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Plant Growth
Regulating Chemicals
Volume I1
Editor
Louis G. Nickell, Ph.D.

Vice President
Research and Development
Velsicol Chemical Corporation
Chicago, Illinois
Boca Raton London New York

CRC Press, Inc.
CRC Press is an imprint of the
Boca Raton,
Taylor & Francis Group, Florida
an informa business
First published 1983 by CRC Press
Taylor & Francis Group
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Boca Raton, FL 33487-2742
Reissued 2018 by CRC Press
© 1983 by CRC Press, Inc.

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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)
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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., ViGe 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 CHEMRAWN 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
Unit
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 Physiologi st 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
U.S. Department of Agriculture W. C. Wilson
Pasadena, California Research Scientist Ill
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 I
Growth Regulator Usage in Apple and Pear Production . . .. . .. . .... . . . . . . . .... . .... ...... 1
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) . .... . ... . . .. ..... .. 4 1
Kenneth Bridge
Chapter 4
Bioregulation of Rubber Synthesis in Guayufe 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. Jung and W. Rademacher
Index .. . . .. .. . . .. . .. .... . .... .... . ..... ..... . . .. . .. . . .. . . .. . . . ... . .. . . . . ... . ..... .. . . . .. 273
Volume II
Chapter 1
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
Gro wth 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 II 1
Chapter 1
TROPICAL FRUIT AND BEVERAGE CROPS
D. P. Bartholomew and R. A. Criley
TABLE OF CONTENTS
I. Introduction .... ... .... . ....... . ..... . ..... ...... . . ....... . ... ... .... . . ... ........ 2
II . Ananas comosus (L.) Merr. (Pineapple) ....................... . ......... . ........ 3

A. Propagation .... ... .............. . .............. .. ..... ... ...... .. .... ... .. 3
B. Vegetative Growth . .. ...... .... .... .......... ...... ... .. .. ..... .. ......... 4
C. Induction of Flowering (Forcing) ... .... ...... ............ .. ...... ..... ... 4
1. History of Growth Regulator-Induced Flowering . .... .... .... ... .. 4
2. Absorption and Mechanism of Action ... .... .... .. . .. ... ..... .... . 4
3. Plant Susceptibility to Forcing ....... ... ..................... .... . 7
4. Effects of Forcing on Fruit and Propagule Production ..... . . . ..... 8
5. Forcing as a Commercial Practice ... . . ... . ...... .... .. ..... ... .. .. 8
D. Fruit Development .................................... .. .................. 9
E. Fruit Ripening .. .. ... .. . ... .. . ........... .. ...... .... .. . ... ..... ...... . . . 10
F. Postharvest Treatments ........... . ...... . ....... . ... ... ...... .. ..... .... 11
G. Future Research .. .. ... . .. ..... . .................... . ... ..... ... ..... ... . 11
III. Carica papaya L. (Papaya) ................... .. ..... .. ...... .... .... .. .......... 11
A. Propagation . . . ...................... . ..... .. ..... ... ..... .... .... .. ..... . 11
B. Vegetative Growth ....................................................... 12
C. Flowering ... . ...... .. ....... .. .... ... ...... . ...... . ..... .... ... .......... 12
D. Fruit Set and Development .. .................... .. ...................... 12
E. Fruit Ripening ................................ . .......................... 12
IV. Coffea arabica L., C. canephora L. (Coffee) .. ......... .. ...................... 13

A. Propagation ... . .. .. ... .... .... .... .... .. .. .. . .... ... . .......... . ......... 13
B. Vegetative Growth . . . .......... .... .. .. ... ...... .. . . ... .. . . . .... .. . ... .. . 13
C. Flowering . ..... .. ...... ... ..... , ...... ... ...... . . .. ...... . ...... . ..... ... 13
D. Fruit Set and Development ... .. ......................................... 14
E. Fruit Ripening ........................................ ...... ............. 15
V. Mangifera indica L. (Mango) ................................ . .. . .. . ............ 15

A. Propagation ................... . ..... .. .. . ...... .. .... .... .... ... ......... 15
B. Vegetative Growth ....................................................... 16
C. Flowering .. ... ................ . .. . . . .... . . .. .. . .... .. . . . ....... . ......... 16
D. Fruit Set and Development .................. .... .... .. .................. 18
E. Fruit Ripening and Postharvest Handling ...... . ... ....... .. .......... . .. 19
VI. Musa spp. (Banana, Plantain) .. ...... ............. .... ... .... ........... .. ...... 19
A. Propagation .. .. ............ . . .. .... . . . ... ....... ....... .. ....... ...... .. . 19
B. Vegetative Growth ...... .. .. . .. .. ..... .... ............ .. ...... .. ..... ... . 19
C. Flowering ..... ..... . ..... . .. .... . . ... ....... . .... .... .. ............... .. . 20
D. Fruit Set and Development ..... .. .. .... .............. .. ..... .. ....... . .. 20
E. Fruit Ripening .. .... .. ..... ... ...... .... .. ........... .. . . .... .. ........ .. 20
2 Plant Growth Regulating Chemicals
VII. Psidium guajava L. (Guava) ... . .. . . ... . . ....... .. . ....... . ... .................. 21
A. Propagation ....... ..... .. .................... .. .. . ...... .. ..... .... ...... 21
B. Vegetative Growth ................ . ..... .. ............................... 21
C. Induction of Flowering ............ ... .... . ... .. ......................... 21
D. Fruit Set and Development ............... ... ............................ 22
E. Fruit Ripening ........................... . ............................... 22
VIII . Theobroma cacao L. (Cacao) .............. ..... ............................... . 22

A. Propagation .... .. .... .. ... . . .. ..... . . . .. . .... . ... ................ ..... . .. 22
B. Vegetative Growth ...... ... ... ... .. . .. .. .......... ............ . ......... . 22
C. Flowering ..... .... .. . . ...... ... . .. .. . ...... .. ... ... . .... . . ....... .. .... .. 22
D. Fruit Set and Development ............. . .. .... .. ..... .. ...... . .......... 22
Acknowledgments ....... .. . . .. .... ..................... .. . . . ...... ................ ..... . 23
References ................ .. .. ...... .............. . ...... .... .. .... ................... . . . 23
I. INTRODUCTION
A wide variety of fruit and beverage crops are grown in the tropics but few of them are
intensively cultivated using modem crop production practices . The commercial use of growth
regulators on tropical fruit and beverage crops appears to be restricted to those crops that
have been grown on sufficient scale to justify the research support required to evaluate crop
responses to them.
The crops selected for this review are Ananas comosus L. Merr. (pineapple), Carica
papaya L. (papaya, pawpaw) , Mangifera indica L. (mango), Musa spp. (banana, plantain),
Psidium guava L. (guava), and the beverage crops Coffee arabica L., C. canephora L.
(coffee), and Theobroma cacao L. (cacao). The above crops are either important in inter-
national commerce and, thus , are grown on large plantations, often controlled by multina-
tional corporations (e.g., banana, cacao, coffee, pineapple), or are grown more or less
intensively on a somewhat smaller scale to supply local and regional markets. World pro-
duction in 1980 for the crops in 1000 metric tons are the following: pineapple - 7636,
papaya-1917, mango-14,342, dessertbananas-39,254, plaintains-21,265, guava
-no data reported, coffee- 4821, and cacao- 1557.4 It is likely that the above figures
do not include significant quantities of the fruit crops that are grown by home owners and
on small farms for direct consumption or for local markets .
Growth regulators are used on the above crops to enhance or speed propagation, induce
flowering, increase fruit size, promote ripening on the plant, and delay ripening after harvest.
Much work has been directed towards induction of flowering for year-round production and
on ripening to concentrate harvesting because fruit bearing tends to be seasonal, or biennial
as in some mango cultivars, even though the environment may be suitable for production
throughout the year. Commercial use of growth regulators has reduced costs and extended
supplies of commodities with significant benefits to both the producers and consumer.
Volume II 3
II. Ananas comosus (L.) Merr. (PINEAPPLE)
A. Propagation
Pineapple (Ananas comosus [L.] Merr.) is the only member of the family Bromeliaceae
that is cultivated as a food crop. Pineapple is grown from about 30° south latitude through
the equator to 30° north latitude. The major cultivar of commerce is the Smooth Cayenne
and there are several clonal selections of that cultivar being grown in Hawaii and in other
parts of the world. Several other cultivars of pineapple are grown, 22 · 112 · 113 · 122 but none
approach the prominence of Smooth Cayenne.
Pineapple is propagated exclusively by asexual means using the tops of fruits (crown),
offshoots borne at the base of the fruit which morphologically are crowns of vestigial fruits 78
and are referred to as slips, and shoots which arise from buds that develop in the leaf axils
(suckers), presumably as a result of the loss of apical dominance that occurs coincidentally
with inflorescence initiation. Pineapple is not propagated by seed because cultivars must be
highly self-incompatible to be commercially successful, 89 and cross pollination results in
progeny which shows variability typical of heterozygous parents.
Because vegetative propagation dependent on the production of crowns, slips, or suckers
is slow, various stem-sectioning techniques have been devised to increase the number of
propagules. 22 ·35 •46 •88 • 128 · 132 • 147 Pineapple also has been propagated by meristem budding
techniques 39 ·90 •91 ·93 · 104 • 134 • 140 • 160 and plantlets have been regenerated from callus. 92 · 115 • 146
There is no indication that important clones have been mass propagated by tissue culture.
In fact, a pineapple company in Hawaii abandoned attempts to propagate clones of pineapple
by tissue culture techniques because variability was observed among the progeny. Others
also have observed variants among pineapple plants propagated from tissue culture. 92 · 146
Plantlets regenerated from syncarp tissue exhibited greater variability than did plantlets
regenerated from meristem tissue of slip, crown, and axillary buds. 146 No studies were
reported where tissue-cultured plantlets were grown to maturity to assess the effects of
variability on productivity and fruit quality. Until such data become available, the mass
propagation of pineapple clones by tissue culture techniques should be approached with
caution. Tissue culture techniques may be most useful for the propagation of potentially
valuable new clones produced in a breeding program so long as roguing is practiced to
remove offtypes.
Rapid propagation of field-grown pineapple can be accomplished by spraying plants with
the rnorphactin Multi-prop® (Celamerck GmbH & Co.) or Maintain CF125® (U.S. Borax
Co.). 124 Multi-prop®, a mixture of methyl esters of 2-chloro-9-hydroxyfluorene-(9)-carbox-
ylic acid (chlorflurenol), 9-hydroxyfluorene-(9)-carboxylic acid (flurenol), and 2, 7-dichloro-
9-hydroxyfluorene-(9)-carboxylic acid (dichlor-flurenol), induces the formation of plantlets,
which are normal both in appearance and in growth rate, on the peduncle or on the developing
inflorescence of pineapple. 28 ·51 ·80 •81 • 150 The morphactin mixture is applied as an aqueous
spray over the plants after inflorescence development has been initiated (forced) with ethe-
phon (2-chloroethylphosphonic acid), alpha-naphthaleneacetic acid (NAA), or its sodium
salt (SNAA). The timing of application is determined by the effect of the prevailing envi-
ronment on normal inflorescence development51 •81 and by the number and final size of the
plantlets desired. Plantlet number is also influenced by plant size with larger plants producing
greater plantlet numbers than small plants. 150 Morphactin-induced plantlet number and mass
were greater when plants were forced with ethephon or SNAA than with ethylene or beta-
hydroxyethylhydrazine (BOH). 51
In Australia, the application of Multi-prop® 1 to 4 weeks after forcing resulted in 10 to
30 plantlets per plant weighing an average of 200 to 50 g each when harvested 43 weeks
after forcing. 5 1 Application of Multi-prop® 1 week after forcing resulted in the production
of more plantlets on the fruit peduncle, while later application increased those borne on the
fruit and reduced those borne on the peduncle. suso Sucker numbers were reduced by the
application of Multi-prop® 7 days after forcing, but not to unacceptable levels. Where large
plantlet numbers were produced, sucker growth was inhibited until the plantlets were har-
vested. The current recommendation for use in Australia is to force plants with ethephon
followed by the application of 2500 to 3000 e ha-I of a solution containing 22 ppm active
ingredient (a.i.) of Multi-prop® 10 to 14 days after forcing in early May (forced for winter
harvest). 51 · 149 In South Africa, 81 two applications of Multi-prop® have been made at 7- or
12-day intervals. Both 1.6 and 2.0 e of the chemical (11.2% a.i.) were applied per hectare
initially followed by 3.2 or 4.0 €/ha. No data were given on the spray volume used. Timing
of application after forcing was important with application being made sooner after forcing
in summer than in winter. Results indicated that up to one million plantlets could be produced
from 1 ha containing 43,000 ratooned pineapple plants. 81 Chlorflurenol is used commercially
in Hawaii and in Brazil but no data are available on specific practices.
Chlorflurenol is labeled as a poison but it has low toxicity to wildlife, fish, and mammals. 5
B. Vegetative Growth
There is no commercial practice utilizing growth regulators to promote vegetative growth
of pineapple nor were any published reports found suggesting that there is interest in, or
potential for, developing such practices.
C. Induction of Flowering (Forcing)

1. History of Growth Regulator-Induced Flowering
The artificial induction of flowering in pineapple dates to about 1874 when it was acci-
dentally discovered that wood smoke uniformly forced pineapple plants to flower. 22 This
discovery was exploited commercially in glasshouse culture of pineapple in the Azores 22
and in field culture in Puerto Rico. 42 • 119 Rodriquez 119 showed that a component of smoke,
ethylene, was as effective as smoke as a flower inductant. Acetylene gas, an acetylene-
saturated solution of water, and calcium carbide, which releases acetylene on contact with
water, were shown to force pineapple in the 1930s. 82 •83 · 87 In 1941, Johnson showed that an
ethylene-saturated water-oil emulsion containing colloidal earth could also force pineapple. 79
The events leading to the discovery that growth regulators with auxin activity could force
pineapple apparently are not documented. Several years of research must have prefaced the
first published reports in 1942 because data from Hawaii 20 were compared with data for
IAA and NAA from Florida in the same year. 25 A few years later it was shown that 2,4-D
also could induce flowering of pineapple and that Cabezona pineapple could be forced every
month of the year. 142 • 143
Beta-hydroxyethylhydrazine (BOH) which is thought to be an ethylene releasing agent 133 •138
was reported to induce flowering in Smooth Cayenne in 1955. 64 More recently, ethephon
(2-chloroethylphosphonic acid) and CGA-15281 (beta-chloroethyl-methyl-bis-benzyloxy-sil-
ane), which also degrade to release ethylene 19 •26 and stimulate the plant to produce ethylene, 148
have been shown to force pineapple. 15 •24 • 109
Many other compounds have induced flowering in pineapple, 41 •55 •65 - 68 but little or no
additional work has been done with many of them and none is of commercial importance.
The literature on those compounds studied in some detail is summarized in Table 1. Only
four compounds - acetylene (and calcium carbide), ethephon, ethylene, and NAA (or
SNAA) - have been used in commercial practice to force pineapple.
2. Absorption and Mechanism of Action

Green leaf tissue is required for flower initiation with gaseous ethylene. 141 Flowering did
not occur if only the basal portions of older leaves and the etiolated young leaves in the
center of the plant were retained. Plants defoliated to one large green leaf before treatment
or defoliated completely 2 days after treatment were induced to flower. 141
Volume II 5
Table 1
SUMMARY OF STUDIES ON THE USE OF GROWTH
REGULATORS AS FLOWER INDUCT ANTS OF PINEAPPLE
State,
Growth regulator country Cultivar Ref.
Acetylene and calcium Australia Smooth Cayenne 48, 87

carbide Brazil Pernambuco 43, 94
Perola 44
Colombia Smooth Cayenne 121
Cuba Red Spanish 53-55
Smooth Cayenne 53-57
Florida Abachi 26
Ghana Sugarloaf 98-100, 102
Smooth Cayenne I, 101
Guinea Smooth Cayenne 107, 113
Hawaii Smooth Cayenne 83
India Kew, Giant Kew 33, 70, 96
(Smooth Cayenne)
Kenya 41
Malaysia Sarawak (Smooth !56
Cayenne)
Singapore Spanish !51' 153, 155
Mexico Smooth Cayenne 2
Puerto Rico Smooth Cayenne 135
Sri Lanka Kew (Smooth 126
(Ceylon) Cayenne)
South Africa Smooth Cayenne 30
Taiwan Smooth Cayenne 75, 157, 158
Alpha-naphthalene-acetic Australia Smooth Cayenne 48, 69, 97
acid (ANA, NAA, SNA)
Bangladesh Giant Kew (Smooth 3, 129
Cayenne)
Honey Queen 130
Brazil Pernambuco 43
Florida Abachi 25
Ghana Sugarloaf 98, 99
Guinea Smooth Cayenne 107, Ill, 113
Hawaii Smooth Cayenne 20
India Kew, Giant Kew 31, 32, 70, 114
(Smooth Cayenne)
Kenya 41
Malaysia Singapore Spanish !55
Puerto Rico Cabezona 142-146
Red Spanish 146
Sri Lanka Kew (Smooth 126, 127
(Ceylon) Cayenne)
Mauritius 127
Taiwan Smooth Cayenne 75, 158
Beta-hydroxy-
ethylhydrazine
Australia Smooth Cayenne 48
Ghana Sugarloaf 98-100
Malaysia Mauritius 19
Puerto Rico Smooth Cayenne 133
Taiwan Smooth Cayenne 158
Table 1 (continued)
SUMMARY OF STUDIES ON THE USE OF GROWTH
REGULATORS AS FLOWER INDUCTANTS OF PINEAPPLE
State,
Growth regulator country Cultivar Ref.
2 ,4-Dichlorophenoxy- India Giant Kew (Smooth 76

acetic acid Cayenne)
Puerto Rico Cabezona 142, 143
Red Spanish 142, 143
Peru 58
Ethephon Australia Smooth Cayenne 48 , 52
Colombia Smooth Cayenne 121
Cuba Red Spanish 53, 54
Smooth Cayenne 53-57
Ghana Sugarloaf 98-!00
Guinea Smooth Cayenne !09, 113
India Kew , Giant Kew 34, 36, 70, 114
(Smooth Cayenne)
Malaysia Singapore Spanish 154
Philippines Smooth Cayenne !3, 14
West Africa Smooth Cayenne 71 , 72, 109 , !38
Ethylene Australia Smooth Cayenne 48
Brazil 47
Florida Abachi 141
Red Spanish 141
West Africa Smooth Cayenne 27, 37, 109
Indoleacetic acid Florida Abachi 25
India Smooth Cayenne 32
Naphthaleneacetamide Hawaii Smooth Cayenne 20
Florida Abachi 25
India Giant Kew (S mooth 114
Cayenne)
It is not known whether acetylene and ethylene are absorbed through the leaf trichomes 120
and the cuticle, throu gh the stomates , or all three . Evidence that forcing with unsaturated
hydrocarbons is much more effective at night 1•2 •30 •75 · 158 and in the absence of wind 75 indicates
that the stomata are an important avenue for entry into the leaf. The effectiveness of night
application is explained, at least in part, by the fact that pineapple is an obligate Crassulacean
acid metabolism plant so the stomata are closed during the day , open in late afternoon , and
remain open throughout the night.9
NAA and ethephon are rapidly absorbed and absorption is presumed to be through the
trichomes , or cuticle, or both. Within 5 min after labeled NAA having an activity of 60,000
cpm was applied to a leaf, 200 cpm of activity was detected in the stem apex. 86 Rain
following the application of NAA does not alter the induction of flowering . 11 3 Rapid ab-
sorption of ethephon is demonstrated by the fact that flushing the plant with 6 of water e
from l min to 2 hr after application has no effect on forcing. 50
Flower induction with N AA and ethephon is a response to a specific quantity of growth
regulator rather than to a volume or concentration. 7 1• 143 Forcing has been achieved by the
application of 5.0 mg of ethephon in a 20-,.L€ volume into the plant heart or to a l -cm 2 spot
on a single leaf. 10
The mechanism by which the various forcing agents initiate flowering in pineapple is not
known. Basic research on the physiology of flower induction has not extended knowledge
beyond the point where it is generally believed that induction occurs in response to ethylene.
Volume II 7
Although Gowing hypothesized that NAA induced flowering in pineapple by acting as an

antagonist of native IAA, 62 more recent data suggest that flowering occurs as a result of
ethylene production by the plant which is stimulated by NAA. 17 NAA induced evolution of
ethylene from pineapple attained a peak about 1 week after treatment 17 and fruit from
pineapple forced with NAA are harvested a week or more later than fruit from plants forced
with acetylene, ethylene, or ethephon. 30 ·9 H The approximately 1-week delay in harvesting
of fruit from plants forced with NAA may be due to a delay in the NAA forcing response.
However, it was recently shown that bract and flower primordia development began at the
same time in the Masmerah and Smooth Cayenne cultivars regardless of whether they were
forced with ethephon or NAA. 95 · 156 Significant amounts of ethylene are produced within 16
hr by pea (Pisum sativum) 18 and mung bean (Vigna radiata) 159 tissue incubated in media
containing IAA, so it is likely that sufficient ethylene is present at the apex of pineapple
within a few hours after treatment with NAA to induce flowering. Therefore, it seems likely
that the delayed maturation of pineapple fruits on plants forced with NAA is a side effect
of auxin forcing.
Recently, silver ion, an inhibitor of ethylene action, 12 was found to inhibit flower induction
of pineapple with ethephon, 123 further suggesting that ethylene is in some way involved in
the induction process.
3. Plant Susceptibility to Forcing

In order to force pineapple into flower, a minimum plant size is required. 14 •23 •33 Das et
alY found that a minimum of 12 leaves about 30 em in length was required before fruit
formation could be induced. In another study, 23 2-month-old plants (514 g fresh weight,
62.9 g dry weight) could not be forced with ethephon regardless of the night temperature
at which they had been conditioned. Four-month-old plants (878 g fresh weight, 114.6 g
dry weight) were readily forced if the night temperature was controlled at 20°C. 23 In com-
mercial practice the minimum plant size required for forcing is attained well before the plant
is large enough to produce a fruit of marketable size and plant and fruit mass are highly
correlated. 33 ·36 ·45 ·83 •87 ·97 · 110 • 137 • 143 The pineapple plant produces a terminal inflorescence so
the initiation of new leaves ceases at the time that flower parts begin to be laid down.
Because of this fact, comparisons of fruit characteristics of forced plants with plants which
flower naturally much later and, therefore, are much larger are not particularly meaningful
though such comparisons are fairly common. 55 ·98 - 100
Variation in susceptibility of pineapple to forcing agents is observed even after plants are
large enough to produce a marketable fruit. In some cases, larger plants were easier to force
than smaller ones, 23 •25 •33 but very large plants are also reported to be difficult to force. 14 •46 •97 •99 • 127
In many cases, the effects of plant size were evaluated by planting at one time followed by
sequential forcing. The forcing results, thus, were confounded with changes in environment
which could also influence plant susceptibility.
Pineapple plants are more susceptible to forcing near the time for natural inflorescence
initiation and this is especially evident when NAA is used. 25 •30 •31 •63 •69 •75 •99 • 107 •113 • 155 Plants
well supplied with water, having a high nitrogen content, and growing vigorously are more
difficult to force than those that are mildly stressed for nitrogen. 2 • 14 •41 •45 •72 • 107 · 109 • 113 Based
on data of Guyot and Py 72 for Smooth Cayenne, percent nitrogen on a fresh weight basis
in the basal white tissue of the youngest physiologically mature leaf (termed the "D"
leaf) 84 • 108 • 131 should be below 1.6% with good forcing being obtained when the value was
1.3%. It is usually recommended that no nitrogen be applied for 2 to 3 months prior to
forcing, especially if difficulty in forcing is likely or if NAA is used. 113
Forcing is more difficult in warm humid equatorial climates at most times of the year 139
and during the warmer seasons in other localities. Night temperatures of 25°C or greater
reduced the susceptibility of Smooth Cayenne to forcing with ethephon. 11 •23 When shelter
temperatures exceed 30oC during the day forcing is not recommended, as forcing percentages
obtained with ethephon 49 and NAA 50 declined approximately linearly with increasing tem-
perature. There is need for additional research on the effects of day and night temperature
on forcing success to improve the understanding of how the plant and the environment
interact.
Differences in susceptibility between cultivars exist; a greater quantity of growth regulator
was required to force Smooth Cayenne and Cabezona than was required for Red and Sin-
gapore Spanish.s3.144.153
4. Effects of Forcing on Fruit and Propagule Production

Forcing has little or no direct effect on fruit quality although fruit which develops during
cold or cloudy periods will have reduced sugar and increased acid levels. Fruit shape and
cannery recovery can be affected by forcing. NAA forcing resulted in a slightly larger fruit
than that produced by forcing with other growth regulators, 69 ·155 but generally the fruit is
more conical 112 ·123 so slice recovery may actually be reduced. Forcing with NAA also
increases peduncle length" 2 · 113 which increases susceptibility to lodging but, because of
better fruit exposure, may also be responsible for the reduced incidence of fruit disease
observed in comparative studies with ethylene. 123 The use of more than 40 mg of ethephon
per plant can result in fruit that are much shorter than normal. 30 ·50
The production of slips is consistently reduced with the use of NAA/5·112 but data for
ethephon are inconsistent. Some workers report reduced slip numbers when compared to
the control, 14 ·53 ·77 •99 ·100 while others report no effect. 24 The effect is less pronounced on large
plants than smaller ones 34 ·100 and control plants for all the above studies were larger and
flowered later than those forced with ethephon. The obvious conclusion is that the effect of
ethephon on slip numbers is due primarily to the fact that forced plants are smaller at the
time of flowering than those that flower naturally.
5. Forcing as a Commercial Practice

Though few data are available, ethylene generally is reported to be a more effective forcing
agent than acetylene, dry calcium carbide, ethephon, or NAA, especially in warm, humid
climates. 139 Ethylene is applied at night at a rate of about 800 g/ha in 7000 e of water
containing 0.5% activated charcoal in West Africa. 37 It is also used on one plantation in
Hawaii but no details are available on the specific quantities of material used. Special
techniques are required for its application 37 so the use of ethylene generally has been restricted
to larger plantations.
Water saturated with acetylene is used commercially by smaller growers in West Africa 139
where the scale of the enterprise is not large enough to justify the purchase of sophisticated
spray equipment. The recommended rates in Reunion and West Africa are to add 200 g of
calcium carbide to 75 or 100 e of cool water and apply 50 to 80 me of solution in the center
of the plant. 6·139 Py and Tisseau 112 reported that the percentage of plants forced increased
progressively as water temperature was reduced from 25 to 7°C. However, Aldrich and
Nakasone 2 consistently obtained higher forcing with water at 25 to 26°C than was obtained
at 3 to 5°C. They speculated that reduced forcing resulted from too slow a rate of acetylene
evolution from the cold water solution. A second application is often made 2 to 5 days after
the first to increase the forcing percentage. 2.43·44 ·75 ·98 - 100·113 • 139 The procedure and precautions
for mixing calcium carbide with water have been described in detail. 26 ·87 ·113
Dry calcium carbide has also been used but it can be less effective than acetylene in
solution 2·26 and may cause plant injury. 2·26 ·139 The details on the use of dry carbide in the
literature are often sketchy with amounts often given in qualitative terms. In Taiwan 0.5 to
0. 7 g of calcium carbide is placed in the center of the plant; 75 in South Africa a 7-mm pebble
of calcium carbide is used 30 while in Cuba 2.0 g per plant was applied. 55 If there is no water
Volume II 9
in the center of the plant and no rain or dew occurs soon after the application then water
must be poured into the center of the plant to generate acetylene.
Ethephon is more costly and less effective than acetylene 70 ·139 but has been reported to
be as, 99 and less effective 100 ·139 than dry calcium carbide. Because it is a stable liquid, it is
easily used by both large and small growers. Refinements in the use of ethephon such as
the addition of2.0 to 5.0% by weight of urea to enhance absorption, 6 ·30 ·48 .49 ·77 • 138 and sodium
carbonate34 or sodium borate49 ·52 to raise the pH of the solution to 9.0 increases the efficacy
of ethephon.
Commercial practices with ethephon vary in different parts of the world. In South Africa,
e
approximately 0.5 kg a.i. ethephon and 80 kg urea in 2250 to 3000 of water are sprayed
over 1 ha with the higher volume being used on ratoon crops. 30 Similar rates and volumes
are used in Hawaii. In Australia, consistent results are obtained by spraying 2000 C/ha of
a solution containing 120 to 180 ppm ethephon, 5% urea, and 0.5% sodium borate. 50 ·149 If
forcing is expected to be difficult, a second application is recommended 14 to 20 days later;
however, if the second application is made earlier than 7 days, forcing may actually be
depressed. 52 Ethephon is also effective if applied in the center of the plant in volumes of
20 to 50 me containing concentrations of ethephon and adjuvents comparable to those used
for spray applications.
NAA is less effective than acetylene, 2·69 ·70 ·75 ·155 calcium carbide, 2·98 ·99 or ethe-
phon,70·75·98·99·114 and its use in commercial practice may be waning. It is no longer used in
Hawaii. Its primary advantages are low cost and ease of storage and handling. The disad-
vantages were discussed previously. The range of effective quantities or concentrations is
quite narrow with poor or no forcing resulting from too low or too high a quantity of
chemical. 62 ·112 In most studies and in commercial practice, 20 to 50 me of a solution
containing 5 to 20 ppm of NAA or SNAA is poured into the center of the plant or 2000 to
3000 e/ha are sprayed over the foliage. 30 .48·75 ·155 The greater spray volume is recommended
for use on ratooning plants in Australia. A second application is made 10 to 20 days later
when forcing is expected to be difficult. 30 ·48 As with ethephon, if the second application is
made too early the percentage of plants forced may actually be reduced.
D. Fruit Development
Early work on the effects of growth regulators on fruit development of pineapple showed
that the application of NAA or Fruitone® (composition not known) to the developing inflo-
rescence when it was less than 2.5 em in diameter increased fruit size as much as 25% but
decreased fruit-soluble solids and reduced slip and sucker numbers at high concentrations. 21
There was no effect on fruitlet number. Fruit maturity was also delayed. Studies with NAA
and Fruitone CPA® (active ingredient, 2-[3-chlorophenoxy] propionic acid) since that time
have confirmed the earlier results and extended them so that commercial exploitation is now
possible. Both Fruitone CPA® and NAA significantly increased fruit
mass 3·8·29 ·73 •85 ·103 ·105 ·118 ·125 ·135 ·145 ·152 with Fruitone CPA® producing somewhat greater fruit
weights than NAA. 135 Application can be made up to 8 weeks before harvest but most
workers recommend application after anthesis when the corolla has dried. The increase in
fruit mass at growth regulator concentrations which were not phytotoxic resulted from
significant increases in fruit length 103 or diameter 145 ·152 or possibly from increases in both
dimensions, since Dalldorf29 and Vieira et al. 145 show average fruit length from Fruitone-
treated plants was 0.7 to 2.0 em greater than the control, though the differences were not
always statistically significant. If fruit length is consistently increased by a treatment even
though the increase is not statistically significant, it may be very significant in terms of
processing because a 1.0-cm increase in fruit length could increase cannery slice recovery
by as much as 10%.
The use of NAA and Fruitone CPA® delays fruit maturity by 1 to 4 weeks 3·73 ·75 ·118 ·135 ·145 ·152
with the amount of delay depending primarily on the stage of development at the time of
application. 135 Fruit crown length and mass 29 • 103 • 105 • 145 or average size 118 are decreased by
NAA at concentrations of 200 ppm or greater and by Fruitone CPA® at concentrations of
75 to 100 ppm. 29 •5 ° Fruit crowns show symptoms of phytotoxicity at 500 ppm of Fruitone
CPA ®29 and lower concentrations applied 7 to 8 weeks before harvest cause the crowns to
become brittle 135 which may affect their suitability as planting material.
NAA and Fruitone CPA® have been reported to delay slip or sucker development, or
both, especially at higher concentrations of the growth regulators, 21 ·73 ·75 •85 • 105 · 135 but to pro-
mote slip or sucker production, especially at low concentrations of growth regulator, 21 • 103 • 135
or to have no effect on sucker production. 103
Fruit treated .with Fruitone CPA®, and especially with NAA, often ripens internally before
the shell turns yellow, making it difficult to determine the proper time to harvest and adversely
affecting juice quality and color. 16 •73 ·75 Peduncle and core diameter are increased by
NAA 21 .7 3 ·75 •85 • 105 and total soluble solids is either unaffected by the two chemicals 145 • 152 or
reduced, 16 •21 ·73 •75 • 103 • 105 • 135 while titratable acids have been reported to increase, 16 • 145 • 152 remain
the same, 103 or decrease 73 •75 •85 with treatment. The preponderance of evidence suggests that
a reduction in soluble solids would be the typical reponse.
Maximum increases in fruit size with a minimum of side effects are obtained with a single
application of 100 ppm of NAA 75 or 100 to 200 ppm of Fruitone CPA®, 29 • 118 and adjuvents
such as urea, sodium carbonate, or a spreader-sticker do not enhance the effectiveness of
Fruitone CPA®. 145 Fruit sizing hormones are not used in Australia and NAA is banned by
the cannery because of its adverse effects on fruit quality. 16 Skirmish tests with Fruitone
CPA® in Australia indicate it increases fruit size with fewer adverse effects than NAA but
fruit are already larger than is considered optimum by the cannery, so the likelihood of a
fruit sizing chemical being supported for registration is remote. 50 • 149 Fruitone CPA® is
registered for use in South Africa as a fruit sizing agent or to delay harvest of plant crop
fruit to be processed or marketed fresh and for first and second ratoons. Recommended rates
are 6 m€ Fruitone CPA® per liter as a directed spray on the fruit at the dry petal stage, 4
€ Fruitone CPA® per I 000 € applied per hectare up to 6 weeks before harvest if the crowns
are to be used for planting, and 6 €12000 €/ha if the crowns are not required. There is much
interest in the potential for Fruitone CPA® in Hawaii and an Experimental Use Permit has
been obtained. Slice recovery is said to be increased significantly with the use of Fruitone
CPA®.
A spray application of 50 m€ of a 50-ppm a.i. solution of Promalin® (Abbott Laboratories,
a mixture of gibberellins GA4 , GA 7 , and the cytokinin 6-benzyladenine) increased fruit
weight 13%. 10 Fruit top diameter was unaffected or slightly increased while fruit length was
increased by 1.4 em. The effect of Promalin® on soluble solids was not significant and the
effect on titratable acids, though significant, ranged from 0.86 to 0.96%. The compound
appeared to offer sufficient promise to warrant further study.
E. Fruit Ripening
Labor costs for fruit harvesting operations can be quite large. The use of forcing agents
on pineapple narrowed the fruit ripening peak so that fruit could be harvested with as few
as three passes through the field, thus, greatly reducing harvesting costs. With the discovery
that ethephon could ripen fruit ahead of the normal date of maturation, 40 interest was aroused
in the possibility of ripening pineapple so that all fruit could be harvested at one time.
Ethephon applied from 1 to 6 weeks before the projected harvest date induced early
ripening of the fruit, but it was evident that premature ripening essentially fixed the quality
of the fruit at the time of ethephon application. 7 · 38 • 102 • 106 · 116 · 117 · 154 Fruit total soluble solids
increase more or less linear! y with time during the 30 days prior to the completion of ripening
while flesh color changes due to an increase in carotenoids during the final 20 days of
Volume II 11
ripening. 60 Artificial ripening of Smooth Cayenne fruit before the normal maturation time
results in a uniformly colored fruit shell, but fruit yield , sugar content, flavor, and flesh
color are typically those of an unripened fruit at the same stage of development. 7 · 27 · 106 · 11 6 • 117
In contrast with the results for Smooth Cayenne, Norman 102 and Wee and Ng 154 reported
that the application of ethephon to fruit of Singapore Spanish and Sugarloaf cultivars up to
4 weeks before normal ripening had little or no effect on fruit yield or quality.
Ethephon is registered as a fruit ripener for pineapple in Hawaii and is regularly used on
ratoon crop fruit. The extent of its use on plant crop fruit is determined in part by the
probability that ripening will also initiate flowering of suckers that would be relied upon for
a ratoon crop. Ethephon is also used in the Ivory Coast to ripen fruit and the recommended
practice is to apply 1.5 kg a.i ./ 1000 f /ha directly to the fruit or the same amount of ethephon
in 2000-f water as a spray over the whole plant. 139 Ethephon is not registered for use as a
ripener in Australia but studies on preharvest ripening have been conducted. The addition
of urea is not recommended because it increases the effectiveness of ethephon as a forcing
agent while having a minimal effect on its efficacy as a ripener. 50 Although once-over
harvesting is possible with ethephon ripening, the best practice is reported to be to make an
initial picking, usually a light one, after which the remaining fruit are ripened with a 1000-
ppm spray solution applied at the rate of 1000 f/ha. 149 With the recommended practice, fruit
quality was found to be high with no adverse effects on fruit yield or cannery recovery. 149
F. Postharvest Treatments
As a part of studies to examine the effects of growth regulators on pineapple, 6 5 . 68 the
effects of several hundred chemicals on postharvest ripening and senescence were also
evaluated. 61 It was found that a number of compounds including NAA and 2,4,5-trichlo-
rophenoxyacetic acid (2,4,5-T) could delay the onset of senescence, thus, extending the
shelf life of pineapple after harvest. 5 9 Additional studies in Taiwan and Australia 74 · 136 have
confirmed these results and shown that dipping fruit in NAA will extend the shelf life
sufficiently so that fruit can be shipped to more distant markets. Dipping in NAA also
reduced losses to rot by as much as 50%. 136 NAA is registered in Australia for use as a dip
at a concentration of 100 ppm . For optimum shelf life, the fruit is picked "mature green"
or with just a tinge of color and dipped to immerse the fruit but not the crown.
G. Future Research
Although growth regulators provide a greater degree of control over fruiting and devel-
opment in pineapple than is possible for most other crops, there is still need for additional
research. Forcing success can at times be unpredictable. Increased knowledge about the
physiology of the process could increase the reliability of forcing during periods when the
probability of success is low. There also is a need for a chemical forcing agent that is as
easy to use as ethephon but is as successful as ethylene and acetylene for use in the warm
humid tropics. Flower inhibitors are needed to prevent precocious natural flowering and
such compounds might have use in preventing the induction of suckers by ethephon applied
as a ripener. Growth regulators used to increase fruit size could be improved by the elim-
ination of their adverse effects on fruit quality. Ripening agents which do not force suckers
are needed. Finally, the development of growth regulators that could further extend shelf
life would greatly expand the market potential for fresh fruit in temperate zone areas far
from the centers of pineapple production.
III. Carica papaya L. (PAPAYA)
A. Propagation
Papaya is reproduced exclusively from seed for commercial plantings . Germination varies
from under 20 to 90%. Consistently high germination is desirable and growth regulators
have been applied to the seed in an effort to improve germination. 164 · 169 ·170 · 172 ·176 ·178 • 179 The
effects of gibberellin were not consistent but, in general, the percent of germination was
not improved although the rate of germination and of stem elongation were increased. 176
An increased rate of germination over the range 50 to 500-ppm GA 3 was reported, while
NAA and IAA depressed germination. 176 Presence of the sarcotesta (a gelatinous aril sur-
rounding the seed) reduced the effect of both gibberellin and the auxins. 176 • 178 • 179
The rooting of cuttings is possible 168 • 171 · 177 using auxins such as IAA and IBA. The practice
is not used for extensive commercial plantings.
The papaya growth habit is excurrent and flowers are borne in the axil of each leaf. There
is no need for pruning or establishment of a fruit-bearing scaffold. Normal cultivars are
usually too tall to harvest from the ground after about 3 years, but there is little need to
control vigor with retardants as semidwarf cultivars are available which begin to bear within
6 to 7 months from seeding at heights under 1 m. 165 · 180 Since these do not elongate as rapidly
as normal cultivars, the time that harvest can be effected from the ground is prolonged.
Sex determination tests have been developed for use on vegetative seedlings. 166 This
would obviate the need to rogue unproductive male plants. It also suggests the possibility
for sex modification by plant growth regulators.
C. Flowering
Precocious flowering has been a trait selected by breeders; 167 thus, chemicals have not
been used to accelerate flowering. Flowering can begin as early as 4 months from seeding
in precocious strains 165 • 167 or as late as 13 to 14 months 167 at a height of 1.2 to 2.0 m.
Papaya is a polygamous plant in which commercial strains have been selected which bear
perfect flowers. Factors such as heredity and environment affect both the sex expression
and fertility of the flowers 15 with short days and cooler temperatures favoring femaleness
in hermaphroditic flowers.
Ethephon has been used to modify sex expression towards femaleness. 174 Spray treatment
of the two-leaf-stage seedling with 100 to 300 ppm ethephon repeated 15 to 30 days later
improved femaleness to near 90% vs. only 30% for control plants. Treatment at an early
stage gave a more marked effect than later, and three or more treatments were unnecessary.
D. Fruit Set and Development

No growth regulators have attained commercial use in fruit set as every flower sets a fruit
in the hermaphroditic strains. Development of female-sterile flowers occurs in some cultivars
during warm, dry weather, but the hormonal basis has not been investigated. Fruits which
are carpellodic or misshapen are thinned manually. Ethephon (300 ppm) applied at the brown
stigma stage of flowering and twice again over the next 15 days is reported to have increased
fruit size and weight, 161 although it would seem that caution should be exercised due to the
usual abscisson-inducing effect of this chemical on other fruits in an early stage of development.
Green fruits are used in the production of papain which is extracted from the latex. Papain
yields were increased fourfold by the application of 34 mM ethephon in coconut oil to the
fruits. The effect was due to the increased flow of latex. 162 • 163 • 173
E. Fruit Ripening
There is little need to enhance ripening as natural ethylene production stimulates rapid
ripening. Ethephon will enhance ripening and is reported to increase total sugars and reducing
sugars and to decrease acidity 173 • Delay of ripening with plant growth regulators to permit
rougher handling during shipment and to prolong shipment time has apparently not been
studied.
Volume II 13
IV. Coffea arabica L., C. canephora L. (COFFEE)
A. Propagation
Coffee may be established from seed or from vegetatively increased clones. Seed prop-
agation is common as fresh seed germinates readily, and seedlings of the arabic a coffee are
very uniform, but there is little commercial use of growth regulators to improve either
percentage or rate of germination. Seed soaks in auxin (IAA, IBA at 25 and 100 ppm) were
reported to improve germination percentage and 0.1% folic acid, 0.1% ascorbic acid, and
1500-ppm thiourea each hastened germination 5 to 7 days. 199 Exogenous gibberellin and
abscisic acid inhibit germination while kinetin reverses the effect. 220 •221
An early review 197 reported all auxins and commercial formulations, such as Rootone®,
Hormodin®, and Seradix®, had been used to improve rooting of cuttings, but specific
recommendations were not derived. Leafy cuttings of partially hardened wood can be rooted
under mist using 1000- to 4000-ppm IBA or NAA in talc dusts or in alcohol as a dip, 202 •203 •217
but care must be taken to choose vertical rather than horizontal shoots as a new growth
perpetuates the habit of the original cutting.
Several systems are used to develop the framework of the coffee plant. Growth regulators
have not been used commercially to stimulate suckers or basal breaks. Under high density
planting conditions, chlormequat has been helpful experimentally in retarding growth and
setting flower buds. 205 ·206 The rate of use was 1000 ppm applied four times at weekly intervals.
Daminozide also retarded growth but had less effect on flowering and yields. 206
Vegetative growth is controlled largely by the availability of moisture, although daylength
and temperature have also been implicated. 208 Flushes of growth occur following periods of
significant rainfall while growth ceases during dry periods. Growth cannot be induced in
the dormant state with gibberellins, but during flushing gibberellins may be limiting and
shoots are more sensitive to their application. 190 ·207
A pruning system has been developed to encourage uniform shoot development. 187 This
factor is important to uniform flowering and fruit development. On established plants with
a vigorous root system, growth regulators are not needed to stimulate shoots, but their
potential for shoot stimulation should be examined for younger plants and old, poorly tended
plants.
C. Flowering
Flowering initiation and development in coffee has been the subject of much research and
the resulting papers and reviews 183 . 185 • 191 •204 •207 •208 have become important examples of the
effects of environment upon the hormone systems involved in flowering. Flower initiation
must be considered separately from development as different environmental factors control
these processes.
It has been experimentally demonstrated that most varieties of Coffea arabica are short
day plants with a critical photoperiod of 13 hr. 214 However, mature plants seem to hold the
short day stimulus for long periods and are not as sensitive to long days as the seedlings
used to establish the critical photoperiod. 196 Between 20° Nand 20° S inductive photoperiods
prevail nearly the year around and permit flower initiation throughout the year. While
initiation can apparently occur at any time photoperiod permits, other factors such as tem-
perature, 195 · 196 •224 light intensity, 200 and rainfall 183 •200 ·225 also modify flower formation. Floral
initiation is favored by cool temperatures, 195 · 196 •224 while high temperatures decrease flower
production. 184 ·200 Following initiation flower development proceeds slowly until the buds
are 4 to 6 mm long at which point they enter a state of dormancy.
With respect to growth regulator use to enhance or delay flower initiation, both gibberellins
and growth retardants have been examined. Repeated applications of 100-ppm GA 3 over a
4-month period caused greater extension of the branches, making a more open plant, but
flower bud formation and development was retarded. 195 Where chlormequat was applied to
coffee, bean production was ultimately enhanced 30 to 35%, suggesting a favorable effect
on flower initiation. 205 It was theorized that this was due to inhibition of gibberellin synthesis
by the retardant.
It is of importance that flowers initiate and develop uniformly and that anthesis occurs
uniformly in order to concentrate the harvest period. The dormancy of the flower buds is
under the control of endogenous hormones, notably ABA, gibberellins, and cytoki-
nins.188·189·19u07 ABA, and perhaps other inhibitors, maintain the dormant state for about 2
months after which flowering can occur unless an imposed dormancy is maintained by an
inadequate supply of water. In response to irrigation or rainfall, an increase in gibberellin 183 ·
185 ·188 and cytokinins 189 overcomes the dormancy imposed by ABA. Coupled with rainfall
may be sudden 5 to 10°C drops in bud temperature which also seem to trigger development
leading to flower opening. 188 Exogenous gibberellin seems to replace these stimuli, 182-184 ·194 ·222
but development still requires adequate water. Exogenous ABA can prevent bud break, 191 ·193
although the effect decreases after 2 months of dormancy, and may do so by regulation of
the free-bound state of gibberellins 188 or by preventing the differentiation of xylem strands
to the flower bud. 191 ·207 Because ABA levels in the flower buds do not change until after
bud break, it is thought that the balance of promoters (BA and cytokinins) to inhibitors
(ABA) is the mechanism of control for bud break. 191 ·207
Despite our understanding of the physiology of dormancy and bud break to this point,
actual application of gibberellin in the field has yielded variable results. GA 3 has been less
effective on C. canephora 219 and has not always yielded definitive results on C. arabica. 194
Two key elements must be integrated and controlled: water supply and uniform shoot
development. Given these, it is possible to concentrate flower initiation and flower opening.
The use of GA 4 + 7 or PromaJin® (20 ppm applied three times at 2-week intervals or until
flowers open) permits normal floral development 194 where rainfall has been insufficient to
stimulate uniform flowering following bud break.

Many flowers are borne in a cluster and a need to enhance fruit set is seldom necessary.
On the contrary, thinning could be helpful in increasing grade by reducing fruit competition.
In certain high elevation growing areas, growth regulator use to overcome poor fruit set
caused by frosts would be desirable. To this end, it may be possible to use gibberellic acid
to increase fruit grade and weight. Since fruit growth is nearly completed 12 to 14 weeks
after flowering, 208 gibberellin is likely to be effective only in early stages of fruit growth.
GA 3 (Progibb® formulation of GA 3 ) at the rate of 100 ppm applied two to four times at 2-
week intervals to trees bearing 4- to 10-week-o1d fruits increased yields due to increases in
both size and weight. 207 •209 ·211 The crop was pushed back to a "late" crop in one instance
of its use, but yields of early and midseason crops have also been increased. Timing of
gibberellin application with respect to stage of fruit growth must be worked out if altered
bearing patterns are desired. 209
Early fruit drop, occurring 8 to 12 weeks after flowering during rapid enlargement of the
fruit (stage 2), may be reduced by application of 2,4-D or 2,4,5-T. 198 Hormonal imbalances
are, thus, implied. The concentration used must be low to avoid phytotoxicity.
Ethephon has been used experimentally as a thinning agent. Applied 6 days before flower
opening, it had little effect. Related experiments showed that 10- to 13-week-old fruits could
be induced to abscise following treatment with 800- to 1600-ppm ethephon although there
was about 10% leaf loss as well. The potential for some selectivity- thinning the products
of particular flowerings - seems to exist. 192 ·212
Volume II 15
Evaluation of new products will continue. The application of sodium mono-nitroguaiacol

(Atonik®) at 250 ppm has improved yields over three seasons of testing. 223 A biostimulant
known as Ergostim® is said to have caused an increase in fruit number and size and more
uniform maturity. 186
E. Fruit Ripening
Sprays of 2,4,5-T (50 ppm) were shown to accelerate ripening when applied to nearly
mature green berries (I . 6 to I . 8 em) of C. canephora, but induced abscission of small fruits.
GA 3 (10 to 500 ppm) delayed ripening. 181
The use of ethephon to hasten maturity and concentrate ripening has received attention
everywhere coffee is grown. 181 · 192 •201 •212 •213 •215 •216 •218 •219 The fruits need to be at 75% of their
full development for the optimum result. 207 •210 • 211 While concentrations as high as 1400 ppm
have been used, 212 rates of 250 to 480 ppm also resulted in significant acceleration of
ripening. 201 •213 ·215 The recommended time of application to accelerate ripening is about 6 to
8 weeks before the main flush of ripening, but if trees are bearing multiple crops from
different flowering times, some abscission of immature fruits is induced and not all crops
can be concentrated for one picking. 207 •211 Thus, flowering must be concentrated in order
to use ethephon for concentrating the harvest. The coupling of hand picking of the early
crop with a subsequent spray of ethephon to ripen the rest of the crop may be the direction
that plant growth regulator use would take. 210
Ripe berries do not normally abscise, but could be induced to loosen with ethephon. 192 •218
Coupled with TIBA at 50 ppm, 1400-ppm ethephon induced loosening of about 50% of the
ripe berries so that they could be shaken off.2 11 •212 This would make possible mechanical
harvesting.
Concerns that accelerated ripening or abscission of younger fruits would have an adverse
effect on yields and and beverage quality have been investigated. 192 • 194 •201 •213 ·219 The beverage
quality did not suffer when applications were made to uniform crops in which the fruits
were in their last stages of maturation, 192 •201 but in uneven crops, the outer flesh of younger
fruits ripened while the bean remained immature and the final product was judged
unsatisfactory. 194
The potential for plant growth regulator use on coffee requires consistent results in tests
over many seasons in different geographic locations. The approximate stage of maturity for
the desired response must be known. The quality of the final beverage product must not
suffer, nor should there be adverse effects on the plant or on subsequent fruit set. Much
progress has been made with ethephon towards resolving these problems, and its commercial
use seems likely to be realized.
V. Magifera indica L. (MANGO)
Among tropical tree fruits, the mango is the most widely grown. Some 87 countries
produced almost 14 million metric tons in 1977 237 with 65% of this produced in India. A
significant contribution to expansion of the industry, especially in the Philippines, has been
the development of a chemical treatment to induce flowering and relieve the problem of
alternate year bearing. The extent of developmental research in this area has made the mango
the "apple" of the tropics.
A. Propagation
While mangos may come true from seed if the cultivar is a nucellar apomict with po-
lyembryonic seed, much of the plant material used for commercial orchards is vegetatively
propagated by grafting, budding, stooling, air layering, or cuttings. An extensive literature
review has been prepared. 247
One of the preferred approaches for cuttings involves etiolation for 3 to 4 weeks of a
portion of the branch which will be the base of the cutting. After it is cut from the tree, the
base of the cutting is given a 30-sec dip in auxin. Similarly, girdling the branch 3 to 4 weeks
prior to taking the cutting and treating with auxin has produced a high rate of success. The
auxins most commonly used are NAA and IBA at 0.5 to 1% concentrations. Among other
growth regulators, phenolics 276 ·280 (p-hydroxybenzoic acid and p-coumaric acid at I0- 3 M),
flavonols 276 (rutin and quercitin at 2000 ppm), and retardants and ethephon 279 ·281 (chlormequat
at 1%, ethephon at 250 ppm applied to the stock plant; ethephon at 50 ppm as a 24-hr soak)
have also been reported synergistic in improving rooting.
Many techniques such as etiolation, girdling, and application of auxin have been successful
for stooling and air layering as well. 234 ·258 ·268 •282 A 2% preparation of IAA in lanolin applied
to an incision above a shield bud on stock budded during its rest period is reported to have
improved take. 257
Growth regulators have not been used commercially in the development of the framework
on which the crop is carried nor in the control of vigor. Experimental results with daminozide,
chlormequat, and maleic hydrazide have shown some retardation and inhibition of growth,
often coupled with enhanced flower bud induction. 248 ·26 H 65 •274 •286 •289 •293 The use of ethephon
to induce flowering apparently has not caused a noticeable decline in vigor. 243
Growth retardation by chlormequat and daminozide may be due to countering gibberellin
production. In the "off" year in biennially bearing trees, endogenous gibberellin content is
high 270 and treatment with chlormequae 93 and daminozide 248 after fruiting in an "on" year
increased the number of flowering shoots the following year. Gibberellic acid has been
shown to delay or inhibit flowering 260 ·283 as well as stimulate bud break. 286 In the latter case,
50 ppm of GA 3 or NaGA 3 was used with two to four applications.
Research on the timing of retardant use for vegetative growth control and to induce
flowering suggests application over a period of several months with 2 to 4 weeks between
spray treatments. Chlormequat has been applied at rates of 3000 to 5000 ppm. 274 •293 Con-
centrations of at least 7000 ppm daminozide seem to be needed for significant retardation
of growth. 248 Maleic hydrazide delays bud break at 5000 ppm. 248 Such uses have not been
commercially accepted.
C. Flowering
It has long been known that the smoke from smudging fires could cause flower bud
differentiation 226 •255 •256 in mango as in pineapple with the ethylene component considered
the induction stimulus. 277 That flowering in mango is hormonally controlled now seems well
established. 270 ·275 ·287 When ethephon was developed, experiments in the Philippines showed
that it was effective in inducing flowering in the mango. 250 Since then, extensive research
has been conducted in mango-growing areas to utilize this response in controlling flowering
and fruiting. 230.235,242,243.269
Since concentrations of ethephon in excess of 500 ppm cause defoliation, it has been
recommended that concentrations of 150 to 250 ppm 227 • 242 · 243 · 252 be applied at 2- to 3-week
intervals with up to four or five applications. The timing of application is related to the
stage of growth and time of previous harvest, and recommendations vary from country to
country. The mango is at least a facultative short day plant, and ethylene or ethephon
application during the fall-to-winter period assures that terminal flower buds are set on a
high percentage of shoots, including young shoots. 238 Even auxiliary buds have been observed
to set flower buds when no terminal growth was made during a heavy "on" year. 243
Gibberellin applied just before flower bud differentiation is reported to have delayed
development although it did not prevent it. Among the benefits reported were an increased
number of perfect flowers and improved fruit set. 283
Volume Jl 17
FIGURE I. Exampl es of commercial preparations for forcing fl owering of mango .
The need to repeat applications of ethephon poses its own set of problems as bearing trees
achieve a considerable size. Attempts to devise a formulation which could be applied one
time with a prolonged period of activity without defoliation side effects led to the development
in the Philippines of Flower Set. It was reported effective in inducing flowering on shoots
younger than those normally responsive to smudging and on branches where fruits were just
harvested. 230
While ethephon was quite effective, it was also an expensive manufactured chemical.
Research in the Philippines led to the discovery that I% KN0 3 sprays were effective in
inducing flowering. That it is induction rather than merely breaking dormancy of preformed
flower buds was circumstantially established by Barba, 230 who suggested an effect on the
nitrate reductase system with ultimate production of ethylene by the plant as a possible mode
of action. The very rapid appearance of flowers 229 •230 ·2 3 8 ·240 •24 1·267 as few as 7 days after
spraying in some cultivars results because flower initiation and development and panicle
elongation progress without the intervening dormant period 262 •275 common in temperate fruit
tree species. Thus, flowering may occur several weeks ahead of nontreated trees and could
be induced out of season where weather conditions will permit fruit set and development.
A number of flower-inducing materials have been marketed in the Philippines (Figure 1) .
Most of these contain potassium nitrate (Table 2) . The recommended concentrations vary
from 1 to 2% although higher concentrations are effective but may cause burning . 2 30 •240 ·267
No reports of the effect of KN0 3 -ethephon combinations were found .
A floral malformation occurs in some cultivars which may be due to a hormone imbalance
at the time of flower initiation . 259 NAA sprays (100 to 200 ppm) reduced the problem
somewhat as did deblossoming , 245 while gibberellin (100 to 250 ppm) and ethephon (50 to
150 ppm) increased the incidence of malformed panicles.253 Coupling the fungicide , Bay-
fol an®, with a chlormequat spray (500 ppm) also reduced malformation. 253 The root of the
problem remains unsolved as Fusarium infections and high temperatures have also been
implicated. 245
In Mangifera a/tis sima , forcing with KN0 3 was successful in only 2 out of 11 trials. 239
The concentration range for success was from 10 to 40 g!€ .
The wind-borne pollen of the mango can be allergenic to some persons. Where the crop
is not desired , as in city streets , eliminating the flowers will eliminate a subsequent fruit
Table 2
MANUFACTURER/DISTRIBUTOR
(PHILIPPINES) AND ANALYSIS FOR
NITRATE OF FLOWER INDUCERS•
Manufacturer/ Analysis for

Flower inducer distributor nitrate
Agriblum Agrix +
Angromangobloom Agricor
Flowerblum Valenzuela +
Golden Crownb Union Carbide +
Hormudge Shell +
Mangobloom Samsom +
Mangobloom Planters +
Mango Flowerset Bayer +
Manggrow Kabukiran +
Mangotone Agchem +
Mangovit Conserve +
Miracle blum Philippine Orchard +
Reb loom Atlas +
Surebloom Alden
Courtesy of N.D. Bondad.
" Presence of nitrate is based on company report.
drop problem and reduce the source of pollen. Ethephon at 500 to 1000 ppm applied as a
spray to the panicles at an early stage of blossoming caused deblossoming on one cultivar
and killed the panicles of another. 244 While this is an isolated report, such chemical de-
blossoming could be of value in landscape situations and possibly in reducing the size of a
crop during an "on" year in a biennial cycle.

The use of auxins to enhance fruit set in temperate fruit trees finds it parallel on mango
as well. 228 •284 •288 •290 Despite the large number of flowers on a panicle ( 1200 to 1500), very
few fruits set, and this may be as low as one fruit per panicle for some cultivars. Much
natural drop occurs. Extensive early fruit drop stimulated examination of a variety of growth
regulators to effect its control. 246 •273 •278 •284 •285 Prominent among these have been studies with
auxins: NAA, 2,4-D, 2,4,5-T, and 2,4,5-TP. Reduced fruit drop was observed for NAA in
the concentration range of 10 to 40 ppm. 228 •271 •284 Similar concentrations of 2,4,5-T and
2,4,5-TP were not as effective 246 •261 although the combination of 2,4,5-T with 2 to 4% urea
was reported to increase set. 284 Among the side effects reported have been hastened
maturation 261 and increased size. 246 •285 There appear to be differences in cultivar responses.
The commercial preparation, Planofix®, which contains NAA, improved fruit set when 10
to 30 ppm a.i. was applied twice 15 days apart to the fruits at marble size, with 20 ppm
giving best results, 228 but in another experiment it was not effective. 271 The growth retardant,
daminozide, is credited in one experiment with reducing drop when 100 ppm was sprayed
on pea-sized fruits. 271
Reports are mixed on the use of auxin sprays to reduce late fruit drop. Sprays of 2,4-D
(10 to 20 ppm) and NAA (30 to 40 ppm) were reported to reduce fruit drop when applied
to fruits 10 to 12 em long 285 and had no effect on 7- to 8-cm fruits. 246 Similarly, 2,4,5-TP
applied 4 weeks before harvest had no effect on drop. 261
Volume II 19
There is evidence for a role of gibberellin in fruit development with 100-ppm GA 3

enhancing fruit size and weight when applied at the pea stage. 272 In another experiment,
three applications of gibberellic acid at weekly intervals to young fruits increased fruit size
and weight, total soluble solids, sugars, and ascorbic acid content. 292 All concentrations up
to 200 ppm were effective. Commercial application is unlikely because of cost.
While parthenocarpic fruit development is occasionally noted and could be used to avoid
certain seed-eating pests, there has been no commercial development of growth regulators
for this purpose.
E. Fruit Ripening and Postharvest Handling

Application of 200-ppm ethephon when the fruit was pea sized advanced ripening by 10
days; 80 days vs. 90 for the control. 272 Most reports on ripening of mango, however, deal
with postharvest use of ethylene or ethephon.
Since mangos are often picked and shipped when mature-green and do not show desirable
external color development even when ripened in storage, the possibility of using ethylene
as for bananas was proposed and found to be adaptable, 231 •232 as the mango is a climacteric
fruit. 266 The conditions for ripening of mature fruit with 10- to 20-ppm ethylene include a
complete air exchange of the ripening room every 2 hr and 92 to 95% relative humidity. A
temperature of 21 °C is used to achieve a balance between the rate of color development and
ripening although a temperature as high as 30oC during exposure to ethylene accelerates the
ripening process. Optimum treatment times ranged from 12 to 24 hr depending on maturity
of the fruit. The exudate from the stem end must be removed as covered areas did not
degreen. Ripening was accelerated over control fruits by 2 to 3 days depending on cultivar
and color development was more uniform in ethylene preripened fruit. Ethylene treatment
has been used commercially in the U.S. to ripen Florida mangos. 232
Mango fruits can be dipped in ethephon solutions to enhance ripening. 231 •251 •291 Effective
concentrations of ethephon ranged from 480 to 5000 ppm a.i. A solution temperature of
26oC during ethephon treatment was used, and following removal from the solution, the
fruits were held at 21°C. As with ethylene treatment, a warmer temperature accelerates
ripening. A sticker adjuvant was added to the solution to improve coverage. Immersion
times ranged from 1 to 2 min up to 10 min. As long as the fruits were mature, the quality
of ethephon-treated fruits was considered equal to that of naturally ripened fruits. A regis-
tration for this use of ethephon has not been issued.
Postharvest dips of mature fruit for 2 min in 2,4,5-TP at concentrations of 1000 and 1500
ppm followed by draining, drying, and storage at 18 to 29.5oC and 45 to 85% relative
humidity for 10 days showed that this auxin treatment retarded color development. 249 It was
suggested that such treatment could be used to extend the storage life of mangos. However,
ripening changes continued in the fruits, and the combination of green, but ripe, fruit was
deemed less than satisfactory. No commercial use has been made of this approach.
VI. Musa spp. (BANANA, PLANTAIN)
A. Propagation
The dessert banana and plantains generally are propagated vegetatively by corm or rhizome
divisions called suckers. Growth regulators are not used. Difficulties in seed germination
in breeding programs have been noted, but growth regulators are not used to enhance
germination of the hard-coated seeds.
The upright growth upon which fruit is borne is called a pseudostem. A pseudostem arises
from a sucker which develops from the corm of a maturing fruiting pseudostem. Depending
on cultivar and planting density, two to four suckers or more can arise. The competition
among the developing stems may limit yields. Thus, it is desirable to reduce the sucker
population. In Australian banana plantations, 2,4-D at the rate of 10 to 15 me of a l% a.i.
solution is injected into the base of a pseudostem near the growing point to destroy it.
Desuckering, a process of thinning the sucker population, is carried out by pouring about
4 me of a l: 16 solution of 50% 2,4-D amine into the vase formed by the emerging leaves. 295
Injection of kerosene is as effective, however. 294
C. Flowering
The number of leaves preceding the development of flowers is constant for different
cultivars and flowering can be anticipated by close observation to ascertain the appearance
of the last leaf. The flower cluster develops from the apical meristem at the base of the
pseudostem at ground level, differentiating female to hemaphroditic flowers to male flowers
acropetally in the axils of fleshy bracts. The number of flower-bearing bracts is determined
by genetic traits, the environment, and hormonal regulation. 301 There is potential to modify
numbers of female flowers per cluster and number of clusters hormonally; 301 however, no
definitive results have been published.

The number of fruits which can be set is limited by the number of female and herma-
phroditic flowers formed. Parthenocarpic development is the rule. The effects of growth
regulators have been studied to try to improve fruit length and diameter.
Since nearly all bananas mature their bunches in about 3 months under tropical conditions,
any efforts to increase final fruit size must relate to increasing growth rate rather than
extending the time of development. 313 However, gibberellins may alter fruit length 302 •309
and, by delaying maturation, 303 •310 also extend the time period for development.
Gibberellic acid 4 + 7 (as ProGibb 47®) has been applied as a spray to emerging clusters
and found to increase fruit length slightly. 302 Promalin® increased yields as reflected in
greater bunch, hand, and finger fresh weight as well as longer and fatter fruits. 302 To date,
no commercial application has been made of this information, and other reports indicate
gibberellin application had little effect. 312
E. Fruit Ripening
Banana is well known as a climacteric fruit, responding to ethylene by characteristic
changes in color and firmness as well as increase in soluble solids, total sugars, and reducing
sugars. Indeed, the rapidity of ripening during shipping has stimulated research aimed at
delaying ripening. 305 •307 •308 Kinetin 316 and 2,4-D 314 delay degreening but only gibberellin has
been used commercially to delay the climacteric and color development. 315
Gibberellins have been shown to delay color development, 298 ·304 and one formulation (ICI
Regulex, 1% a.i. GA 4 + 7 ) is used commercially in Jamaica under defined conditions to
lengthen the green life of exportable bananas. 303 •304 Fruit must be graded by maturity as thin,
immature fruit is normally less responsive to degreening procedures and will not require
treatment while fuller fruit should be treated. A fresh 10-ppm solution is prepared in a
suspension of benomyl fungicide each day. A volume of 45 e is used as a dip for up to 100
13.8-kg boxes of packed hands before the solution needs to be renewed.
Green fruits are ripened by wholesalers in special ripening rooms under controlled con-
ditions. For rapid ripening, ethylene is injected into the rooms at the rate of 15 to 30 e/28.3
m 3 • The fruit is pre warmed to a pulp temperature of 20°C and held at an air temperature of
20°C for the first 24 hr of exposure to ethylene. Subsequently, as fruit begins to develop
color, the room temperature will be dropped lower, with final temperatures of 13.5 to 16°C.
A high relative humidity is required: 90 to 100% until colored with 90% recommended after
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CARRIAGE BY BEARERS.
Out in the country there are practically no roads, as we understand

roads. It is necessary to cultivate every inch of available ground, and
the farmer begrudges anything taken from the fields for the paths,
which are but a foot or two wide. It is easy to understand that, under
such conditions, the almost universal mode of passenger transit is by
chairs and bearers. The narrowness of the paths is a source of
trouble. When two parties of bearers approach each other, there is
much shouting to induce one or other to return and make way; but
when both come on, one has to get off, or be pushed off, into the
swamp by the sides. When one is a foreigner his portion is invariably
the swamp.
The bearers are patient, much-enduring people, who do their work
thoroughly and without complaining, in the face of mud, and rain,
and difficult roads. They will carry a traveller from twenty to twenty-
five miles a day. When a lady occupies the chair the curtains are
rigidly closed. It would be at the risk of her life to travel in an open
chair. There is much etiquette connected with the getting in and out
of chairs, which wise travellers never neglect. The photograph is of a
lady’s chair.
CARRIAGE BY BEARERS
A TRAVELLER ARRIVING
AT AN INN
IN MANCHURIA.
There are various ways of carrying a traveller’s baggage.

Sometimes it is slung in the centre of bars and carried as the
traveller’s own chair is carried. More often a package is slung at
each end of a bar, which is placed across the shoulders of a coolie.
Constant change of shoulder is necessary, and the stopping to make
this change becomes a serious matter in a journey of any length. It is
trying work, and the shoulders of the coolies generally show it by the
callositis produced by the constant carrying of heavy burdens. The
illustration shows Mrs. Bishop’s baggage arriving after a day’s
journey.
A TRAVELLER ARRIVING
AT AN INN
IN MANCHURIA
CARRIAGE
OF
MERCHANDISE.
It will be seen that two coolies, by means of these bars, can carry a
great weight—as much as two hundred pounds is carried between
them—and they will cover with this weight twenty to twenty-five miles
a day. Chair-carriers will, with the attendant luggage-carriers, cover
as much as twenty-five miles, but their burdens are less heavy.
CARRIAGE
OF MERCHANDISE
THE MODE OF CARRYING
OIL AND WINE.
In wicker baskets lined with oiled paper of extraordinary toughness,

which is much used everywhere. The oil is obtained from various “oil
seeds,” the tough paper by macerating bamboo. Beneath the basket
will be noticed a long cylinder. This is the coolie’s purse, in which he
carries his “cash,” the small copper or brass coin of the country,
which is of such small value that nine pounds weight of copper cash
is only worth one English shilling.
THE MODE OF CARRYING
OIL AND WINE
WHEELBARROW TRAFFIC
ON THE
CHENGTU PLAIN.
This Chengtu Plain, with its 2,500 square miles of country and
4,000,000 population, is perhaps the best cultivated and most fertile
spot in the world. It owes its fertility to the work of two engineers,
who, more than two thousand years ago (250 years B.C.), designed
and carried out the most perfect system of irrigation. They were Li
Ping, the father, and his son, and are familiarly known to-day as the
first and second gentlemen of China. The land bears four crops in
the year. With all this produce and population, the traffic is
enormous, and it is mainly carried on by means of wheelbarrows,
which are so contrived, by placing the wheel in the centre and
platforms at the side and behind it, as to enable one man to wheel
five hundredweight with ease. The narrow roads of the plain are
covered by an almost endless procession of these wheelbarrows,
which are often preceded by one man pulling in addition to the man
behind.
WHEELBARROW TRAFFIC
ON THE
CHENGTU PLAIN
THE WHEELBARROW
OF NORTH CHINA.
This is another form of the same baggage-carrier which is in use all

over the Empire. It is much larger than that in use on the Plain of
Chengtu, but is constructed on the same principle; by means of it
one man can wheel as much as half a ton. It is a vehicle well
adapted to the narrow roads of the country.
THE WHEELBARROW
OF NORTH CHINA

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Textbook Plant Growth Regulating Chemicals Volume 2 First Edition Nickell Ebook All Chapter PDF

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Plant Growth Regulating Chemicals -

Volume 2 First Edition Nickell

Plant Growth Regulating Chemicals - Volume 1 First

Brassinosteroids Plant Growth and Development Shamsul

Plant Genome Diversity Volume 2 Physical Structure

Plant Health Under Biotic Stress Volume 2 Microbial

Identifying and Regulating Carcinogens First Edition

Weed Physiology - Volume 2- Herbicide Physiology First

Wood Adhesives Chemistry and Technology Volume 2 First

Revival: Alcohol Interactions with Drugs and Chemicals

Louis G. Nickell, Ph.D.

Boca Raton London New York

Reissued 2018 by CRC Press

© 1983 by CRC Press, Inc.

No claim to original U.S. Government works

Library of Congress Cataloging-in-Publication Data

Plant growth regulating chemicals.

Includes bibliographies and index.

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

ISBN 13: 978-1-315-89664-9 (hbk)

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

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

TROPICAL FRUIT AND BEVERAGE CROPS

D. P. Bartholomew and R. A. Criley

II . Ananas comosus (L.) Merr. (Pineapple) ....................... . ......... . ........ 3

IV. Coffea arabica L., C. canephora L. (Coffee) .. ......... .. ...................... 13

V. Mangifera indica L. (Mango) ................................ . .. . .. . ............ 15

VIII . Theobroma cacao L. (Cacao) .............. ..... ............................... . 22

Acknowledgments ....... .. . . .. .... ..................... .. . . . ...... ................ ..... . 23

References ................ .. .. ...... .............. . ...... .... .. .... ................... . . . 23

II. Ananas comosus (L.) Merr. (PINEAPPLE)

C. Induction of Flowering (Forcing)

2. Absorption and Mechanism of Action

Acetylene and calcium Australia Smooth Cayenne 48, 87

2 ,4-Dichlorophenoxy- India Giant Kew (Smooth 76

Although Gowing hypothesized that NAA induced flowering in pineapple by acting as an

3. Plant Susceptibility to Forcing

4. Effects of Forcing on Fruit and Propagule Production

5. Forcing as a Commercial Practice

III. Carica papaya L. (PAPAYA)

D. Fruit Set and Development

IV. Coffea arabica L., C. canephora L. (COFFEE)

D. Fruit Set and Development

Evaluation of new products will continue. The application of sodium mono-nitroguaiacol

V. Magifera indica L. (MANGO)

FIGURE I. Exampl es of commercial preparations for forcing fl owering of mango .

Manufacturer/ Analysis for

Courtesy of N.D. Bondad.

" Presence of nitrate is based on company report.

D. Fruit Set and Development

There is evidence for a role of gibberellin in fruit development with 100-ppm GA 3

E. Fruit Ripening and Postharvest Handling

VI. Musa spp. (BANANA, PLANTAIN)

D. Fruit Set and Development

Out in the country there are practically no roads, as we understand

There are various ways of carrying a traveller’s baggage.

In wicker baskets lined with oiled paper of extraordinary toughness,

This is another form of the same baggage-carrier which is in use all

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