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66 views65 pages

Complete Plant Growth Regulating Chemicals - Volume 1 First Edition Nickell PDF For All Chapters

Regulating

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Plant Growth
Regulating Chemicals
Volume I

Editor

Louis G. Nickell, Ph.D.


Vice President
Research and Development
Velsicol Chemical Corporation
Chicago, Illinois

Boca Raton London New York

CRC Press is CRC Press,


an imprint of the Inc.
Taylor & Boca Raton,
Francis Group, Florida
an informa business
First published 1983 by CRC Press
Taylor & Francis Group
6000 Broken Sound Parkway NW, Suite 300
Boca Raton, FL 33487-2742

Reissued 2018 by CRC Press

© 1983 by CRC Press, Inc.


CRC Press is an imprint of Taylor & Francis Group, an Informa business

No claim to original U.S. Government works

This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish
reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the
consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this
publication and apologize to copyright holders if permission to publish in this form has not been obtained. If any copyright material has
not been acknowledged please write and let us know so we may rectify in any future reprint.

Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form
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Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and
explanation without intent to infringe.

Library of Congress Cataloging in Publication Data


Main entry under title:

Plant growth regulating chemicals.

Includes bibliographies and index.


1. Plant regulators. I. Nickell, Louis G.,
1921-
QK745.P56 1983  631.8  82-22832
ISBN 0-8493-5002-6 (v. 1)
ISBN 0-8493-5003-4 (v. 2)

A Library of Congress record exists under LC control number: 82022832

Publisher’s Note
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may be apparent.

Disclaimer
The publisher has made every effort to trace copyright holders and welcomes correspondence from those they have been unable to
contact.

ISBN 13: 978-1-315-89665-6 (hbk)


ISBN 13: 978-1-351-07575-6 (ebk)

Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the
CRC Press Web site at http://www.crcpress.com
PREFACE

The need to increase the world food supply substantially by the end of this century poses
one of the greatest challenges yet faced by man. Many agricultural scientists believe that
this challenge can be met , and it is expected that plant growth regulators will play an
increasingly important role in meeting this challenge.
Plant growth regulating chemicals are used to modify crops by changing the rate or pattern
or both of their response(s) to the internal and external factors which govern all stages at
crop development from germination through vegetative growth, reproductive development,
maturity, and senescence or aging , as well as post-harvest preservation.
The purpose of this two-volume work is to make available both to the investigator and
to the user , on a crop by crop basis, the latest information on the use of chemicals to regulate
plant growth and development. Emphasis is given to the major crops and to those with which
the most success has been achieved. Since the degree of practical success with each crop
varies, primary attention is given to chemicals registered for specific use(s) with the particular
crop discussed. Also included is information concerning chemicals not yet registered , but
for which practical results are available. In some cases information concerning active com-
pounds in the exploratory stages is included. Where known and pertinent, information
concerning mode of action is included .
The obvious classifications to use in presenting data on effectiveness of plant growth
regulating chemicals are (I) by crop, (2) by chemical class, and (3) by plant function or
process . Essentially all major summary or survey publications to date have been based on
the plant function or process approach . This is primarily an academic approach and is not
nearly as useful for practical purposes as a presentation by crops, as is done in this publication .
THE EDITOR

Louis G. Nickell, Ph.D., Vice President of Research and Development, Velsicol Chem-
ical Corporation, Chicago, Illinois, was born July 10, 1921, in Little Rock, Arkansas. He
received his B.S. degree in botany from Yale University in 1942. After serving 4 years in
the U.S. Marine Corps as a regular commissioned officer, he returned to Yale University,
receiving his M.S. in microbiology in 1947 and his Ph.D. in plant physiology in 1949. He
is married to Natalie Wills Nickell and has three children and four grandchildren. His first
professional experience was as Research Associate at the Brooklyn Botanic Garden from
1949 to 1951 where he was engaged in research on plant tissue culture and plant growth
substances. He joined industry in 1951, going to Pfizer, Inc. in Brooklyn as its Plant
Physiologist and Assistant Mycologist. There he specialized in antibiotics and their effects
in agriculture as well as plant tissue and cell culture. In 1953 he became Head of Pfizer's
Phytochemistry Laboratory and received the first patent issued for the use of plant cell
cultures for the production of secondary products. In 1961 he moved to Hawaii to become
Head of the Plant Physiology and Biochemistry Department of the Hawaiian Sugar Planters'
Association, becoming its director of research in 1965. His first commercial success with
plant growth regulating chemicals was the registration of diquat for the prevention of flow-
ering in sugarcane in the early 1960's. This was followed successively by the registration
of gibberellic acid for increasing the sugar yields in cane and later by the development of
the first commercial product for the ripening of sugarcane, glyphosine. In 1975, he joined
the Research Division of W. R. Grace & Company as Vice President of its Research Division
in charge of agricultural, biological, and medical research, development, and commercial-
ization. In 1978, he joined Velsicol Chemical Corporation as Vice President of Research
and Development- his present position. His publications (over 300) and patents (over 30)
have been primarily in the area of plant cell and tissue culture and the regulation of plant
growth through the use of chemicals. He is the author of a book published in early 1982
entitled Plant Growth Regulators -Agricultural Uses.
He has served as President of the Hawaii Academy of Science, as Chairman of the Hawaiian
Section of the American Chemical Society, as Vice Chairman and Chairman of the Plant
Growth Regulator Society of America, as Council Member of the Society for Economic
Botany, and has been the Treasurer of the American Society of Plant Physiologists since
1976. He has served as Chairman of the Governor's Advisory Committee on Science and
Technology in Hawaii, as a member of the National Academy of Sciences - National
Research Council Committee on Agricultural Production Efficiency, and is Chairman of
the "The Forward Edge" Session of CHEMRA WN II, the International Conference on
Chemistry and World Food Supplies: The New Frontiers, and a member of the Editorial
Board of the Journal of Plant Growth Regulation.
CONTRIBUTORS

James E. Baker, Ph.D. Martha Davis, Ph.D.


Research Plant Physiologist Postdoctoral Research Associate
Plant Hormone Laboratory Agronomy Department
Plant Physiology Institute University of Arkansas
Agricultural Research Service Fayetteville, Arkansas
U.S. Department of Agriculture
Beltsville , Maryland E. F. Eastin, Ph.D.
Professor of Weed Science
Duane P. Bartholomew Texas Agricultural Experiment Station
Associate Agronomist Beaumont, Texas
Department of Agronomy and Soil
Science Donald M. Elkins, Ph.D.
Unversity of Hawaii Professor of Plant and Soil Science
Honolulu, Hawaii Southern Illinois University
Carbondale, Illinois
Wolfgang D. Binder, Ph.D.
Tree Physiologist Gail Ezra, Ph.D.
Research Branch Research Associate
British Columbia Ministry of Forests Department of Environmental Biology
Victoria, British Columbia University of Guelph
Canada Guelph, Ontario
Canada
Kenneth Bridge
E. Hayman, Ph.D.
Product Development Manager
Research Chemist
Plant Growth Regulators
Agricultural Research Service
Union Carbide Agricultural Products
Fruit and Vegetable Chemistry Laboratory
Company, Inc.
U.S. Department of Agriculture
Raleigh , North Carolina
Pasadena, California
George W. Cathey
W. J. Hsu
Plant Physiologist
Research Chemist
Cotton Physiology and Genetics Research
Agricultural Research Service
Unit
Fruit and Vegetable Chemistry Laboratory
Agricultural Research Service
U.S. Department of Agriculture
U.S. Department of Agriculture
Pasadena, California
Stoneville, Mississippi
Johannes Jung, D.Sc.
John A. Considine, Ph.D. BASF Agricultural Research Center
Senior Viticultural Research Officer Limburgerhof
Department of Agriculture Federal Republic of Germany
Ferntree Gully , Victoria
Australia Darold L. Ketring
Plant Physiologist
Richard A. Criley, Ph.D. Agricultural Research Center
Professor of Horticulture U .S . Department of Agriculture
Department of Horticulture Department of Agronomy
University of Hawaii Oklahoma State University
Honolulu, Hawaii Stillwater, Oklahoma
N. E. Looney, Ph.D. Gilbert F. Stallknecht, Ph.D.
Pomologist and Plant Physiologist Superintendent! Agronomist
Agriculture Canada Research Station Montana State University
Summerland, British Columbia Huntley, Montana
Canada
George L. ·Steffens
Louis G. Nickell, Ph.D.
Plant Physiologist
Vice President
Plant Hormone and Regulators Laboratory
Research and Development
U.S . Department of Agriculture
Velsicol Chemical Corporation
Beltsville, Maryland
Chicago, Illinois

EdwardS. Oplinger, Ph.D. Gerald R. Stephenson, Ph.D.


Professor of Agronomy Professor of Weed Science
University of Wisconsin Department of Environmental Biology
Madison, Wisconsin University of Guelph
Guelph , Ontario
Richard P. Pharis, Ph.D. Canada
Professor of Biology
University of Calgary
Charles A. Stutte, Ph.D.
Calgary, Alberta
Distinguished Professor
Canada
Altheimer Chair for Soybean Research
Agronomy Department
S.M. Poling
University of Arkansas
Chemist
Fayetteville, Arkansas
Agricultural Research Service
Fruit and Vegetable Chemistry Laboratory
U.S. Department of Agriculture W. C. Wilson
Pasadena, California Research Scientist III
Acting Harvest Coordinator
Wilhelm Rademacher, D.Sc. Florida Department of Citrus
BASF Agricultural Research Center Lake Alfred, Florida
Limburgerhof
Federal Republic of Germany
S. H. Wittwer, Ph.D.
Director
Stephen A. Ross, Ph.D.
Agricultural Experiment Station
Senior Tree Physiologist
Michigan State University
British Columbia Ministry of Forests
East Lansing, Michigan
Research Branch
Victoria, British Columbia
Canada Henry Yokoyama, Ph.D.
Research Chemist
Otto John Schwarz, Ph.D. Fruit and Vegetable Chemistry Laboratory
Associate Professor of Botany Agricultural Research Service
University of Tennessee U.S. Department of Agriculture
Knoxville, Tennessee Pasadena, California
TABLE OF CONTENTS

Volume I

Chapter 1
Growth Regulator Usage in Apple and Pear Production .. . . .. ........................... .
Norman E. Looney

Chapter 2
Growth Regulator Use in the Production of Prunus Species Fruits ........ ..... . . . . . . ... 27
Norman E. Looney

Chapter 3
Plant Growth Regulator Use in Natural Rubber (Hevea brasiliensis) .............. . ..... 41
Kenneth Bridge

Chapter 4
Bioregulation of Rubber Synthesis in Guayule Plant ... . . . . ....... ................... . .. 59
H. Yokoyama, W. J. Hsu, E. Hayman, and S.M. Poling

Chapter 5
Tobacco . ... . ... . .. .. .... . . . . .. . ... . .. . . . . . .... . . . .. ... . . .. . .. .. . . . . ..... . ... . ....... . . .. 71
George L. Steffens

Chapter 6
Concepts and Practice of Use of Plant Growth Regulating Chemicals in Viticulture . . . . 89
John A. Considine

Chapter 7
Sugarcane .............. .. . .... .. ... .. ... . . . . .. .... .. ..... . . ... . ... ..... . ...... .. ....... 185
Louis G. Nickell

Chapter 8
The Use of Exogenous Plant Growth Regulators on Citrus .............. . ......... ... .. 207
W. C. Wilson

Chapter 9
Cotton ............... ... . . . ... . ... . . ... ................ ... .. .... . . .. ............. . . ... .. 233
George W. Cathey

Chapter 10
Cereal Grains .. ....... . .. . . .. .. . . .. .. .. ....... . . . .. .. .. . . . . ............. .... . . . ... . . .. . . 253
J. J ung and W. Rademacher

Index ...... . ........... ... ... . ... . ........................... . .......................... 273

Volume II

Chapter I
Tropical Fruit and Beverage Crops ..... .. . ..... . . ..... . . . ... .. . . ..... . . ...... . . . . . ... ... .
D.P. Bartholomew and R. A. Criley
Chapter 2
Growth Regulators and Conifers: their Physiology and Potential Uses in Forestry .. .. .. 35
S. D. Ross, R. P. Pharis, and W. D. Binder

Chapter 3
Paraquat-Induced Lightwood Formation in Pine ............ . . .. ......................... 79
Otto J. Schwarz

Chapter 4
Growth Regulators in Soybean Production .... .... . .. . ........ .... . . . . . .. . ... . ... .. . . .. . 99
Charles A. Stutte and Martha D. Davis

Chapter 5
Growth Regulating Chemicals for Turf and other Grasses .. . .. . ....... . ...... . ... . .. ... 113
Donald M. Elkins

Chapter 6
Com . .............. . .......... ..... . . ................. . ........... .. ...... . ......... . .. . 131
Edward S. Oplinger

Chapter 7
Peanuts . . .. . .. .. ..... .. . .. .. . . . .. . .. . ... .. ... . ... .. . .. . . . ... . ... .. . . . .... .. .. . ... . ...... 139
Darold L. Ketring

Chapter 8
Plant Growth Regulators in Rice . ....................... . . . ..... . .............. . .... ... 149
E. Ford Eastin

Chapter 9
Application of Plant Growth Regulators to Potatoes: Production and Research .. . ... .. . 161
Gilbert F. Stallknecht

Chapter 10
Preservation of Cut Flowers . . .. .. .. .................. . .. . . ... . ... ................ ...... 177
James E. Baker

Chapter 11
Herbicide Antidotes: A New Era in Selective Chemical Weed Control ........... . . .... 193
G. R. Stephenson and G. Ezra

Chapter 12
Vegetables . . .. . . . .. . . ... . .. . . .. .. . . .. ... . . . . . .. . . .. .. . . . . .. . . .. . . ... .. .. . .. .. . . ... .. . . . . 213
S. H. Wittwer

Index . ................. .. .. . .. . ........................ . . .. . . . . .................... ..... 233


Volume I I

Chapter 1

GROWTH REGULA TOR USAGE IN APPLE AND PEAR PRODUCTION

Norman E. Looney

TABLE OF CONTENTS

I. Introduction ............ . ... .. ...... . . . ... .. . . .... . . . . . .. .. ...... .. ...... . .... . .. . 2

II. Growth Regulator Usage in the Nursery .. .... . . ... ..... ...... ....... ... ..... . . .. 2
A. Production of Clonal Rootstocks and Self-Rooted Trees . .... .. .. . .. ... .. . 3
B. Defoliation of Nursery Stock ........ . ............ .. ...... . ............... 5
C. Promotion of Feathering of Nursery Trees . . . . .. ... . . .......... ..... . . .. .. 6

III . Controlling Growth and Modifying the Structure of Orchard Trees ... .. . . . . .. . .. 6
A. Regulation of Tree Branching Habit . ... ..... ....... ....... . .. . ....... .... 6
B. Promotion of Spur Development and Flower Initiation . ..... . .. .. ........ 9
C. Suppression of Root Suckers . ...... .. .... . .... .... . . . . . ..... . . ... . ... . . .. 9
D. Suppression of Water Sprouts ... . ....... . ............ .. .. . ... . .......... 10

IV. Regulation of Flowering and Fruit Set .......... .. . . .... ...... ................. . 11
A. Improving Return Bloom . ... ......... . .. . ..... . .. . .... ... ......... . . . . .. 11
B. Increasing Fruit Set ... . . .. . . .. . . .. . .... . . .. . . . . . ... . ... .. .. . . . . . . . . . . . ... 11
C. Chemical Thinning ... ... ..... ..... .... .... ... .. ...... .. ...... .. ......... 12
l. Blossom Thinning of Apples .. .... ...... .. ....... .. ... .. ... . .... . 14
2. Postbloom Thinning with Hormone Materials .... ... ....... . . .. . . 15
3. Postbloom Thinning of Apples with Carbaryl . . ..... ... .. .. .... . . 16
4. Flower and Fruit Thinning with Ethephon . .... ........ .. .. . ..... . 16

V. Improving Fruit Condition and Appearance .... ..... ... ....... ....... . .. . ... . . .. 17
A. The Effects of Daminozide on Apple Fruits ........... .. .... .... ........ 17
B. Preventing Premature Softening of Pear Fruits .... ...... . .. . ..... .... . ... 18
C. Reduction of Fruit Russeting . .. ...... .. ....................... ..... ..... 18
D. Increasing the Length to Diameter Ratio of Apples . . .. ..... . .. ... ....... 19

VI. Preventing Preharvest Fruit Drop .. ..... . .. . . .... ..... .. . . ..... . . . . . . . .......... . 19
A. Control of Fruit Drop with Synthetic Auxins ...... ..... .... ... .. .. .... .. 19
B. Control of Fruit Drop with Daminozide ..... ...... .. .. .. .... . .. .. ....... 21

VII. Regulation of Fruit Ripening . ... .... . . . . .. . . ... . . ............ . ... .. .... . .. . ..... 22
A. Delay of Fruit Ripening with Daminozide ..... .. .............. .. ........ 22
B. Advancing Fruit Ripening with Ethephon ... . .. .... ..... .... ...... ....... 22
C. Product Interactions . . . ... . . . .... .. .. .. . . ................... . .... .. . . . . ... 22

VIII. Future Development Opportunities .... . . . ....... . ... . ..... . . . . . . . ............... 23

References .. . .. . ... . . .... ..... . . .. ...... .. . .. . .. .... . .. . .. . .. . . . ... . . .. . .. ... ....... .. .. . 25


2 Plant Growth Regulating Chemicals

I. INTRODUCTION

The range of plant growth regulating chemicals used by apple producers around the world
is unmatched by any other crop. Plant growth regulators are used in the production of nursery
trees, to regulate the shape and size of orchard trees, to promote flowering, and to increase
and reduce fruit set. Other treatments control the growth of unwanted shoots or suckers,
and still others prevent preharvest fruit drop, regulate fruit ripening, and influence various
aspects of fruit quality . The number of products and practices available to pear growers is
smaller but still impressive.
This high degree of interest among apple and pear producers in chemical growth regulation
is quite understandable. These are high value crops produced on expensive land with in-
creasingly intensive management systems and costly equipment. A relatively small increase
in orchard productivity or improvement in fruit quality will cover the cost of the treatment.
Secondly, apples and pears are produced on long-lived trees. The use of a growth regulator
to overcome a tree productivity or fruit quality deficiency in an existing planting is almost
always more economical than replanting the orchard. Thirdly, there are strong market pref-
erences for a few established apple and pear cultivars and introducing new cultivars is a
very slow process. Some growth regulator techniques have been developed specifically to
improve the performance of these well-established cultivars which , of course, perpetuates
both the cultivar and the growth regulator procedure. And finally , fresh market sales of all
fruits depend heavily on their appearance. Treatments that either directly or indirectly enhance
fruit appearance and condition are valued highly by producers and their marketing agents .
This discussion of growth regulator usage in apple and pear production is organized by
practice rather than by chemical. There are several reasons for choosing this format, an
important one being that some materials are used to achieve very diverse aims. For example,
naphthaleneacetic acid (NAA) in various formulations promotes the rooting of cuttings, is
an ingredient of fruit setting mixtures for some cultivars yet is used to reduce set of both
apples and pears, and it prevents abscission of mature fruits of both species. Applied as a
paint , NAA prevents the growth of unwanted sprouts and suckers.
The reader is cautioned that some of the practices and several of the materials mentioned
are either unimportant or unavailable to some large sections of the fruit growing community .
An example is the thinning of pear fruits. Producers in many areas use NAA or naphtha-
leneacetamide (NAAm) yet this practice is considered unnecessary in California, the state
that produces the most pears in North America. Obviously, it is difficult to generalize about
growth regulator practices . The materials, and especially the procedures for their use, differ
considerably from region to region. On the other hand, it is impossible to be as thorough
as the subject deserves. It is hoped that by surveying these practices and regional differences
the reader will see some opportunities for expanding the useful role of plant growth regulators
in worldwide apple and pear production.

II. GROWTH REGULATOR USAGE IN THE NURSERY

Commercial orchardists usually purchase finished trees from a local nurseryman. These
trees are produced by grafting or budding a scion piece of the cultivar of interest onto a
seedling or, preferably, a clonal rootstock with known characteristics. This rootstock is 1
or 2 years old before it is budded or grafted and the two-part tree is grown for at least
another year before it is sold to the orchardist. The production of self-rooted trees by tissue
culture methods may shorten this operation somewhat, but it introduces still other oppor-
tunities for error into what is already a highly technical business . Obviously, the successful
operation of a fruit tree nursery requires highly skilled technicians and constant attention to
detail. It is not surprising that nurserymen are keenly interested and innovative participants
in the plant growth regulator field .
Volume I 3

FIGURE I. Commercial production of fruit tree rootstocks by in vitro meristem culture. (Bottom) Mother cultures
of various apple and cherry rootstock clones held in a controlled environment room. (Top left) A proliferating
culture of EMLA 7 apple ready to ''harvest''. Rooting of these harvested shoots is accomplished by placing them
first on an auxin-containing and then on an auxin-free medium. (Top right) Rooted plants ready for potting.
(Courtesy of Dr. D . I. Dunstan , Kelowna Nurseries Ltd. , Kelowna , British Columbia.)

A. Production of Clonal Rootstocks and Self-Rooted Trees


The clonal rootstocks so important in modem apple and pear culture are mainly produced
by stooling or layering procedures. However, the growth regulator-assisted rooting of hard-
wood cuttings is growing in importance, a development stemming from problems with the
stool-bed procedures ranging from low productivity to a lack of flexibility in responding to
changing market demands .
Successful techniques for rooting large winter cuttings of various rootstock clones have
been developed by English researchers in the last decade. 1 The specific recipe varies some-
what with cultivar and location, but one essential step is the treatment of the basal end of
the cuttings with an auxin - most commonly 0.25 to 0.50% indoyl-3-butyric acid (IBA)
- to encourage root initiation. Rooting occurs within 2 weeks in a carefully constituted
rooting medium held at 21°C. Ideally, these rooted cuttings can then be handled like rooted
4 Plant Growth Regulating Chemicals

FIGURE 2. Defoliation of Cox's Orange Pippin nursery trees with a 2%


CuEDT A spray applied September 25, 1981. The trees in the left row were
untreated. The photograph was taken October 14 , 1981. (Courtesy of Dr.
J. N. Knight , East Mailing Research Station, Maidstone , Kent , England.)

layers, but more commonly they require additional attention to encourage the development
of a strong root system before being used as rootstocks by the nurseryman. Several important
apple rootstock clones, including M26, M27, MM106, and MMlll, respond well to this
procedure . The quince rootstocks used to control pear tree size and the Old Home x
Farmingdale series of Pyrus communis rootstocks are also readily propagated by hardwood
cuttings .
It is also possible to produce self-rooted trees of a number of commercially important
apple and pear cultivars by the rooting of hardwood cuttings. 2 It has been suggested that
the cost of finished trees can be reduced by this technique, but commercial application to
date has been very limited .
Perhaps the most exciting developments in fruit tree propagation technology are occurring
in the in vitro propagation area. Production of virus-free rootstock materials of apple and
pear and of own-rooted trees of important cultivars of each species is now a commercial
reality in both North America and Europe (Figure 1). This procedure offers great flexibility
in adjusting to market demands since plants of desirable clones can be rapidly multiplied
from mother cultures maintained by the laboratory .
The procedure 3 ·4 is to culture surface-sterilized shoot meristems on a nutrient medium
Volume I 5

1M

FIGURE 3. Feathering of Gloster apple trees following a single spray


of 1000 ppm propyl 1-3-t-butyl phenoxyacetate (M & B 25-105) applied
at approximately 60-cm tree height. The tree on the right was not treated.
(Courtesy of Dr. Henk van Oosten, Research Station for Fruit Growing,
Wilhelminadorp, The Netherlands.)

containing enough 6-benzyladenine (BA) (about 5 X I0- 6 M) to stimulate shoot proliferation


from axillary buds. These shoots are periodically harvested and "planted" on a second
medium lacking BA but containing NAA or IBA to stimulate root initiation. A third transfer
onto a medium free of cytokinins or auxins encourages root development. These small
complete plants are eventually potted and grown into salable trees using standard nursery
practices.

B. Defoliation of Nursery Stock


Finished nursery trees intended for autumn or spring orchard planting, or rootstock layers
intended for sale or further use by the nursery, are dug in the autumn after the plants have
stopped growing and are largely defoliated. However, since nursery stock is encouraged to
grow very vigorously, these physiological processes are delayed. Natural defoliation may
occur too late to make trees available for autumn planting, or in some locations, to permit
digging before the soil freezes.
Therefore, there is great interest among nurserymen in chemical treatments to advance
defoliation although no universally suitable procedure has been developed yet. Most effective
defoliants also cause bark and bud injury from time to time. Furthermore, the results obtained
with a given procedure can differ greatly from year to year, and a number of important
apple and pear cultivars are remarkably resistant to chemical defoliation. Nonetheless, there
are chemical defoliation procedures in commercial use in Europe and North America that
are worthy of mention here.
A I% CuEDT A spray applied in mid-October effectively defoliates nursery trees of Cox's
6 Plant Growth Regulating Chemicals

Orange Pippin apple as well as some other cultivars and is finding acceptance among
nurserymen in England (Figure 2). All apple rootstock clones also respond satisfactorily to
CuEDT A, but certain other apple and pear scion cultivars will sustain bark and bud injury
yet resist defoliation. 5
In North America the combination of a particular wetting agent (Du Pont® WK at about
1.5%) with I00 to 200 ppm (2-chloroethyl)-phosphonic acid (ethephon), applied two or
three times in late autumn, has led to satisfactory defoliation in Washington State. 6 Another
material, 2,3-dihydro-5 ,6-dimethyl-1 ,4-dithiin-1, I ,4,4-tetraoxide (Harvade®, Uniroyal
Chemical Co.), combined with DuPont WK is reported to give comparable results. 7 How-
ever, commercial experience is still quite limited, and variable defoliation and occasional
tree injury continue to be serious concerns.

C. Promotion of Feathering of Nursery Trees


In some fruit growing regions of the world, especially in Europe, orchardists are accus-
tomed to planting trees that have five to ten side branches or "feathers" when they come
from the nursery. Some cultivars branch readily (e.g., the Cox's Orange Pippin shown in
Figure 2), while others branch reluctantly. Feathered nursery trees develop a greater fruiting
volume early in the life of the orchard and are therefore more productive. 8
A growth regulator introduced in the mid-1970s by May and Baker Ltd., n-propyl-3-t-
butylphenoxyacetate marketed as M & B 25-105, is proving to be a highly effective promoter
of feathering in nursery trees (Figure 3). Applied early in the growing season of the final
nursery year at a concentration of about 750 ppm, this material can increase the number of
feathers by threefold or more (Table I). This treatment is even more beneficial if the feathers
developing too low on the trunk are removed by hand. 9
M & 8 25-1 05 is effective on several important apple and pear cultivars, but is considerably
less promising with others. 10 It acts by temporarily interfering with basipetal auxin movement,
thus reducing apical dominance. 11 The feathering effect is achieved most satisfactorily on
vigorously growing nursery trees.
In North America there appears to be greater interest in a treatment to promote branching
of young trees after they are established in the orchard. The growth regulator techniques
being developed for this purpose will be discussed in a later section, but this difference in
attitude toward the production of well-branched nursery trees warrants a brief discussion
here.
Tree planting in Europe occurs in the autumn or very early spring. In the latter case the
trees remain in the nursery over winter or are heeled-in in the orchard in the autumn. Thus,
cold storage of finished trees is less common than is the case in much of North America.
Feathered trees are more difficult and expensive to store than tight bundles of nonbranched
trees. Furthermore, the European climate is generally more conducive to the establishment
of young trees, and less severe pruning is practiced at the time of planting. Thus, feathered
trees are less of a problem for nurserymen and relatively more beneficial to the orchardist.

III. CONTROLLING GROWTH AND MODIFYING THE STRUCTURE OF


ORCHARD TREES

The growth regulator treatments discussed in this section are those applied to orchard
trees, but generally not to fruits or flowers. They are applied to young nonbearing trees to
assist in tree training and to promote spur development; to nonbearing parts of the tree (such
as treatments to control the growth of root suckers); or they are applied to the upper tree
parts during the dormant season.

A. Regulation of Tree Branching Habit


Unfortunately, materials like M & B 25-105 used by nurserymen in Europe to induce
Volume I 7

Table 1
FEATHERING OF NURSERY TREES AS INFLUENCED BY A
SINGLE EARLY SUMMER SPRAY OF M & B 25-105

Number of branches
per tree

M&
B 25·
Untreated 105

Apples on MM106 (850


ppm)
Spartan 3.2 9.3
Crispin (Mutsu) 3.2 9.6
Bramley's Seedling 2.0 10.2
Pears on Quince A (750
ppm)
Conference 0.0 3.5
Williams (Bartlett) 1.3 6.5
Doyenne du Cornice 1.1 3.7

From Quinlan, J. D. , Acta Hortic ., 120, 55, 1981. With permission .

Table 2
STIMULATION OF GROWTH AND BRANCHING OF YOUNG MACSPUR APPLE
TREES TREATED WITH MIXTURES OF 6-BENZYLADENINE (BA), GA 4 + 7 AND
GA 3

Total
No. of new growth
Concentration shoots per Average shoot per tree
No. Treatment (ppm) tree length (em) (em)

I No spray 5.5 7.61 41.9


2 BA 500 12.0 8.28 99 .3
GA. ", 250
3 BA 500 20.8 4.30 89.3
GA" +' 250
GA, 250

From Costante, J. F .. Proc. New Engl. Fruit Meet., 86, 65, 1980. With permission.

feathering of maiden trees are much less effective when applied to trees in the orchard.
Therefore, considering that poorly branched nursery trees are the norm in many countries
and that the value of early branching is being increasingly recognized , treatments to encourage
branching of young orchard trees are beginning to emerge .
With many strong-growing cultivars and strains of apple and pear, not only the number
but also the location and angle of attachment of lateral branches can cause concern . When
a young unbranched tree is planted, the height at which it is headed (cut back) depends to
a large extent upon where the first whorl of lateral branches is desired. Unfortunately, these
branches often emerge from a very narrow zone below the heading cut, with two or more
upright shoots competing for leadership. These branches are often spaced too closely and
are otherwise unsuitable as scaffold branches unless they are mechanically spread.
This unsatisfactory growth habit can be altered by treating dormant buds well below the
heading cut with a mixture ofbenzyladenine (BA) and GA 4 + 7 in a lanolin paste. 12 Treatments
QC
v'
/
,,_ , _,-
". ""~.....
-~ :' ·~~ .._
'~ ......-. ~,. -- .-._ ,..- " - ..... J ~· --
-~ .... - ..,
iS"
~
CJ
....
<:::>
~
::-
~
~
:0:::
""
iS"
§·
~

Q
::!
""
§'
1:;"

FIGURE 4. Modification of growth and fruiting habit of Cox's Orange Pippin apple trees with annual daminozide spray treatments. Both trees were planted in
November 1970 and photographed late in the summer of 1973. The tree on the right was treated with 1700 ppm daminozide in 1971 and 1972 and 1000 ppm in
1973. (From East Mailing Research Station Report for 1973. 64, 1974. With permission.)
Volume I 9

applied to individual buds or to all of the buds and bark in the desired trunk zone give
comparable results. The result is more branches , wider branch angles, and increased total
shoot growth. However, to date there has been very little use of this promising technique
in commercial orcharding . Correct timing is important and may be difficult to achieve ; or
perhaps the treatment method is considered too costly and tedious.
Quite a different tree branching problem is encountered with some of the increasingly
popular "spur type" cultivars of apple. These compact trees tend to form fruiting spurs
rather than branches, particularly when tree vigor is low. It is not uncommon for trees of
these cultivars to resemble a "fruiting pole", thus making very inefficient use of the allotted
orchard space.
An early season application of Promalin® (Abbott Laboratories Inc.) , a proprietary mixture
of BA and gibberellins A4 and A 7 (GA 4 + 7 ), is showing considerable promise in tests in
North America and Belgium. It stimulates the development of lateral branches and increases
the total length of shoot growth on treated trees (Table 2). The addition of a third gibberellin
(GA 1 ) may further enhance treatment effectiveness. 13

B. Promotion of Spur Development and Flower Initiation


Several important apple cultivars (e.g., Rome Beauty, Cox 's Orange Pippin, Tydeman's
Early Worcester) form long thin branches with a paucity of fruiting spurs. This condition
can be partly corrected, and long-term tree productivity improved, by annual applications
of butanedioic acid mono-(2,2-dimethylhydrazide) (daminozide or Alar-85®, Uniroyal Inc.) .
A program of 500 to I 000 ppm daminozide applied for several years within a month of bud
break leads to shorter, sturdier branches supporting more spurs per unit of branch length
(Figure 4). Flower initiation is also increased by these treatments but the more important
aim is to modify growth habit.
Other apple cultivars, while displaying a suitable compact and potentially productive tree
shape and growth habit, are nonetheless reluctant to commence full bearing. The growth
regulator treatment prescribed in Washington State and elsewhere in the U.S. is a combination
of daminozide and ethephon applied 4 to 5 weeks after bud break. Typical rates are 1000
to 1500 ppm daminozide and 300 ppm ethephon in a high volume spray. When the flowering
problem is less severe, daminozide alone (applied somewhat earlier) is the more common
treatment. These are likely to be one-time treatments occurring in the fourth or fifth orchard
year.
A comparable reluctance to commence flowering and fruiting occurs in young pear trees.
A chemical registered in several European countries to control vegetative growth and improve
flowering of pear trees is (2-chloroethyl) trimethylammonium chloride (chlormequat, Cy-
coceJ®, Cyanamid Inc.). The Dutch recommendation for controlling growth and improving
fruitfulness of Cornice pear is to apply I 000 to 1600 ppm chlormequat when the new shoots
have five to eight leaves and repeat in 2 to 3 weeks if necessary .

C. Suppression of Root Suckers


A troublesome characteristic of some otherwise desirable apple and pear rootstocks is
their tendency to initiate strong growing shoots from adventitious buds near the soil surface
(Figure 5). Root suckers are unproductive and unsightly ; they hinder access to the tree and
can harbor a wide range of orchard pests. Fire blight entry through root suckers is a particularly
serious concern in some countries.
Procedures for the chemical control of root suckers are slowly being developed. A product
originating in England and marketed in western Europe and the Southern Hemisphere com-
bines 12.7% by weight of the free acid of NAA in an emulsion with decanol. Tipoff® (Midox
Ltd., U.K.) is applied at a rate of about 4% in water when the suckers are about 10-cm
long (Figure 5) . At least 4 weeks should have elapsed since full bloom and spray drift must
be avoided.
10 Plant Growth Regulating Chemicals

FIGURE 5. Control of sprout and sucker growth with NAA-based products. Top: Control of sprout growth on
the trunks of 15-year-old Delicious apple trees with Trehold Inhibitor A-112® in 20% white latex paint applied in
April. The active ingredient is the NAA ester at 10000 ppm. (Courtesy of Dr. T. J. Raese, USDA Tree Fruit
Research Laboratory, Wenatchee, WA). Bottom: Control of root suckers on 14-year-old Beurre Hardy/Quince A
pear trees with 4'7< Tipoff'"' applied in early summer. The active ingredients are alpha naphthaleneacetic acid (2240
ppm) and N-decanol (2.78%). (Courtesy of Dr. S. J. Wertheim, Research Station for Fruit Growing, Wilhelmi-
nadorp, The Netherlands). The treated tree (both top and bottom) is on the right.

In North America the material more commonly used for this purpose is Trehold Sprout
Inhibitor A-112® (Union Carbide Inc.). This product contains 13.2% of the ethyl ester of
NAA and is diluted to a concentration of 0.5 to 1.0% NAA with water or white latex paint,
depending on use. For root sucker control it is usually diluted with water and used as
described for Tipoff®. With both materials, best results are obtained when the sucker growth
from previous seasons is cut back so that only current season growth is treated.

D. Suppression of Water Sprouts


Water sprouts are strong upright-growing shoots arising from the trunk and major scaffold
Volume I 11

limbs. They are particularly prevalent in vigorous trees subjected to containment pruning.
On less vigorous trees they may be confined to regions near major pruning cuts. Water
sprouts interfere with spray movement and coverage within the tree and seriously reduce
light penetration. Removing them by hand significantly increases annual pruning costs.
Trehold Sprout Inhibitor A-I J2® diluted with 25 to 50% white latex paint in water to a
final NAA concentration of 0.5 to 1.0% is used commercially in North America and parts
of Europe to control water sprouts. It is applied as a paint to problem areas on the trunk
and scaffold limbs. This may involve a complete coverage or spot applications to major
pruning cuts . NAA applied to uncut surfaces is not translocated so thorough coverage is
essential. However, applied to cut surfaces it will move about 30 em into the tree and towards
the root system , concentrating in the outer ring of xylem . 14 This NAA paint treatment is
about equally effective on apple and pear trees. 15

IV. REGULATION OF FLOWERING AND FRUIT SET

Apple and pear growers also use chemicals to regulate cropping by increasing "return"
flowering of cultivars tending to bear biennially; by promoting fruit set on certain problem
cultivars; and perhaps most importantly, by reducing set on trees that tend to overcrop in
any given season. Interestingly, the use of one practice does not necessarily preclude the
use of either of the others on the same tree and even in the same season.

A. Improving Return Bloom


Early fruit thinning is a powerful tool for increasing return flowering. However, the
growth-retarding chemicals, daminozide, ethephon, and chlormequat, when applied during
the flower initiation period provide additional help in this regard. Daminozide is the favored
material for bearing apple trees because, unlike ethephon, early season use does not involve
the risk of excessive fruit abscission. On cultivars tending to flower biennially, daminozide
is often applied with one of the fruit thinning sprays. However, in England and parts of
Europe a slightly different use pattern is developing . Annual mid-summer daminozide sprays ,
at considerably reduced rates , are said to improve flower strength or "quality" and thus
improve cropping the following year.
With pears, increasing numbers of European growers are using multiple applications of
chlormequat to improve the productivity of mature trees . Relatively low rates of chemical
(generally less than 500 ppm) are applied several times at about 2-week intervals commencing
shortly after petal fall. This treatment regime effectively controls shoot growth and improves
return cropping without any appreciable reduction in fruit size. Increases in both flower
numbers and fruit set explain the improved cropping. 16

B. Increasing Fruit Set


Spring frosts and other climatic hazards periodically reduce the set of pome fruits in all
major producing regions . This, coupled with the fact that growth regulator techniques have
been developed to improve fruit set on crops as diverse as tomatoes and grapes, has en-
couraged the search for fruit setting aids applicable to apples and pears. Unfortunately,
progress to date, especially with apples, has been rather disappointing with cultivars differing
widely in their responsiveness to the various chemicals. Furthermore, treatments developed
specifically for a given cultivar and region all too frequently fail to achieve the desired result
in wider commercial practice .
Probably the most successful practice now in commercial use aims at improving the
cropping of pears with blossom-time sprays of GA 3 or GA 4 + 7 . 17 Rates of 15 to 25 ppm
in a high volume spray are commonly suggested . Cultivars which normally set a high
proportion of seedless fruit, such as Triomph de Vienne and Beurn! Hardy, respond partie-
12 Plant Growth Regulating Chemicals

ularly well to gibberellin treatments and are treated annually by many orchardists in England ,
Holland , and Belgium. Some other cultivars (e.g . , Conference) are treated only when spring
frost has been severe enough to prevent the natural production of seeded fruits. This rescue
operation can be highly cost effective even though some fruits may be misshapen, and
blossom density the following season is likely to be reduced.
A second method for increasing fruit set of pears involves the use of 2,4,5-trichloro-
phenoxypropionic acid (fenoprop) as a postharvest (5 to 10 ppm) or a blossom-time (3 to
6 ppm) spray. This spray is safely applied to a small number of cultivars and dates back to
early work in Washington State with Anjou pears. 18 This use of fenoprop no longer appears
in North American spray guides , perhaps because positive responses often reveal other
cultural ills , such as inadequate pollination , but it continues to find favor in some other
countries. South African and Australian growers of Packhams Triumph pears continue to
use fenoprop in this manner.
Chemical promotion of apple fruit set is even more problematic. The severity of the fruit
set problem with Cox's Orange Pippin apples has led a group of English researchers to seek
a growth regulator solution. 19 · 20 The present state-of-the-art fruit setting mixture for Cox
combines GA 4 + 7 , N 1N 1-diphenylurea, and 2-naphthoxyacetic acid. Applied at blossom
time, it has increased the set of seedless fruits in a number of experiments. However, seedless
fruits, or those with very few seeds, compete poorly with normally seeded fruits and the
aim of increasing yields year in and year out by setting a proportion of such fruits is proving
difficult to achieve. The key to success may be to grow Cox without cross-pollination and
rely completely on hormone-induced set. This novel approach is now under test.
Clearly, the development of chemical techniques to reliably improve apple and pear fruit
set is an important area for continued research. Early results with the ethylene biosynthesis
inhibitor, aminoethoxyvinylglycine, have been outstanding in this regard. 21 This use of an
inhibitory rather than promotive growth substance could be the conceptual breakthrough that
will lead to some long-awaited practical solutions.

C. Chemical Thinning
Each year it becomes more difficult to sustain apple and pear production where fruit
thinning is done entirely by hand. Hand thinning is costly , and because of labor availability
problems, it is increasingly difficult to achieve at any price. However, chemical thinning is
more than simply a cost-effective way of removing excess fruit. Its major advantage over
hand thinning is that fruit removal is accomplished early in the growing season when the
return bloom and fruit sizing benefits are greatest.
With the exception of the more recent introduction of ethephon, the chemicals now in
commercial use for flower and fruit thinning have been widely available since the 1950s.
Not surprisingly, some very localized usage patterns have evolved during these years of
field experience. Table 3 illustrates the range of chemical thinning advice directed at growers
of a single widely grown apple cultivar, Golden Delicious, in a number of countries and
regions within countries . Unfortunately, a more complete presentation of the latest published
advice concerning chemical thinning of apples and pears cannot be attempted here . The
sheer volume of information indicates the importance placed on chemical thinning technology
around the world and suggests an interest in, if not an absolute requirement for, a great deal
of local advice.
However, some important regional differences are worthy of elaboration. For example,
blossom thinning with sodium 4,6-dinitro-ortho-cresylate (DNOC) is an integral part of the
apple thinning programs used by growers in western North America, yet its use is rare in
other regions. Some interesting exceptions include Victoria, Australia, and Nova Scotia,
Canada. Of the postbloom thinners, NAA is applied at petal fall to a range of apple cultivars
in Queensland and Western Australia, although the common advice elsewhere is for a much
Volume I 13

Table 3
CHEMICAL THINNING OF GOLDEN DELICIOUS APPLES: "SPRAY
CALENDAR" ADVICE FROM 12 LOCATIONS

Concentration
Location Chemical Timing (g/100 t)• Remarks

Canada
British Columbia DNOC Full 40 For "concentrate" spraying apply
bloom 4. 7 kg/ha in 1700 {; of water
NAAm 8-12 days 1.7 Use a wetting agent; for "concen-
AFB' trate " spraying apply 118 g/ha as
above
NAA plus 15 days AFB 0.5 Apply as a full volume spray
carbaryl 50
U.S.
Washington DNOC See remarks 25-38 Apply when 3 blossoms per spur
cluster are open on the N side of
the tree
NAAm 7-14 days 1.7-3.4 Add a wetting agent
AFB
NAAm plus I 0---20 days 1.7-3.4 Ethephon enhances thinning and re-
ethephon AFB 30---40 tum flowering
NAA plus 15-25 days 0.3
carbaryl AFB 45---60
Ohio NAA 14--21 days 1.7-2.0 Reduce to 0.8 g if a wetting agent is
AFB added; NAAm not recommended
New York NAA Up to 21 days 1.5-2.0 Without a wetting agent
AFB or 18-
mm fruit
length
NAA plus As above 0.5-1.0
carbaryl 60
Virginia NAA 14--18 days 0.8 Add a wetting agent; carbary I use
AFB can lead to russeting
England Carbaryl 80% petal fall 50 Wet the tree thoroughly and apply
to 21 days under slow drying conditions
AFB
The Netherlands NAAm 7 days AFB 6.0 Without wetting agent; apply as a
full volume spray
Carbaryl 12-mm fruit di- 75 As above
ameter; 19-
28 days AFB
France NAAm 7-10 days 5.0 Apply at least 1500 £/ha
AFB
NAA 9-mm fruit 1.7 As above
diameter
Carbaryl 15- to 16-mm 150 Apply 1500---2000 t/ha
fruit diameter;
18-25 days
AFB
West Germany NAAm 75% petal fall 5.0---5.8 NAA is not available for use in
to 7 days AFB Germany
South Africa NAAm Full bloom to 7.0 Reported to thin less than carbaryl
90% petal fall
Carbaryl 14--24 days 75 May aggravate russeting
AFB
14 Plant Growth Regulating Chemicals

Table 3 (continued)
CHEMICAL THINNING OF GOLDEN DELICIOUS APPLES: "SPRAY
CALENDAR" ADVICE FROM 12 LOCATIONS

Concentration
Location Chemical Timing (g/100 £)• Remarks

Australia
Tasmania NAA Full bloom to 1.0 Add a wetting agent
14 days AFB
Western Australia NAA Late petal fall 1.0 Add a wetting agent; carbaryl aggra-
vates russeting

" Values are grams of active chemical. These values multiplied by 10 yield ppm.
" After full bloom.

later spray. North Carolina growers are specifically advised that early NAA sprays may
stick the fruit on rather than remove it. And in British Columbia, Canada successful thinning
is achieved using low volume or "concentrate" spray equipment, whereas only 100 km
away in Washington State the advice is to apply all chemical thinning sprays to run-off.
Some of these procedural differences relate direct! y to climate. Others relate to the orchard
management systems used in various regions. Furthermore, it has been demonstrated re-
peatedly that cultivars of both species differ greatly in their responsiveness to various chemical
thinning agents.
Undoubtedly, other explanations for this diversity could be listed, but suffice it to say
that widely applicable recommendations for chemical thinning are seldom attempted. The
following paragraphs describe in general terms the more important procedures.

I. Blossom Thinning of Apples


This is predicated on the following general observations:

1. Apple trees normally produce 5 to 20 times more blossoms than are required to produce
a full crop.
2. Individual fruiting spurs produce 5 or 6 flowers yet should support no more than 2
fruits.
3. If a tree is to maintain annual cropping, a sizable proportion of these spurs should
"rest" in any given season.
4. The flowers in each cluster open predictably, i.e., the central flower opens first.
5. With certain cultivars the position of the fruit within this cluster influences an important
aspect of fruit quality, i.e., fruit shape.

Selective blossom thinning can be achieved with DNOC since: (I) fertilized flowers (fruits)
are resistant to DNOC because the stigmatic tissues have achieved their function; (2) uno-
pened flowers are protected from the spray; and (3) open but unfertilized flowers are damaged
and therefore fail to set.
Obviously, timing of the spray is very important. Factors to consider include the stage
of bloom and the conditions for pollen transfer and pollen tube growth that existed during
the 24- to 48-hr period preceding the spray. If the spray is too late, very few flowers will
be affected. If it is too early, the "king" or central flower may be thinned but too many
lateral flowers will persist. This would be unwise for Delicious apples where the elongated
"king-bloom" fruits are valued, but may be the correct procedure for Mcintosh where this
fruit is often misshapen. Weather conditions at the time of, and following spraying are also
important in that wet weather extends the activity of DNOC. It is not surprising that DNOC
is not widely used in maritime fruit growing regions.
Volume I IS

Table 4
THINNING OF PEAR FRUITS WITH NAA AND NAAm: "SPRAY
CALENDAR" ADVICE FROM 8 LOCATIONS

Concentration
Location Cultivar(s) Chemical (ppm)• Timing"

Canada
British Columbia Bartlett NAAm 16 13-21
Ontario Bartlett and NAAm or 10 2-3
Anjou NAA 10 5-9
Keiffer and NAAm and 15-20 2-3
Winter Nelis NAA 10 5-9
U.S.
New Jersey Bartlett and Bose NAAm 25-50' 7-10
New York Bartlett NAAm or 25 3-5
NAA 2 5-10
Ohio Bartlett NAAm 25-35 3-7
Washington Bartlett NAAm or lG--15 15-21
NAA lG--15 15-21
Denmark Conference and NAA 15' 3-10
Bonne Louise
Clapps Favorite NAA 22' 3-10
Australia
New South Williams NAA lG--15 lG--14
Wales

Concentration of chemical in a full volume spray with added wetting agent.


• Days after full bloom.
Wetting agent requirement not clearly specified.

Some suggested rates and times for DNOC application to thin Golden Delicious are listed
in Table 3. Short-season cultivars and those prone to biennial bearing are most commonly
treated with DNOC. Evidence to date suggests that pear trees cannot be safely thinned with
this chemical.

2. Postbloom Thinning with Hormone Materials


Postbloom thinning with hormone materials, primarily NAA and naphthaleneacetamide
(NAAm), is the most widely used approach to chemical fruit removal. These chemicals are
sold under numerous trade names. NAA has greater auxinic activity than NAAm and this
is usually reflected in lower application rates. Because of its attenuated auxinic activity
NAAm is less likely to overthin, but because it appears to retain its auxinic activity for a
longer period, NAAm usage can lead to some unexpected side effects. The thinning activity
of both NAA and NAAm is usually enhanced when they are applied with wetting agents.
The mode of action of NAA and NAAm in inducing selective fruit abscission is far from
clear. One would expect the same mode of action yet, as was suggested above, certain
cultivars respond differently (e.g., NAAm is more likely than NAA to lead to "pygmy"
Delicious apples). NAAm is typically, but not always, applied somewhat nearer blossom
time than is NAA.
Most cultivars of apples appear to be responsive to NAA and NAAm as fruit thinning
agents commencing at petal fall and continuing for 2 or 3 weeks. 22 Early sprays are suggested
in some regions; later ones in others. Rather typical advice for thinning apples with NAA
and NAAm on the eastern seaboard of North America is found in the 1981 Virginia Spray
Bulletin for Commercial Tree Fruit Growers. For Grimes, Jonathan, Stayman, York Im-
perial, and Rome Beauty apples this bulletin suggests 50 ppm NAAm applied between 4
16 Plant Growth Regulating Chemicals

and 18 days after full bloom or 5 ppm NAA plus a wetting agent applied between 14 and
18 days. It suggests that early NAAm sprays (between 4 and 8 days) are particularly
appropriate for short-season apples such as Yellow Transparent, but can also be used for
the above-mentioned cultivars. NAAm is not recommended for Delicious or Golden Delicious
and NAA is not recommended for the summer apples. The comparable bulletin from the
neighboring state of North Carolina warns growers that NAA at rates higher than 5 ppm
can lead to the retention of pygmy Delicious fruits.
Growers in New Jersey are advised that NAAm levels can be reduced to 25 ppm if Tween
20 is added to the spray mixture. This bulletin, 1981 Commercial Tree Fruit Production
Recommendations for New Jersey, also suggests that Golden Delicious apples can be thinned
with 8 to 10 ppm NAA (plus Tween 20) applied 14 to 16 days after full bloom. This material
is also suggested for petal-fall sprays for the early maturing apples and in the 7- to 14-day
time period for mid-season apples like Mcintosh, Spartan, and Empire.
Both NAA and NAAm are used to thin Bartlett or Williams pears in North America and
parts of Australia and a number of other cultivars are similarly thinned in Europe and
elsewhere (Table 4). And, as has been shown for apples, the specific advice concerning
timing and application rates can vary rather widely, particularly within North America. It
is interesting, for example, that in Washington State and Ontario, Canada NAA and NAAm
are applied at the same rates (10 to 15 ppm), whereas in New York the suggested NAA rate
is lower by a factor of I 0. The recommended time of application of each of these chemicals
ranges from petal fall in several eastern states to 15 to 21 days after full bloom in the west.
Generally speaking, the use of chemicals to thin pear fruits is less successful and less
common than with apples. In some very important pear growing regions such as California
and Victoria, Australia chemical thinning is either considered unnecessary or not reliable
enough to recommend.

3. Postbloom Thinning of Apples with Carbaryl


Postbloom thinning of apples with carbaryl (!-naphthyl N-methylcarbamate), and more
recently with carbaryl-containing combinations, is practiced worldwide. It dates back to the
observation by Batjer and Westwood 23 that carbaryl, applied as an insecticide, reliably
reduces fruit set. Advantages of carbaryl include the fact that it seldom causes overthinning
yet is active well into the postbloom period. It is widely used on the Delicious cultivar
because, unlike NAAm, it does not encourage the retention of pygmy apples. The insecticidal
activity of carbaryl is a disadvantage when it leads to the destruction of beneficial insects
and mites. However, careful monitoring of insect and mite populations can be used to avoid
problems. For example, the excessive reduction of predaceous mite populations can lead to
severe problems with spider mites, but mites resistant to carbaryl are becoming increasingly
common. Consequently, this problem occurs less frequently.
The mechanism by which carbaryl causes fruit thinning, while not completely understood,
appears to differ from that of NAA and NAAm. 22 ·24 Therefore, more effective and reliable
thinning can be achieved by combining carbaryl with NAA (Table 5). This approach is
rapidly gaining popularity.
Carbaryl is not a satisfactory pear fruit thinner.

4. Flower and Fruit Thinning with Ethephon


Applied as a blossom-time or postbloom spray, ethephon can be used to induce flower
and fruit abscission. However, despite early promise as a thinning agent, ethephon is not
widely used for this purpose in commercial fruit growing. It has proven more likely to
overthin than the alternative materials, a problem that researchers hope to overcome by
refining the procedures for its use. One of the very few published suggestions for its use as
a fruit thinner in commercial orcharding appears in the 1981 Spray Guide for Tree Fruits in
Volume I 17

Table 5
CHEMICAL THINNING OF
SPARTAN APPLES WITH
CARBARYL AND/OR NAA,
SUMMERLAND, BRITISH
COLUMBIA 1979 TO 1981

Final fruit set as a


percent
of the water controls"

Treatment• 1979 1980 1981

Carbaryl (500 ppm) 86 95 74


NAA (10 ppm) 71 84 63
Carbaryl plus NAA 50 69 51

Sprays applied IS days after full bloom each year


in the same orchard.
• The control fruit set values (fruit per 100 blossom
clusters) were 88.9, 129.2, and 199.0 in 1979,
1980, and 1981, respectively.

Eastern Washington. A combination of NAAm (17 to 34 ppm) and ethephon (300 to 400
ppm) applied 10 to 20 days after full bloom is suggested for Golden Delicious to help
counteract a biennial bearing tendency.

V. IMPROVING FRUIT CONDITION AND APPEARANCE

Modifying fruit condition and appearance is a relatively new but steadily expanding area
of growth regulator usage in apple and pear production. Growth regulators can be used to
improve the firmness, color, and shape of apples and to decrease the incidence or severity
of several disorders of apple and pear fruits. Daminozide, introduced in the mid-1960s, is
responsible for many of these effects, but some imaginative new uses for gibberellins and
cytokinins are gaining in importance.

A. The Effects of Daminozide on Apple Fruits


Daminozide is now available in most fruit growing countries and, as will become in-
creasingly apparent, is proving to be a very useful chemical. Its effects on fruit color and
condition are still generally viewed as welcome side effects of sprays applied for more urgent
reasons such as fruit drop control or the promotion of flowering. However, it is increasingly
common for apple growers to apply daminozide in early to mid-summer with the primary
aim of improving red color, increasing fruit firmness, or perhaps protecting the crop from
a specific disorder such as "watercore" (Table 6). 25 - 33
Daminozide is usually applied as a full volume spray containing 500 to 2000 ppm of
active chemical. Low volume sprays have also proven effective in many regions. The amount
of chemical applied is usually less with mature trees and is commonly reduced after several
years of use on the same tree.
Some adverse carry-over effects of daminozide on fruit size, shape, and length of the
pedicel have been noted and these are more likely to occur if the chemical is applied at high
rates or too late in the growing season. Apple cultivars differ considerably in their suscep-
tibility to these carry-over effects.
Another precaution is that a few storage disorders of apples can be advanced or intensified
by daminozide treatment. These effects are often specific for a given cultivar and location,
18 Plant Growth Regulating Chemicals

Table 6
THE EFFECT OF DAMINOZIDE TREATMENT ON
SOME APPEARANCE AND CONDITION
PARAMETERS OF MCINTOSH AND DELICIOUS
APPLES

Control Treated Ref.

Mcintosh
Surface red color(%) 63 75' 25
37 62 26
62 80 27
Flesh firmness (kg) at harvest 8.1 8.7 28
6.9 7.5 25
Flesh firmness (kg) after 3--4 5.3 5.9 28
months of 0°C storage 7.9 7.7 n.s. 29
Flesh firmness (kg) after 5 months 4.3 4.9 29
of CA storage 4.5 5.7 25
Titrateable acidity (mt 0.1 N 34.3 25.6 28
NaOH/25 mt of juice)
Juice soluble solids(%) 11.8 11.6 n.s. 28
Development of brown core in 0°C 30 53 30
storage (% afflicted)
Delicious
Flesh firmness (kg) at harvest 7.8 8.6 31
8.7 9.1 28
8.0 8.5 32
Flesh firmness (kg) after 3--{i 6.7 8.4 33
months of 0°C storage 7.3 7.5 n.s. 28
Titrateable acidity (mt 0.1 N 11.4 11.4 n.s. 28
NaOH/25 me of juice)
Juice soluble solids (%) 12.6 12.0 31
II. I 10.9 n.s. 28
Incidence of water core at harvest 56 8 31
(% afflicted)
Incidence of storage scald (% 84 20 33
afflicted)

Differences due to treatment are statistically significant at the 5% prob-


ability level unless indicated by n.s.

but nonetheless can be of serious concern. For example, daminozide sprays are reported to
shorten the storage life of Mcintosh apples in New York because they promote internal
browning. 30 This effect has not been observed with Mcintosh grown in British Columbia.

B. Preventing Premature Softening of Pear Fruits


A disorder of Bartlett pear fruits referred to as "pink-end" can be avoided or reduced by
a daminozide spray (about 1000 ppm) applied 18 to 24 days before anticipated harvest.
Pink-end is characterized by softening of the calyx end of fruits still on the tree and is
induced by field temperatures below 10°C during the month preceding harvest. 34 This use
of daminozide is presently confined to Oregon and Washington.

C. Reduction of Fruit Russeting


Most apple cultivars are susceptible to skin russeting when grown in humid climates.
Severe fruit russeting frequently reduces the value of Golden Delicious apples, and an
effective protectant would receive widespread acclaim. Australian work indicating that GA4 + 7
sprays reduce the severity of russet 35 is very promising and is being followed up at other
Volume I 19

Table 7
REDUCTION OF APPLE FRUIT RUSSETING BY 4
SPRAYS OF GA 4 + 7 APPLIED WITHIN 1 MONTH OF
BLOOM

Orchard: 2 3 4 5 6
Cultivar: Karmijn de Sonnaville Golden Delicious
No spray 82" 50 79 58 18 40
GA4 + 7 (10 ppm) 41 10 68 42 4 17

Values are the percentage of moderately to severely russeted fruits in


each treatment and orchard lot.

From Tromp, J. and Wertheim, S. J., Proc. 15th Colloq. Int. Potash lnst.,
p. 137, 1980. With permission.

locations. 17 •36 Relatively high concentrations of GA 4 + 7 and multiple applications appear to


be necessary for complete control, 36 but considerably lower rates can lead to a marked
improvement in fruit finish (Table 7). 17 Assuming no adverse side effects, eventual regis-
tration and widespread commercial use can be anticipated. Interestingly, GA 3 is considerably
less effective and very recent evidence suggests that it is the GA 4 in the GA4 + 7 mixture that
reduces russeting. And since GA 7 is the mixture component most likely to inhibit flower
initiation, 38 a pure GA4 product for russet control would be ideal.

D. Increasing the Length to Diameter Ratio of Apples


A proprietary mixture of GA4 + 7 and 6-benzyladenine (6-BA) (Promalin®, Abbott Lab-
oratories) is now registered in several countries and is used primarily to promote fruit
elongation of Delicious apples. Historically, elongated or angular fruit of this cultivar have
received a premium in the marketplace, and it has been clearly demonstrated that blossom-
time sprays of Promalin® can lead to longer, more "typey" fruit. 39 It is also suggested that
Promalin® treatment increases average fruit weight without noticeably reducing fruit num-
bers, However, other reports show quite clearly that this treatment enhances fruit thinning; 40
in fact, this effect is sometimes viewed as being beneficial and can be enhanced by adding
DNOC to the spray mix. 41
The time of Promalin® application has a great influence on the fruit elongation effect. 39 ,4 1 .4 2
It is presently suggested that the spray be applied at the 80% bloom stage and that, unless
fruit thinning is desired, high volume sprays and rainfall near the time of application are to
be avoided. A medium volume spray containing 25 ppm of the active ingredients in Pro malin®
(approximately equal amounts by weight of 6-BA and GA4 + 7 ) is used by Washington State
apple growers.

VI. PREVENTING PREHARVEST FRUITDROP

A major production problem with a number of important apple and pear cultivars is the
tendency for fruits to loosen and drop before harvest can be completed. It is not uncommon
for crop losses of 20 to 30% to be experienced unless some form of chemical protection is
provided.
One of the earliest nonherbicidal uses of plant growth regulators was the use of synthetic
auxins to control preharvest drop of apples and pears and these uses continue today (see
Edgerton43 for a review of this subject), More recently, daminozide has proven to be effective
in this regard and, while more costly than the auxinic materials, is gaining in popularity.

A. Control of Fruit Drop with Synthetic Auxins


NAA and 2,4,5-trichlorophenoxypropionic acid (2,4,5-TP or fenoprop) are the synthetic
20 Plant Growth Regulating Chemicals

Table 8
SOME SELECTED "SPRAY CALENDAR" ADVICE FOR CONTROLLING
PREHARVEST DROP OF APPLES AND PEARS WITH SYNTHETIC AUXINS

Location Crop Material Specific advice

Canada
British Columbia Bartlett pear NAA 20 ppm in a high volume spray or 1.4 kg
Early- to mid-season of a 6.34% active NAA product per hec-
apples tare; apply 2 days before expected drop
Ontario Apples NAA 10 ppm applied before drop commences;
repeat in 7 days if necessary
England Conference pear NAA 10 ppm in a high volume spray
Early apples Fenoprop 10--15 ppm in a high volume spray
Other apples NAA I 0 ppm 6-14 days before expected drop
The Netherlands Apples and pears NAA 5-7.5 ppm for early apples and pears-
10 ppm for all others; apply under fast-
drying conditions to cultivars susceptible
to skin damage (e.g., Boskoop); also, ap-
ply as a high volume spray 7-10 days
before expected harvest
South Africa Apples and pears NAA 20 ppm in a high volume spray at first sign
(all cultivars) of drop; normally 10--14 days before
harvest
Delicious and Granny Fenoprop I 0 ppm as above
Smith apples
Israel Apples Fenoprop 10-20 ppm depending on cultivar; NAA
added when quick action is desired
Japan Japanese pear 2,4-DB' 10--15 ppm 7-14 days before
(all cultivars) plus harvest; 30--40 ppm just before harvest
MCPBh
U.S.
North Carolina Delicious apple Fenoprop 10 ppm in a high volume spray 10--14
days before peak harvest date
Delicious and other NAA I0 ppm 4--6 days before expected fruit
apples drop
Ohio Early apples NAA 5 ppm 4---5 days before heavy drop
Apples later than NAA I0 ppm as above; repeat in 7 days if
Mcintosh necessary
Delicious and Grimes Fenoprop 10 ppm in a high volume spray 10 days
apples before peak harvest
Late apples Fenoprop 20 ppm as above
Washington Bartlett pear NAA 10 ppm in a high volume spray or 0.46-
0.94 f of a 5.6% active NAA product per
hectare in a low volume spray
Delicious and Winesap Fenoprop 10 ppm in a high volume spray 7-12 days
apples before expected drop; concentrate spray-
ing can lead to blind wood
Delicious apple NAA 15-20 ppm in a high volume spray or 218
g actual NAA per hectare applied by
aircraft

2,4-Dichlorophenoxybutyric acid.
2-Methyl-4-chlorophenoxybutyric acid.

auxins used most widely for this purpose (Table 8). Important advantages of these materials
are their low cost and quick action. Relatively low levels of chemical (10 to 20 ppm),
applied just before fruit drop commences, will delay abscission for several weeks. NAA
provides fast but relatively short-lived protection whereas fenoprop, thought to be somewhat
slower acting, gives excellent fruit drop control for up to 1 month.
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