION



The tomato, Lycopersicon esculentum Mill., is one of the most

important commercially-grown vegetable crops in the United States.

With a market value of $517,769,000 in 1979, 19.61% of the total value

of the principal vegetable crops, it was second only to lettuce. In

acreage it was the fourth most important crop with a harvested area of

51,924 hectares, equalling 7.92% of the total commercial vegetable

acreage in the United States (Anonymous, 1980a). Florida is the second

largest producer of tomatoes in the United States and the country's

sole domestic supplier during the months of January, February, and March

(Anonymous, 1979; Zepp and Simmons, 1979). The Dade County tomato

production area accounts for approximately 25% of the state's total

number of harvested acres (Anonymous, 1980b).

For cultivation of tomatoes on the Rockdale soils of South Florida,

the investment in land preparation, planting, and cultivation prac-

tices, including pesticide applications, was estimated at $526.75 per

cultivated acre for a 842-acre farm in 1973 (Walker and Hunt, 1973).

Additional costs for harvesting, processing, and overhead were estimated

at $835.08 per acre. With an investment of this magnitude, it is not

surprising that tomato growers, especially in Dade County, Florida,

would strive to hold yield losses due to pests to a minimum.







Since the end of World War II when chlorinated hydrocarbons came

into widespread use as pesticides, tomato yield has increased from 9.3

metric tons average per hactare for the period 1945-1950 to 17.9 metric

tons average per hectare for the period 1965-1970 (Rose, 1973). Al-

though this near doubling of the yield can be attributed to a number

of cultural practices, effective control of the large number of insect

pests has, undoubtedly, been an important contributing factor.

Leafminers have for the last few decades been considered one of

the most serious pests infesting tomatoes (Wene, 1955; Hayslip, 1961;

Poe et al., 1978). Serious outbreaks of the vegetable leafminer on

tomato occurred for the first time in 1946 and resulted in serious

defoliation of the crop (Anonymous, 1947). An increase in leafminer

populations has been attributed to the general use of DDT and other

chlorinated hydrocarbons as well as organophosphates (Spencer, 1973a).

An accentuation of the leafminer problem has been shown to be the result

of a reduction of the parasite populations by the use of insecticides

(Wene, 1955) as well as a result of the ineffectiveness of the in-

secticides against the leafminers themselves (Hills and Taylor, 1951;

Shorey and Hall, 1963).

The actual damage done by the vegetable leafminer, Liriomyza

sativae Blanchard, to the host plant consists of the consumption of

the leaf's pallisade tissue, which presumably results in a reduction

of the photosynthetic activity of the plant. If the mine density is

high, this damage alone can trigger leaf abscission, ultimately re-

sulting in serious defoliation of the plant. In some cases defoliation

has been reported to have occurred so late in the season that only

little damage was done and an actual benefit in advanced maturity has







been suggested (Michelbacher et al., 1949). However, in fields where

defoliation occurred early in the season serious fruit loss due to

sunburn was observed. Leafminer infestations have been suggested as

the cause of yield reduction in tomatoes on occasion (Wolfenbarger

and Wolfenbarger, 1966; Schuster et al., 1976), but others report no

significant yield decrease despite defoliation by leafminers or in-

creased leafminer density (Levins et al., 1975; Schuster et al., 1976;

Schuster and Everett, 1977; Johnson et al., 1980a,b). The possibility

of negative effects on tomato yield and quality by leafminer popula-

tions larger than those observed was not ruled out, however (Levins

et al., 1975).

Even in the case of low mine densities, defoliation may occur.

The leafminer flies can be directly or indirectly responsible for any

secondary damage occurring as a result of punctures in the leaf

epidermis. The exposure of damaged internal leaf tissue to the

atmosphere provides opportunities for micro-organisms, including plant

pathogens, present on the leaf surface or on the fly's ovipositor to

colonize the leaf (Portier, 1930; Baranowski, 1958; Spencer, 1973a).

Numerous species of micro-organisms have been found in the phylloplane

of healthy plants including tomatoes (Sinha, 1971; Dickinson, 1976)

many of which are able to live as parasites (Dickinson, 1976). Secondary

damage to mined leaves has been noted on sugarbeet (Landis et al., 1967),

alfalfa (Andaloro and Peters, 1977), and grasses (Kamm, 1977).

Discoloration of leaf areas not directly damaged by leafminers

has been observed on various plants and was attributed to either effects

of the iainer itself, its metabolites, or the host tissue (Hering,

1951). Yellowing and necrosis of the mines' vicinity occurs frequently








(Hering, 1951; Kamm, 1977). Leaf injury due to secondary fungal in-

fection has been suggested (Spencer, 1973a) but has been rarely shown

(Haddow, 1941). Bacterial diseases associated with leafminer damage

have also been shown (Sohi and Sandhu, 1968; Leach, 1927).

Yellowing and necrosis of leaves of vegetables, especially tomato,

have been noted frequently. It has been suggested that a pathogen,

presumably an Alternaria species, is responsible for this damage

(Baranowski, 1958). Research was undertaken during the period 1977-1980

to investigate the effects of the vegetable leafminer and associated

pathogen(s) on the yield and fruit quality of the tomato. Specific

objectives of this study were to determine:

(1) the effect of mechanical defoliation of tomato plants in the

field, carried out at different levels and times, on the

production of fruits,

(2) the identity of the pathogen(s) and saprophytes, if any,

present in the discolored leaf areas adjacent to leaf mines

and associated with non-diseased mines, and

(3) the effect of the presence of leaf mines on the photosynthetic

activity of otherwise healthy tomato leaflets.













CHAPTER II
A REVIEW OF THE LITERATURE ON THE IDENTIFICATION AND CONTROL OF
THE VEGETABLE LEAFMINER, Liriomyza sativae Blanchard



Introduction


In order to protect a crop efficiently from insects it is best to

apply a sound pest management strategy instead of relying for the

greater part on insecticides. The successful application of an inte-

grated pest management program for tomatoes has been recently demon-

strated (Pohronezny et al., 1978b). The use of insecticides has

repeatedly led to a decline in the acreage used for the production of

some crops (Metcalf, 1975). A result of the widespread use of insecti-

cides is the selection of secondary pests, formerly controlled by their

natural enemies (Metcalf, 1975). The vegetable leafminer, Liriomyza

sativae Blanchard, is such a pest (Pohronezny and Waddill, 1978) and

it would, therefore, be advisable to control the primary tomato pests

by using methods which allow natural enemies to help reduce the leafminer

populations below economic thresholds.

The present review summarizes the difficulties encountered in

identifying the leafminer and the various methods tried or available

for the control of leafminers infesting tomatoes.








Identification of the Vegetable Leafminer


The vegetable leafminer has been given many names. Frost (1924)

was the first to record a dipterous leafminer on tomato although the

insect was not reared, and, therefore, not identified. Wolfenbarger

(1947) reared Liriomyza pusilla (Meigen) from serpentine mines on tomato

and a number of other crops. L. pusilla was originally described as

Agromyza pusilla by Meigen in 1830 (Frick, 1956). Various workers sub-

sequently reported the serpentine leafminer as occurring on a large

number of host plants (Webster and Parks, 1913; Frost, 1924). Frost

(1924) describes mines of A. pusilla Meigen as a serpentine type on

some hosts and as a blotch type on others. Later Frost (1943) states

that larvae of this fly do not produce serpentine mines and concludes

that Webster and Parks were working with several species. Spencer

(1973a) stated that one of the species involved was undoubtedly

Liriomyza sativae Blanchard.

Each leafminer species appears to produce a consistent pattern in

the construction of its mine (Spencer and Stegmaier, 1973); so it can

be assumed that any report of a leafminer producing mines other than

serpentine ones does not refer to the vegetable or serpentine leafminer,

L. sativae Blanchard. Furthermore, A. pusilla Meigen, synonymous with

L. pusilla (Meigen), is believed to be a European species not occurring
in North America (Stegmaier, 1972) and consequently any reference to

this species name or its synonyms as listed by Frick (1956) in the

United States probably constitutes a misidentification.

Lange (1949) recognized three distinct species, all with a yellow

scutellum, infesting tomatoes in California: Agromyza (Liriomyza)







pusilla Meigen, Agromyza (Liriomyza) subpusilla Frost, and a species

close to Agromyza (Liriomyza) flaveola Fallen. Frick (1957) describes

Liriomyza munda n.sp., L. propepusilla Frost, and L. pictella (Thomson).

Various leafminer species reported earlier by Lange and others are

classified as belonging to one of these three described species.

Agromyza (Liriomyza) pusilla Meigen was considered a synonym for L.

munda Frick; Aqromyza (Liriomyza) subpusilla Frost could be synonymous

with L. munda Frick, L. propepusilla Frost or L. pictella (Thomson);

L. subpusilla (Frost) could by synonymous with L. munda Frick or L.

pictella (Thomson). Most of the records of L. pusilla (Meigen) are

thought to be on L. munda Frick (Stegmaier, 1972) while all references

to L. pictella also include L. munda Frick (Stegmaier, 1966). Spencer

(1965) suggested that the name L. pictella (Thomson) should temporarily

be restricted to the holotype since all species previously identified

as L. pictella proved to be L. munda Frick. Liriomyza munda Frick as

well as L. canomarginis Frick, L. guytona Freeman, L. minutiseta Frick,

and L. pullata Frick are considered to be synonyms of L. sativae Blan-

chard by Spencer (1973a). Liriomyza propepusilla Frost is also thought

to be synonymous to L. sativae Blanchard (Musgrave et al., 1975).

Spencer (1973a,b) lists ten Agromyzid species occurring on

Solanaceous plants. Of these one is a stem miner of the tomato, one

a tuber miner of the white potato, and the remainder leafminers on a

number of plant species. Of the seven species specifically listed as

occurring on tomatoes, only three are reported from the United States:

Liriomyza huidobrensis (Blanchard), L. trifolii (Surgess), and L.

sativae Blanchard. In South Florida only the latter two have been

observed. Liriomvza trifolii (Burgess) may be confused with L. sativae

in infestations 3f crooD (Spencer, 1973a).







Many Liriomyza species are morphologically so similar that often

only examination of the male genitalia will enable one to make a satis-

factory identification (Spencer, 1973a). It is not surprising, there-

fore, that so many names have been given to the tomato serpentine

leafminer when it was discovered with morphological variations dif-

ferent enough from the holotype to make it appear to be a different

species.



Control of the Vegetable Leafminer

Control by Parasites


Parasites have been considered capable of keeping leafminer popula-

tions below economic levels (Baranowski, 1958; Michelbacher et al.,

1951, 1952; Wene, 1955; Getzin, 1960). Musgrave et al. (1975) suggested

a pest management strategy in which the leafminer populations are

allowed to be controlled by their parasites as much as possible. At

least 47 species of Hymenopterous parasites have been reared from L.

sativae Blanchard in various locations in the Western hemisphere. A

list of these parasites with the reported name of the host from which

each one had been reared is given in Table 1.

Of these at least 14 species have been reared from larvae and

pupae of L. sativae in Florida (see Table 1). Also reared were un-

identified species in the following genera: Opius, Chrysocharis,

Achrysocharis, Derostenus, and Diglyphus (Musgrave et al., 1975;

Stegmaier, 1966). In addition one species, Diglyphus pulchripes

(Crawford) is recorded as occurring in Florida (Stegmaier, 1972) and

has been reared from L. sativae elsewhere (Oatman, 1959; McClanahan,






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1977). A Geocoris species has been observed by the author to attack

leafminer pupae on plastic mulch underneath the tomato canopy.



Cultural Control


Abandoned tomato fields could be an important factor in contributing

to the leafminer problem (Adlerz, 1961). It has been recommended that

plants and plant debris remaining in the field after the final harvest

be destroyed to eliminate this fly source (Brogdon, 1961; Wolfenbarger,

1961). However, others feel that abandoned fields contribute few

vegetable leafminers to the agroecosystem (K. Pohronezny and V.H.

Waddill, personal communication, 1980).

Use of various types of mulching has on several occasions been

shown to decrease the number of leaf mines of L. sativae in tomato and

squash (Wolfenbarger and Moore, 1968; Chalfant et al., 1977). An in-

crease, using plastic coated paper as the mulch, was found also (Price

and Poe, 1976).

Staking has been shown to increase the leaf mine density and de-

crease parasitism of the leafminer by Opius dimidiatus (Price and Poe,

1976).



Control by Host Plant Resistance


Differential response of Liriomyza sativae Blanchard in cantaloups,

chrysanthemums, muskmelons, and tomatoes has been found (Kelsheimer,

1953; Wolfenbarger, 1966; Webb and Smith, 1969; Webb et al., 1971;

Kennedy at al., 1975, 1978; Schuster and Harbaugh, 1979). These dif-

ferences in resistance in the tomato cultivars were not great although








several accessions of species related to the cultivated tomato were

virtually immune or demonstrated a considerable antibiosis (Webb and

Smith, 1969; Webb et al., 1971).

A relatively low level of leafminer resistance in tomato may

sufficiently reduce leafminer damage to provide adequate control (Webb

et al., 1971).



Chemical Control


Since the vegetable leafminer became a major pest on various truck

crops in the mid-1940's, many insecticides have been applied to reduce

its populations. DDT has been shown ineffective against leafminers

and, because of the reduction of their parasite populations, actually

caused an increase in the fly population (Hills and Taylor, 1951; Shorey

and Hall, 1963). Research on insecticidal control of the leafminer has

been intensive (Musgrave et al., 1975). A large number of compounds

has been shown to be promising, but the effectiveness of several has

decreased over years of use. Chlordane was one of the first insecti-

cides recommended for leafminer control (Wolfenbarger, 1947). Toxa-

phene, parathion, aldrin, and dieldrin were also effective in the 1940's

(Wolfenbarger, 1948, 1958; Michelbacher et al., 1951; Wene, 1953). All

these products, however, seemed to be losing their effectiveness

(Wolfenbarger, 1958; Baranowski, 1958; Wene, 1955). Toxaphene, para-

thion, and aldrin were shown to reduce the parasite population without

directly affecting the leafminers (Wene, 1955). In the mid-1950's

diazinon became the most effective insecticide (Baranowski, 1958;

Wolfenbarger, 1958), although it was very toxic to some parasites







(Getzin, 1960). Brogdon (1961) and Adlerz (1961) found that diazinon

was no longer effective in 1961 in south Florida but that it still

controlled leafminer populations in north and central Florida. Good

reduction of the fly population on tomato with diazinon was also ob-

tained in California (Shorey and Hall, 1963). Dimethoate was shown

to be very effective in controlling leafminers in the late 1950's

(Getzin, 1960; Hayslip, 1961; Wolfenbarger, 1961). Getzin (1960) found

that dioxathion gave excellent control; on the other hand, Harris

(1962) found it to be ineffective. Harding (1971) noted good control

of the vegetable leafminer on tomato with methamidophos, monocrotophos,

and dimethoate in Texas. These three compounds were also effective on

tomato in south Florida in 1976 (Schuster et al., 1976). However,

dimethoate is not considered effective against the leafminer anymore

(Pohronezny and Waddill, 1978). Oxamyl gave good control of leafminer

on tomato in south Florida in 1974 (Bear, 1975) and in 1975 (Schuster

et al., 1975). In 1977, however, control of defoliation of tomatoes

by applications of oxamyl was not satisfactory (Schuster and Everett,

1977). Janes and Genung (1977) also found no control of the vegetable

leafminer on celery with this insecticide. The use of methomyl for

the control of Lepidopterous pests of tomato destroys the parasite

population and subsequently increases the leafminer densities (Oatman

and Kennedy, 1976; Janes and Genung, 1977; Johnson et al., 1980a,b).

The negative effects of juvenile hormone analogs on biological

control agents appear to outweigh the beneficial effects on target

pests (Poe, 1974; Lema and Poe, 1978). Synthetic pyrethroids like

permethrin seem to give excellent control of leafminer populations

(Schuster et al., 1975; Janes and Genung, 1977; Tryon, 1979) and are







relatively non-toxic to some of the parasites (Waddill, 1978). This

group of compounds may be a good alternative to the conventional in-

secticides because of the spectrum of activity and low toxicity to

parasites and mammals (Schuster et al., 1975).



Conclusions

It would be reasonable to suggest that the vegetable leafminer,

Liriomyza sativae Blanchard, has become resistant to many insecticides

since many of these chemicals have been ineffective in reducing the

leafminer populations. Before the development of DDT and other

chlorinated hydrocarbons, the leafminer had never been considered a

problem in tomato production. Apparently its population was kept suf-

ficiently low by natural enemies. The list of Hymenopterous parasites

recorded for L. sativae illustrates the large number of natural enemies

of this pest species. Since neither host plant resistance nor cultural

methods have as yet been capable of reducing leafminer populations

effectively, the solution to the problem seems to lie in integration

of chemical and biological control.

Allowing the parasite population to build up rapidly as early in

the season as possible would be of great value. This may be accom-

plished by maintaining the parasite population on weeds or crops grown

outside the normal growing season. This is applicable to south Florida,

especially since year-round cultivation of crops is possible. The

application of selective insecticides for control of the Lepidopterous

pests only when needed instead of on the basis of a regular application

schedule would be very beneficial to the leafminer's natural enemies

since it would aid in the buildup of parasite populations.














CHAPTER III

MECHANICAL DEFOLIATION OF THE TOMATO, Lycopersicon
esculentum MILL. CV. WALTER, AND ITS EFFECT
ON YIELD AND FRUIT QUALITY



Introduction


Several important tomato pests are foliage feeders although many

of them inflict damage directly to the fruit as well. Reduction of

marketable yield is, therefore, not solely related to the amount of

foliage consumed. Damage by the vegetable leafminer, Liriomyza sativae

Blanchard, in contrast, is restricted to the leaves and the injury is

different from that caused by most foliage feeders. Only the leaves'

mesophyll is consumed (even the spongy tissue remains for the most part

untouched) leaving both upper and lower epidermis intact. The presence

of a large number of leafminer larvae within one leaf may result in such

a serious impairment of the functions of this plant organ that leaf

death and subsequent abscission occurs. In addition to this type of

damage, yellowing and necrosis of the leaf tissues in the mines'

vicinity may occur, even if the larval population is small, again

possibly resulting in abscission of the entire leaf. It is clear that,

in the case of serious leafminer infestations, partial or even complete

defoliation of tomato plants may occur.

In recent years the population of the vegetable leafminer has become

sc large that growers consider this insect as their most serious pest








(Pohronezny et al., 1978b). Because of the clearly visible damage

inflicted on tomato plants, a negative effect on the yield is often

suspected. However, it has been shown repeatedly that consumption

of leaves and other plant tissues by insects does not necessarily

reduce plant vigor or reproductive capacity (Harris, 1972). In

fact, Harris (1974) suggested that sometimes a certain density of

"pest" insects may be required for a crop to attain its maximum yield.

Potato yield increase following partial defoliation has been

demonstrated (Skuhravy, 1968). Despite many attempts to find a

correlation between leafminer damage and tomato yield no consistent

results were obtained. Naturally occurring leafminer populations and

insecticide-induced populations have been found to have no significant

effect on tomato yield (Levins et al., 1975; Schuster and Everett, 1977;

Johnson et al., 1980a,b) although in some fields and in some years yield

reduction was found (Wolfenbarger and Wolfenbarger, 1966; Schuster et

al., 1976).

Many references have been made to serious leafminer damage of

cultivated plants (Spencer, 1973a; Spencer and Stegmaier, 1973) but only

on a few occasions has an indirect reduction of yield of a crop plant

been demonstrated. A loss in cash value of plants which are grown for

their foliage is obvious. These crops include foliage ornamentals,

celery (Musgrave et al., 1976), cabbage, lettuce (Musgrave et al., 1975),

and alfalfa (Jensen and Koehler, 1970).

The greatest damage by leafminers is often considered to be done

to seedlings or young plants which, as a result of weakening, may die

or become stunted (McGregor, 1914; Elmore and Ranney, 1954; Adlerz, 1961).

Severe damage by leafminers to cantaloups, resulting in complete crop








loss (Hills and Taylor, 1951), and to honeydew melon, resulting in

reduction in yield and fruit quality (Michelbacher et al., 1951), have

been reported.

Defoliation by means other than insect injury has also been found

to have varying effects on fruit production in tomato. The various

levels of defoliation caused by Alternaria blight controlled to

varying degrees with fungicides, appeared not to be correlated with

tomato yield (Richards, 1947). Defoliation by Xanthomonas vesicatoria

(Doidge) Dows. resulted in significant reduction of fruit size

(Pohronezny et al., 1978a). One commercial variety of tomato could

withstand considerable foliar damage due to ozone exposure for a

long period of time (Oshima et al., 1975) without significant

reduction in fruit size, weight or number, even though the fresh

weight of stems and leaves was lowered by 27%. Even a decrease in

fresh weight of 62% seemed to have minimal effects on yield.

Mechanical defoliation of tomato plants to study its effect on

yield has been performed several times. Wiebe (1970) found a

significant yield reduction in greenhouse tomatoes, when all except

the top 2 feet of leaves were removed, when compared to plants with

only the senescent leaves taken off. Selective removal of over-

lapping leaves had no effect on yield. A yield reduction, especially

in the largest fruit size categories was found as a result of

repeated defoliation at high levels (60% or more) in staked tomatc

plants (Jones, 1980).

The effects of mechanical defoliation on yield and fruit

quality of unstaked tomatoes, as they are grown commercially in

Dade County, Florida, have not previously been studied.


__








Actual damage to tomato plants by the leafminer-disease complex

occurs gradually, sometimes over a considerable period of time.

Exact duplication of this damage is virtually impossible so that

simulation by mechanical defoliation may not show the effect of

natural defoliation completely (Capinera and Roltsch, 1980).

The study presented here was undertaken to determine:

(1) the times at which unstaked tomato plants are most

sensitive to defoliation,

(2) the damage threshold at which unstaked tomato plants

will show significant loss in yield and fruit quality

when

(a) defoliated only once, and

(b) defoliated repeatedly.



Materials and Methods



General


Tomatoes, cv. Walter, were planted in 1977 and 1978 at the

University of Florida Agricultural Research and Education Center in

Homestead, Dade County, Florida. After metribuzin was incorporated

into the soil at a rate of 0.84 kg ai/ha, seedbeds were prepared in

groups of seven with their midlines 182 cm apart. Irrigation pipes

with frost protection nozzles were set on the middle bed. The other

beds were fertilized with 7-14-14 at a rate of 2242 kg/ha placed in

two bands 30 cm apart. For the spring crop of 1978 and the spring

crop of 1979 the beds were fumigated w'th Dowfume MC3V at a rate

of 314 kg/ha; for the fall crop of 1978 the rate was 247 kg/ha.








Immediately after the fumigation, the beds were covered with plastic

mulch, and, simultaneously, drip tubing for irrigation was placed

approximately 15 cm in the soil below the plastic. Tomato seeds were

planted with a seed drill 30 cm apart in the rows. One to two weeks

after emerging the seedlings were thinned to one plant per hill.

The foliage was removed by cutting the leaves off at the distal

end of the petiole with scissors. The fresh leaf weight was consistent-

ly found to be highly correlated to the total leaf area (Romshe, 1939).

For the one-time defoliations the fresh weight of the foliage removed

from the completely defoliated plants was used as a reference for

removal of the correct amount from the other plants to be defoliated.

From all but the outer two plants of each plot all mature green

and colored fruit was harvested three times except for the spring crop

of 1979 which was harvested only twice because of poor fruit set. The

first harvest was initiated when approximately 5% of all fruit present

showed color. The fruit was then graded into USDA grade 1 or 2 after

all culled fruit had been removed. These were then sized as extra large,

large, medium, small, and very small according to the measurements given

in Table 2. The culled fruit was subdivided into several types:

misshapen, blemished, sunscalded, decayed, damaged by insects or slugs,

and showing gray wall.


One-Time Defoliation


The defoliation experiments were conducted utilizing a split-plot

randomized complete-block design. Rows were assigned at random within

each of the 4 blocks for defoliation at one particular time.

Defoliation levels were assigned at random to the subplots within each








Table 2. Size ranges and mean weights
of 'Walter' tomatoes.


of the size categories


Size Size range Mean weight
category in mm in grams


very small 48 54 67

small 54 58 99

medium 58 64 142

large 64 73 174

extra large 73 213


Source: Marlowe (1978).








whole plot (row). Each subplot consisted of 12 plants in the spring

crop of 1978, of 22 plants in the fall crop of 1978 and of 17 plants in

the spring crop of 1979. Each subplot of plants except for the control

group was defoliated only once, and those in each row in a block on a

different date. Defoliation levels investigated were total (= 100%),

20%, 40%, 60%, and 80% starting from the top of the plant (= 20% upper

or 20U; 40% upper or 40U; etc.), or 20%, 40%, 60%, and 80% starting

from ground level (= 20% lower or 20L; 40% lower or 40L; etc.).

Yield data were analyzed and comparisons with the control were

made as a two-sided test using the Dunnett's procedure (Steel and

Torrie, 1960).


Experiment 1. Spring crop 1978. The tomato seeds were planted

on November 3, 1977. Beginning November 10, 1977, pesticides were

applied twice weekly by a high volume, low concentrate boom sprayer.

The insecticide permethrin (FMC 33297) was used at alternate rates of

.056 kg ai/ha and .112 kg ai/ha. The fungicide applied simultaneously

with the insecticide was either chlorothalonil (Bravo) at a rate of

1.58 kg ai/ha or mancozeb (Dithane M45 at a rate of 1.34 kg ai/ha.

Form-a-Turf~was applied at a rate of 7.02 1/ha when bacterial

diseases threatened (Pohronezny et al., 1979).

The times of defoliation were: 30 days after planting, 40 days

after planting and so on with 10 days intervals up to and including

100 days after planting. The levels of defoliation were: 100%,

80% upper, 80% lower, 60% upper, 60% lower, 40% upper, 40% lower,

20% upper, and 20% lower.

Harvesting was done between February 14, 1978, and March 23, 1978.








Experiment 2. Fall crop 1978. The tomato seeds were planted on

September 13, 1978. A mixture of permethrin (Ambush) at a rate of

.112 kg ai/ha and either chlorothalonil at 1.68 kg ai/ha or mancozeb

at 1.34 kg ai/ha was applied weekly, and Form-a-Turl on demand as in

Experiment 1.

The times of defoliation were: 30 days after planting, 40 days

after planting and so on with 10 day intervals up to and including

80 days after planting. The levels of defoliation were 100%, 80% upper,

80% lower, and 60% upper.

Fruit was harvested between December 8, 1978, and December 28, 1978.


Experiment 3. Spring crop 1979. The tomato seeds were planted on

December 28, 1978. Pesticide applications were made at the same schedule

and rates as in Experiment 2. Due to the very poor stand of the crop

only a limited area of the field could be used. The number of

defoliations, therefore, had to be limited. The times of defoliation

were: 70 days after planting, 80 days after planting, and 90 days after

planting. The levels of defoliation were: 100%, 80% lower, 60% lower,

and 40% lower.

Fruit was harvested between April 23, 1979, and May 2, 1979.



Repeated Defoliation


The defoliation experiments were conducted on the fall crop of

1978 using a randomized complete-block design. Defoliation levels were

assigned at random within each of the 3 blocks. Plants were treated on

several days by removing the required percentage of the foliage present

on the day of defoliation from the appropriate part of the plants. Each









plot consisted of 22 plants. One plot in each block was not mechanically

defoliated and functioned as the control.

The tomato seeds were planted on September 13, 1978. Pesticide

applications were made as described in Experiment 2.

Fruit was harvested between December 8, 1978, and December 28, 1978.


Experiment 4. Tomato plants were partially defoliated at 30, 50,

and 70 days after planting. The levels of defoliation were 60% lower,

40% upper, 40% lower, and 20% upper.


Experiment 5. Tomato plants were partially defoliated every 10

days, for the first time at 30 days after planting and for the last time

at 80 days after planting. The levels of defoliation were 40% upper,

40% lower, and 20% upper.



Gross Revenue Computation


The computation of the gross income per hectare was based on the

total amount of marketable fruit harvested in the first two pickings

in the sizes extra large, large, medium, and small. The prices used

for each size are listed in Table 3, and are based on market prices

in the season 1978-79 for Dade County, Florida.



Results



Experiment 1


Defoliation from mid-season on had a striking effect on the fruit

set if the defoliation levels were 60% upper, 80% or 100%. In nearly








Table 3. Prices in dollars per 13.6 kg box of tomato fruit of the
four main size categories and two grades.

Grade Size Low Medium High
category price price price


USDA 1 small 3.00 4.50 8.50

medium 4.00 6.50 12.50

large 5.00 9.00 15.00

extra large 6.00 10.00 16.00


USDA 2 small 3.00 4.00 6.50

medium 4.00 6.00 9.50

large 4.50 7.00 12.50

extra large 6.00 8.00 12.50


Source: H. H. Bryan, personal communication, 1980.








all these cases fruit set was significantly reduced to below that of the

control (P < 0.05). The most severe reduction was at the 100% level at

the beginning of the last third of the growing season (Table 4). Early

defoliation had no significant effect on fruit set.

The analyses of all the extra large fruit harvested in both first

and second pickings (Tables 5 and 6) and of the large fruit harvested

in the second picking only (Table 9), showed that very few treatments

resulted in significant yield reductions in these size categories. The

variation among the usually small number of fruit in these sizes was

large. Combining the yields of the two largest fruit categories also

did not reveal any significant reductions (Table 12). Whenever

defoliation took place the yield of the large fruit and the combination

of the two largest fruit classes showed significant reduction in many of

the highest levels of defoliation in the first harvest (Tables 8 and 11).

The plants were especially sensitive to leaf removal early in the first

half (30 days after planting) and early in the second half (60 to 80

days after planting) of the season. The amount of medium size fruit,

especially in the first picking, was affected by defoliation at any

level during the last few weeks before harvesting (Table 14). Generally

speaking defoliation earlier in the season, of 60% or higher led to

serious yield reduction in both first and second pickings (Tables 14 and

15).

Plants defoliated 30 days after planting at the three highest

levels still showed lush growth at the time of the harvests while

leaves of the other plants were senescent to varying degrees. Analysis

of variance of the fresh weight of all above ground parts of the plants

most severely defoliated at 30, 60 and 100 days after planting, after








all fruit had been removed, showed a significantly higher value for the

plants defoliated early in the season than for the plants in the control

(Table 17). No significant difference was found in any of the other

times or levels of defoliation tested.

The analysis of the gross revenue per hectare (Tables 18 and 19)

illustrate the detrimental effect of high levels of defoliation

(60% or more) carried out at any time during the season. In the first

harvest losses can also be expected to occur if the foliage loss takes

place late in the season even at lower levels.

The average weight of all marketable fruit was significantly

reduced only in the first harvest for the 80% upper and 100%

defoliations 30 days after planting. The total weight of all marketable

fruit showed a reduction pattern very similar to that of the gross

revenue pattern, the latter based on different prices for the various

size categories (compare Tables 18 and 20, and Tables 19 and 21). For

both first and second harvests and for all price ranges a highly

significant correlation exists between the two variables, total weight

and gross revenue.

No treatment resulted in significant increases or decreases in the

weight of the culled fruit in the first two harvests or in any

particular cull category. Only in the third picking there were

significantly more misshapen fruit present on plants defoliated at the

80% and 100% levels early in the season (30 and 40 days after planting).

In many cases where defoliation significantly reduced the fruit

weight, this reduction was more noticeable in the USDA grade 1 fruit

than in the USDA grade 2 fruit, especially in the two largest fruit

categories (Tables 5, 8 and 11).








If the data from the first two harvests are combined (Tables 7, 10,

13 and 16) the impact of mechanical defoliation on the grower's yield

can be summarized. It shows that the effect is most pronounced in the

plants at first harvest. However, increases in the second harvest

tend to compensate for the loss in yield in the first.


Experiment 2


In the first harvest the weight of both the extra large (Table 22)

and the large fruit (Table 24) was significantly reduced by defoliation

in the first month of the season at all levels investigated and in the

second month of the season only at the 100% level. No effect at all

was noted for defoliation in the last month of the season except a

possible yield increase. In the second harvest no significant yield

reduction in the extra large fruit was observed (Table 23) while the

weight of the large fruit was reduced significantly (Table 25)

especially in the very high levels of defoliation (80% or more) in the

last two months of the season. The 60% upper defoliation had a

significant effect on the USDA grade 1 fruit only, the most severe

reduction occurring from defoliation at 50 days after planting.

Analysis of the combination of the two largest fruit sizes

(Tables 26 and 27) summarize the differences in effects of defoliation

in the first two harvests.

A reduction in the weight of the medium sized fruit as a result

of foliage removal occurred only in the second harvest (Tables 28 and 29).

The reduction pattern was very similar to that of the large fruit.

Analysis of both the total weight of all marketable fruit and of

the gross revenue per hectare show significant reductions at defoliation








levels and times at which the weight of the extra large and large fruit

was also reduced (Tables 30 and 31; Figures 1 and 2). As in

Experiment 1 a very close correlation existed between the total fruit

weight and the gross revenue based on different prices for different

size categories.

From combining the total yields of the first two harvests

(Figure 3) it appears that the only significant reduction due to 50%

defoliation occurred in plants defoliated 50 days after planting.

The average weight of all marketable fruit was significantly

reduced only in the first harvest by 100% defoliation of the tomato

plants 50 days after planting. In the second harvest no

significant reductions were found.

The weight of all culled fruit together in any of the treatments

showed no significant difference from the control in any of the

harvests. However, more sunscalded and decaying fruit were present

on plants defoliated 15 days before the first harvest at the 30% and

100% levels. Defoliation at 80 days after planting also resulted in more

decaying fruit when 80% or more of the foliage was removed from the

plants.


Experiment 3


Since only late-season defoliations could be examined for effects

on yield and fruit quality, differences in reduction patterns as found

in the first two experiments could not be verified. In fact, analysis

of the weight of the various fruit sizes, total weight, and gross

revenue revealed only very few significant reductions when compared to

the control.








No significant differences in the weight of the extra large

fruit was detected (Figure 4). No extra large fruit was harvested

in the second picking.

The weight of the large fruit was only reduced significantly

by defoliating plants 80 days after planting at the 100% level

in the first picking, while in the second picking the reduction

was only significant when the plants were completely defoliated

90 days after planting (Figure 5). The latter was also the case

for the weight of the medium size fruit in the second harvest

(Figure 6).

Combining the weight of all extra large and large fruit for

analysis showed a reduction by complete defoliation 70 days or

80 days after planting (Figure 7).

Both the weight of all marketable fruit (Figures 8 and 9)

and the gross revenue per hectare (Tables 32 and 33) were signif-

icantly reduced by 100% defoliation at 80 or 90 days after planting.

Total weight of the culled fruit was not significantly different

between any of the treatments and the control. The weight of the

culled fruit as a percentage of the total marketable fruit plus

culls was significantly higher only in the second harvest if the

tomato plants were completely defoliated 80 or 90 days after planting.

Significantly more sunscalded fruit occurred on plants defoliated 90

days after planting at the 60% and higher levels of defoliation.

Weight of the decaying fruit was significantly greater than the control

in plants completely defoliated 70 or 90 days after planting in the


first picking.








Experiment 4


When tomato plants were defoliated three times during the

growing season, the threshold for reduction in fruit weight per

plant in several size categories was lower than that found in

one-time defoliation experiments.

Analysis of all extra large fruit in the first harvest showed

that the repeated removal of 40% from the upper part and 60% from

the lower part of the foliage of the tomato plants had the same effect

on yield in this fruit size. Removal of 40% of the foliage starting

at soil level also reduced the yield, but not as severely (Figure 10).

In the second harvest significant differences in the yield of

extra large fruit also occurred (Figure 10) but because of the low

total yield in this size category its effect on the overall fruit

yield in this harvest was negligible (Figure 14).

The yield of large fruit in the first harvest was also signifi-

cantly reduced at some defoliation levels (Figure 11) but not as

severely as that of the extra large fruit. The effect of defoliation

on the yield in the two largest size categories is summarized in

Figure 12.

Reduction in the weight of the medium size fruit in the first

(Figure 13) had only a minimal effect on the total yield (Figure 14).

In the second picking the increase in the weight of medium size fruit

and large fruit accounted for the significant increase of the total

yield.

The total yield of the first two harvests was not significantly

reduced by defoliation at any level (Figure 15) while the combined

weight of all extra large and large fruit was significantly reduced

only at the 60% defoliation level.








Experiment 5


Frequent defoliation of tomato plants resulted in a reduction of

the yield of extra large fruit at all levels tested (Figure 16) and of

the large fruit at the 40% level (Figure 17) in the first harvest only.

Reduction was evident in all defoliation levels when the two largest

size categories were combined for analysis (Figure 18).

The yield of the medium size fruit was only significantly reduced

as a result of removal of 40% of the foliage from the upper part of the

plants (Figure 19).

In no size category was a yield reduction observed in the second

harvest.

The total yield loss as a result of repeated defoliation (Figure

20) was mainly caused by the fewer extra large and large fruit harvested

and although in the second harvest no significant differences were

observed, the larger fruit weight removed from the plants at that time

compensated for the lesser weight harvested in the first, since

combining the yield of the first and the second harvests showed no

significant differences (Figure 21).



Discussion


Defoliation of unstaked tomato plants revealed a changing

sensitivity to this type of damage in the course of their development

as was demonstrated for potato by Skuhravy (1968) and sugarbeet by

Capinera (1979).

Damage early in the development, before or at anthesis, when most

of the metabolic activity of the plant is directed at vegetative
























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Table 17. Influence of defoliation of 'Walter' tomato plants on the
mean fresh weight of all above ground plant parts, excluding
fruit, at the time of completion of the third harvest of the
spring crop of 1978.


Mean fresh weight per plant (in grams)a'b

Time of defoliation (in days after planting)
Defoliat on
level 30 60 100


80% lower 11062* 8080 6928

80% upper 10960* 7100 8175

100% 14177** 7595 7785


aMean fresh weight per control plant: 8545 g.

bWeight is significantly different from the control if indicated by *
for P < 0.05 or by ** for P < 0.01.
CFor an explanation of the defoliation level codes see page 26.







Table 18. Gross _revenue in dollars per hectare of 'Walter' tomatoes
based on all marketable fruit of sizes extra large through
small. Influence of defoliation on the first harvest of
the spring crop of 1978.

Gross revenue in dollars per hectarea

Time of defoliation (in days after planting)
Defoliation Price
level rangec 30 40 50


20% lower


low
medium
high


20% upper low
medium
high

40% lower low
medium
high

40% upper low
medium
high

60% lower low
medium
high

60% upper low
medium
high

80% lower low
medium
high

80% upper low
medium
high

100% low
medi um
high


8082.09
12637.24
21392.33

6545.31
10280.84
17327.81

5382.35
8378.81
14092.30

4756.31
7543.69
12751.90

5453.36
8520.16
14178.30

3991.70
6143.33
10387.59

4358.11
6787.66
9246.99*

2348.83**
3619.87**
6215.82**

805.63**
1242.04**
2126.91**


6550.75
10303.40
17327.15

6193.58
9754.46
16428.61

4579.86
7279.11
12251.89

6316.15
10130.75
17052.51

4049.70
6440.70
10791.39

5667.20
9069.60
15295.96

3854.30
6185.50
10316.42

2951.32**
4644.44**
7791.31**

1397.73**
2236.14**
3722.67**


5324.03
8493.80
14268.75

3872.43
6281.72
10748.39

6870.04
10919.89
18362.76

4511.49
7076.46
11725.68

5620.91
8852.95
14785.40

5267.03
8346.68
14047.32

6331.63
10011.80
16681.33

5259.78
8123.45
13707.45

3517.23*
5362.91*
8967.95*


aA significant difference from the control is indicated by for
P < 0.05 or by ** for P < 0.01.
For an explanation of the defoliation level codes see page 26.







Table 18. Extended


Gross revenue in dollars

Time of defoliation (in days


per hectarea

after planting)


60 70 80 90 100


6173.47
9767.64
16265.67

6147.28
9617.88
16063.85

5095.03
8046.67
13434.95

4591.06
7398.22
12597.86

5419.59
8656.90
14481.44

4337.84
7106.12
12011.36

2665.81**
4296.49**
7263.29**

2542.74**
4154.81**
7114.36**

1923.61**
3087.23**
5310.85**


CPrices used


6009.88
9644.90
16276.05

5318.10
8434.99
14265.29

5408.71
8472.88
14299.88

4845.43
7845.84
13281.38

2693.48**
4081.66**
6870.53**

4331.91
6742.85
11282.84

2128.23**
3315.41**
5608.72**

3098.60*
4803.75**
8036.95**

1491.64**
2285.07**
3845.90**


6039.37
9527.27
15813.27

6807.10
10794.19
18167.70

5311.34
8200.71
13663.95

4057.60
6443.66
10778.38

4990.19
7772.03
13029.51

2842.58**
4417.75**
7327.38**

3211.95*
4963.23*
8288.86**

2298.74**
3598.78**
6025.53**

1466.60**
2297.76**
3790.05**


2433.01**
3643.43**
6048.10**

3296.79*
5132.76*
8572.06*

2788.05**
4222.35**
7027.70**

3680.82
5693.40*
9493.99*

4065.18
6541.85
10983.16

3612.12
5712.84
9548.36*

3348.36*
5350.39*
9150.98*

2725.11**
4259.09**
7187.50**

2522.31**
4051.18**
6796.89**


3177.35*
4884.31**
8159.86**

2662.18**
4083.64**
6791.28**

3964.52
6119.93
10155.79

2952.63**
4537.35**
7474.33**

5113.31
8183.41
13747.98

4244.26
6678.10
11109.36

3748.70
5976.93
9943.92

2931.05**
4535.05**
7599.87**

2880.97**
4466.68**
7453.91**


in the computations are given in Table 3.


Note: Gross revenue per hectare of the control plants was for the low
price range, $6194.11, for the medium range, $9829.29, and for
the high range, $16607.44


~----








Table 19. Gross revenue in dollars per hectare of 'Walter' tomatoes
based on all marketable fruit of sizes extra large through
small. Influence of defoliation on the second harvest of
the spring crop of 1978.

Gross revenue in dollars per hectarea

Time of defoliation (in days after planting)
Defoliation Pricec
level range 30 40 50

20% lower low 4563.55 5193.22 6543.17
medium 7104.97 8039.75 10308.02
high 12227.51 13900.53 17950.72

20% upper low 4059.58 5135.56 5833.43
medium 6412.53 8048.32 9111.45
high 11250.22 13881.75 16003.55

40% lower low 7001.67 5337.04 6090.78
medium 11108.37 8376.83 9568.95
high 19569.22 14530.70 16476.06

40% upper low 5774.95 5781.54 5732.94
medium 9103.87 9070.75 9001.06
high 15836.17 15851.00 15687.90

60% lower low 7004.47 4477.72 5343.14
medium 11105.89 6946.81 8374.69
high 19362.29 12010.86 14674.69

60% upper low 3737.50* 5059.28 3982.64
medium 5796.53* 8021.46 6185.83
high 10259.42* 14072.86 10740.81

80% lower low 2570.91** 3413.60* 6723.57
medium 3989.40** 5286.63** 10580.51
high 6938.90** 9039.12** 18171.65

80% upper low 2426.76** 3676.05* 1704.49**
medium 3763.36** 5729.15* 2602.21**
high 6585.18** 9997.80* 4614.78**

100% low 1595.76** 3260.88** 2719.68**
medium 2469.10** 5088.44** 4171.45**
high 4322.69** 8866.79** 7113.04**

aA significant difference from the control is indicated by for
P < 0.05 or by ** for P < 0.01.
bFor an explanation of the defoliation level codes see page 26.








Table 19. Extended


Gross revenue in dollars per hectarea

Time of defoliation (in days after planting)

60 70 80 90 100


5998.51
9313.43
16178.68

6105.44
9500.42
16398.13

6074.29
9670.44
16904.73

5612.67
8740.10
15059.05

5828.49
9158.40
16080.99

5720.42
9020.13
15500.08

4547.24
7174.33
12249.09

5702.62
8980.14
15615.57

3101.73**
4760.92**
8075.67**


5101.78
8080.12
14145.02

5359.28
8417.19
14726.59

4353.30
8468.43
14808.63

5186.47
8248.49
14323.77

5634.25
9012.27
15584.60

4041.95
6392.43
10943.62

4602.10
7225.73
12505.11

3233.86**
4956.31**
8444.54**

2773.88**
4247.23**
7173.83**


5819.10
9117.71
15908.33

3796.15
5738.87*
9851.99*

5855.84
9186.73
15841.28

5752.87
9039.45
15933.21

4310.82
6779.92
12185.33

5883.35
9293.65
16007.51

5306.89
8399.23
14574.35

4486.12
7044.50
12366.56

2835.33**
4346.08**
7478.12**


7340.06
11550.22
19939.24

4976.57
7778.30
13439.07

6083.85
9729.58
17019.07

4440.31
6935.28
12216.80

5208.09
8053.43
13803.00

4826.49
7786.70
13607.61

3023.64**
4585.30**
7838.43**

3425.79*
5382.52**
9210.46**

2289.19**
3591.53**
6094.89**


6273.81
9846.72
17153.50

6812.87
10674.09
18326.84

6255.19
9832.39
17188.43

5391.91
8474.69
14736.47

5128.97
7931.51
13724.42

5462.75
8576.18
15051.30

4784.15
7433.97
12846.14

3267.80**
5081.03**
8841.09**

3533.70*
5538.86*
9654.95*


"Prices used in the computations are given in Table 3.

Note: Gross revenue per hectare of the control plants was for the low
price range, $6286.20, for the medium range, S9798.74, and for
the high range, $17064.17.
















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