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Plant Growth Regulators in Citriculture:


World Current Uses

Mohamed El-Otmani*1, Charles W. Coggins, Jr.2, Manuel Agustí3


and Carol J. Lovatt2

1
Department of Horticulture, Institut Agronomique et Vétérinaire Hassan II,
B.P. 18/S, 80000 Agadir, Morocco;
2
Department of Botany and Plant Sciences, University of California, Riverside,
CA 92521-0124, USA;
3
Departamento de Producción Vegetal, ETSIA, Universidad Politécnica,
Camino de Vera 14,46020 Valencia, Spain

* To whom correspondence should be addressed.


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PLANT GROWTH REGULATORS IN CITRICULTURE: WORLD CURRENT USES


Abstract
I. Introduction
II. Concepts of productivity, yield partitioning, and fruit quality
III. Use of Plant growth regulators in citrus propagation
IV. Use of plant growth regulators to alter vegetative growth
V. Use of plant growth regulators to maximize orchard productivity (fruit number and size)
A. Optimization of flowering intensity
B. Manipulation of yield
1. Fruit set
2. Fruit thinning to increase fruit size
3. Stimulation of fruit growth to increase fruit size
VI. Use of plant growth regulators to enhance fruit quality
A. Internal quality
B. Rind quality
1. Rind disorders
2. Rind color
3. Rind softening and senescence
VII. Use of plant growth regulators to reduce fruit drop and for on-tree fruit storage
VIII. Use of plant growth regulators to facilitate harvest
IX. Use of plant growth regulators postharvest to maintain fruit quality
X. Ways to ensure or increase the efficacy of plant growth regulators in citriculture
XI. Conclusion and future prospects
XII. Acknowledgment
XIII. Literature cited
XIV. Tables
XV. Appendix I: Species Names
XVI. Appendix II: Source Contact Information
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ABSTRACT
Plant growth regulators (PGRs), both naturally occurring and synthetic, are used exten-
sively in citriculture not only to help growers solve production problems, but also to increase
the market value of the crop and to improve grower returns. Yield increases are obtained via an
improvement in fruit set, particularly for shy-bearing cultivars, and/or fruit size, especially for
the «small-fruit» group of mandarins. Increased fruit number per tree is achieved indirectly
through an increase in flower number as a result of better flower initiation or by a direct effect
on fruit set. Improved fruit size is brought about directly by stimulating the growth of fruit tis-
sues or indirectly by reducing fruit number by partial inhibition of flower initiation or by sub-
sequent fruit removal. PGRs are also used to hasten maturity, to delay harvest and to maintain
fruit quality postharvest, resulting in greater profits to the grower. This article provides an
overview of current uses of PGRs in citriculture. The reported information is based on an in-
ternational survey and a review of recent technical and scientific literature published on the
subject. Procedures and techniques used to enhance the efficacy of PGRs, including surfactants
and compatibility considerations, are presented and discussed. The detailed information pre-
sented in the text is summarized in tables designed to provide a convenient reference for re-
searchers and growers alike. Included in the tables are the names of the individuals who were
the source of the information. A comprehensive list of these individuals and their addresses
follows the tables. Whereas the results of the survey indicate extensive uses of PGRs in citricul-
ture, the authors admit that the survey is incomplete and extend their apologies to researchers in
countries not represented in the review and for inadvertent omissions.

KEY WORDS: abscisic acid, abscission, alternate bearing, auxins, compatibility, cytokinins,
degreening, earliness, ethylene, flower initiation, fruit color, fruit set, fruit
size, fruit thinning, gibberellins, harvesting, internal fruit quality, postharvest
fruit quality, rind quality, rind senescence, treatment efficacy, yield.

I. INTRODUCTION
Citrus growth and development are governed by plant internal factors (genotype, en-
dogenous hormone balance), the environment in which the plant is grown (climate and soil fac-
tors, including microorganisms in the rhizosphere) and cultural practices used (planting density,
irrigation, fertilization, pruning, exogenously applied plant growth regulators). Since their dis-
covery and commercial availability, plant growth regulators have been used to manipulate plant
growth and development to the advantage of mankind by enabling growers to meet the chal-
lenge of increased demand for food of high quality resulting from the ever-increasing world
population. In citriculture, a large number of plant growth regulators have been evaluated but
only a limited number are used commercially. In this review, emphasis will be on plant growth
regulators used at the commercial scale to solve a given production problem or bring about a
biological response that would enhance the market value of the crop. However, information
concerning plant growth regulators not yet registered but for which practical and valuable re-
sults have been reported have been included. This discussion of plant growth regulator use in
citriculture is organized on the basis of the desired outcome of the management strategy and the
biological response obtained, rather than by chemical, since some materials are used to achieve
several very diverse goals. For example, 2,4-dichlorophenoxy acetic acid (2,4-D) is used in
plant propagation media and in foliar sprays which, by changing concentration and time of ap-
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plication, can be used to improve fruit size, to enhance fruitlet thinning, or to prevent abscis-
sion of mature fruit.
Some of the practices and several of the materials included in this review may not be
available to the entire worldwide citrus community or may not be of practical value in some
citrus-producing regions. The techniques and procedures used may differ considerably from
region to region and from cultivar to cultivar although the same physiological process is tar-
geted. The tables summarizing the commercial uses of plant growth regulators in citriculture
provide the reader with a worldwide overview at a glance. However, the subject is large, pro-
duction regions are numerous, and access to the information is not always possible. The authors
admit the review is less than exhaustive and apologize for their inability to include every coun-
try and every commercial use of PGRs in citriculture. Despite this limitation, this review will
hopefully provide a valuable reference for expanding the use and/or for developing new uses
for plant growth regulators in citrus production, taking into account cultivar differences, local
conditions (environment, harvest window, destination of the crop, etc.) and availability of the
plant growth regulators.
Plant growth regulating chemicals are used in citrus production around the world. They
are used in the nursery or orchard to promote or prevent vegetative shoot growth, to manipu-
late flowering, to modify fruit set and fruit growth, to regulate crop load via thinning, to ma-
nipulate the time of harvest, to improve internal fruit quality (total soluble solids, juice acidity)
and external rind quality (color, firmness), and to improve on-tree and postharvest fruit stor-
age. Other data indicate the use of PGRs to control seed germination. Laboratory uses of plant
growth regulators include media for in vitro production of virus-free plants important for the
production of trees that are used by nurseries as the source of clean budwood for propagation.
Citrus is a long-lived tree producing a high value crop on expensive land with increas-
ingly intensive management systems and costly equipment. The use of a growth regulator to
overcome tree productivity or fruit quality problems in an existing planting is usually more
economical than replanting the orchard (unless the yield-limiting factor is incurable such as
viral disease, irreversible dieback, etc.) and should result in a significant increase in orchard
profitability and grower return. In addition, there is a strong market for the well-established
citrus cultivars, developing and introducing new cultivars is a slow process. Most plant growth
regulator techniques have been developed for and are used to improve the performance of these
well-established cultivars, such as the Valencia orange, Washington navel orange, Satsuma
mandarin, Clementine mandarin, and grapefruit (species names are provided in Appendix I).
Furthermore, fresh market sales of fruit depend heavily on their appearance. Treatments that
either directly or indirectly enhance fruit appearance and quality are valued highly by produc-
ers, packers, and marketing agents.

II. CONCEPTS OF PRODUCTIVITY, YIELD PARTITIONING, AND FRUIT


QUALITY
Productivity is a key element in the success and survival of a citrus-producing enter-
prise. Productivity is limited by factors such as light interception, CO2 fixation and subsequent
metabolism, water and mineral nutrient uptake, and competition among similar or different
plant organs for photoassimilates and nutrients. Plant growth regulators have been used to im-
prove productivity in instances where factors associated with genotype, environment, disease,
or management practices limit economically viable yields. Genetic limitations to productivity of
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cultivars with otherwise desirable characteristics (such as seedless fruit) can include poor fruit
set and/or small fruit size (Krezdorn, 1965). Unfavorable environmental conditions may restrict
many of the processes related to production from flower initiation (Moss, 1969; Southwick and
Davenport, 1986; Lovatt et al., 1988) to the development of inflorescences and flowers or may
cause excessive drop of flower buds, open flowers, entire inflorescences, and developing
fruitlets (Colt and Hodgson, 1919; Krezdorn, 1965). Instances in which diseases cause exces-
sive drop of developing or mature fruit include stem-end decay, which causes summer-fall drop
of navel orange fruit (de Lima et al., 1980).
Management practices such as excess nitrogen leading to excessive vegetative growth
and low flower number, or nitrogen deficiency leading to low flower initiation (see Davenport,
1990) reduce yields. Optimum yield is a yield which is sustainable year after year and which
utilizes the full potential of the available land and the energy and nutritional resources of the
tree. This yield is a function of tree planting density, canopy development, intensity of flower-
ing, fruit set, final number of fruit harvested at maturity, and fruit growth and final size. In
young trees, a high proportion of photosynthate is allocated to vegetative development and
growth. Usually, only a few flowers and fruit are produced during the two to three years after
planting a nursery tree. Canopy development and growth continues and reaches an optimum
between five and 12 years, depending on planting density, at which time yield is at its highest
value (Boswell et al., 1970, 1975b). At this point a natural equilibrium between vegetative
growth and reproductive development is established with variations due to environmental condi-
tions and genotype of the cultivar, provided that nutrition and light interception are optimal.
Maximum fruit production for ‘Valencia’ orange in Cuba, for example, was reached at about
12 years of age when the canopy volume reached its maximum (del Valle et al., 1992). Maxi-
mum production and canopy volume were influenced by the rootstock and were both greater
for Citrus amblycarpa than C. aurantium. However, canopy productivity (kg fruit per m3 can-
opy) was greater for C. aurantium. Plant growth regulators are used to minimize seasonal fluc-
tuations in yield and to maximize energy allocation to harvestable fruit in order to increase fruit
number and fruit size rather than to enhance excessive vegetative growth, which is generally
lost to pruning.
Optimizing the use of available labor is also important in increasing the economic effi-
ciency of a citrus production enterprise. Plant growth regulators are used to hasten or delay
fruit coloring and maturation in order to shorten or extend the harvest season and permit more
efficient use of the labor force in addition to shifting harvest to get the best market price.
Fruit quality is a concept that varies according to the final use of the fruit and at what
point from orchard to consumer the fruit is evaluated. For the grower, any fruit that can be sold
at a reasonably good price is of good quality since his responsibility ends at the farm gate. For
the packer, fruit of uniform size, free of blemishes and with as few culls as possible give the
best return. For the distributor, fruit with longer shelf-life, attractive appearance, and larger
size are considered the best quality. In general consumers of fresh fruit require seedless fruit
with a high sugar to acid ratio, large size and excellent external appearance (color, shape, firm-
ness and freshness). In contrast, the orange juice industry only requires that fruit have high
juice and sugar content; fruit size and external appearance are not major factors. Parameters
important to both the fresh fruit and juice industries have been shown to respond to plant
growth regulators as described below.
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III. USE OF PLANT GROWTH REGULATORS IN CITRUS PROPAGATION


Several plant growth regulators have been tested as aids in citrus propagation and seed-
ling growth (see review by Wilson, 1983). Objectives include shortening the time of seed ger-
mination, increasing the percent germination, and enhancing the subsequent growth rate of the
seedling. The best results were obtained from soaking seeds for 12 hr in a 40 mg∙l–1 solution of
either gibberellic acid (GA3) or naphthalene acetic acid (NAA). These treatments increased
germination and subsequent seedling growth. Other trials included 2,4-D,
2,4,5-trichlorophenoxy acetic acid (2,4,5-T), 2-chloroethyl phosphonic acid (ethephon), and
indoleacetic acid (IAA). The results of our survey provided no evidence that these treatments
are currently used commercially in citrus propagation.
Commercial citrus plantings are generally made with selected scions budded on seedling
rootstocks that are selected for desired resistance to phytopthora, nematodes, and soil salt ac-
cumulations. Citrus scion cultivars, however, can be propagated by rooting of cuttings. Lem-
ons, limes and citrons root easily. Mandarins are difficult to root. Indolebutyric acid (IBA) is a
commonly used rooting aid (Moss, 1975). Other rooting aids, which have shown success on
certain cultivars, are NAA and IAA. Discussion of techniques for the rooting of citrus scion
cultivars and the use of rooted cuttings in commercial propagation are found in Halma (1947),
Erickson and Bitters (1954), Optiz et al. (1968), and Platt and Optiz (1973).
Plant growth regulators also reportedly aid propagation through improved root produc-
tion of air layers of seedless lemon, using solutions containing 1 g∙l–1 of either IBA or NAA
(Patil and Chakrawar, 1979). Success of bud grafting, increased bud growth, and shortening
the time period between bud grafting and outgrowth of the bud using plant growth regulators
were also reported (Coggins and Hield, 1968; Nauer et al. 1979a). IAA and IBA improved bud
grafting success (Coggins and Hield, 1968), whereas 6-benzyladenine (BA) and 6-
(benzylamine)-9-(2-tetrahydropyranyl-)9H-purine increased bud growth (Nauer et al., 1979a).
Growth-regulating chemicals are extensively used as a supplement to in vitro media for
propagation and regeneration of citrus. In the laboratory, 2,4-D (2 mg∙l–1) and kinetin (0.5
mg∙l–1) proved successful in callus production for regeneration of triploids (seedless plants)
from citrus endosperm (Jaskani et al., 1996). Growth regulators are also used in media for the
in vitro production of virus-free citrus plants using shoot-tip grafting (Navarro et al., 1975;
Navarro and Juarez, 1977), nucellus culture, or ovule culture (Navarro and Juarez, 1977).
Nucellus culture is used to obtain nucellar plants from monoembryonic citrus cultivars, whereas
ovule culture can be used to obtain nucellar plants from polyembryonic cultivars. Shoot-tip
grafting can be used to obtain plants free of most viruses from all citrus cultivars (Navarro and
Juarez, 1977). Nucellus and ovule culture result in plants with juvenile characteristics, many of
which are not true to type. Shoot-tip grafting is widely used in citrus for recovering virus-free
citrus plants for commercial propagation. BA and IAA are added to modified Murashige and
Skoog (1962) basal medium for maintaining the grafted plants in vitro. BA at concentrations
0.1 mg∙l–1 and higher inhibits root growth and causes the initiation of numerous buds in the root
region of the rootstock and IAA in the 1 to 10 mg∙l–1 range stimulates lateral root initiation
(Navarro et al., 1975). Using this technique, it is possible to recover cultivars free of tristeza
and psorosis viruses, stubborn spiroplasma and exocortis viroid.
Vegetative propagation of citrus rootstocks by cuttings or microcutting has recently re-
ceived much interest among researchers and nurserymen, particularly for cultivars where seed
is lacking. Rooting procedures include dipping of cuttings in NAA or IBA (Ferguson et al.,
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1985). Rapid propagation of the citremon 1452 rootstock was also obtained in vitro using 1-
cm-long stem segments with 2 buds (Mas et al., 1992). Shoot growth was enhanced when the
medium was supplemented with 0.5 mg∙l–1 BA and 0.5 mg∙l–1 NAA; however, the best rooting
was obtained with 0.15 mg∙l–1 NAA and 0.2 mg∙l–1 thiamine. Using nodal explants to propagate
citrus rootstocks, Bowman (1994) showed that cultivar has a great effect on the response to
plant growth regulators in the culture medium. For Smooth Flat Seville rootstock, the largest
number of shoots was obtained in the presence of a low to moderate level of BA (0.1-1.0 mg∙l–
1
), a moderate level of kinetin (1.0 mg∙l–1) or a moderate to high level of
6-dimethylallylaminopurine (2iP). Yuma citrange rootstock nodal explants produced a larger
number of shoots when the medium included a low to moderate level of BA (0.1-1.0 mg∙l–1), a
high level of kinetin (5.0 mg∙l–1), or a moderate level of 2iP (1.0 mg∙l–1). A high concentration
of BA (5.0 mg∙l–1) inhibited shoot proliferation in both rootstocks. Rooting occurred when
NAA was added to the medium at 0.01 mg∙l–1.

IV. USE OF PLANT GROWTH REGULATORS TO ALTER VEGETATIVE GROWTH


In the nursery, early attempts to increase the size of nursery citrus plants generally were
unsuccessful and results often were opposite to the objective sought (Wilson, 1983). For exam-
ple, one major objective was to increase the growth rate of the main stem of the seedling in
order to obtain a greater stem diameter for earlier grafting and budding, thereby shortening the
time needed to produce a budded tree. Use of GA3 increased stem length at the expense of stem
girth (Abdalla et al., 1979) through an enhancement of photoassimilate translocation to the
growing apex (Ben-Gad et al., 1978). Recently, Khattab and Guindy (1996) reported that GA3
sprayed at 200 mg∙l–1 onto sour orange seedlings increased plant height and girth and produced
plants ready for budding 8 months earlier than untreated seedlings.
Attempts have also been made to alter the branching habit of nursery trees (Abdalla et
al., 1978, Hassaballa, 1979) but the results have not lead to any commercial application. A
common practice to force the growth of the scion bud is to bend down or remove the root-
stock’s seedling stem above the scion bud. The terminal bud (apical meristem) of the rootstock
produces the auxin IAA, which exerts an inhibitory effect on the development of scion bud.
Once the rootstock shoot is bent or removed, growth of the scion is enhanced. However, this
practice can also enhance the growth of lateral and basal buds of the rootstock, which is unde-
sirable. Because removal of these basal buds by hand is costly and time consuming, a commer-
cial product (TreHold®) that contains the auxin NAA (as ethyl-naphthaleneacetate) at 1.15%
was developed to inhibit trunk sprouting of citrus rootstocks (Rouse, 1994). This compound is
used in Florida to prevent axillary bud growth of citrus in the nursery as well as to prevent
sprouting of nonbearing (Knapp, 1996) field-grown trees (Table 1). TreHold® can be used to
control dormancy of individual buds without affecting adjacent buds. It is recommended that
TreHold® be applied to the trunk as a thorough spray or a light brush application (Knapp,
1996). Excessively heavy application may cause tree damage. Under Florida conditions, Tre-
Hold® should not be applied after September 1 since it might inhibit sprouting that would be
desired for tree recovery after a freeze.
In Japan, benzyladenine is used in the greenhouse production of wase ‘Satsuma’ man-
darins. BA is applied at 300 g∙ha–1 in sufficient water to give full tree coverage immediately
following winter heating of the greenhouse in order to accelerate vegetative and flower bud-
8

break. A surfactant is added to the spray solution at 0.075% (I. Iwagaki, personal communica-
tion, address provided in Appendix II).
Citrus trees are hand- or machine-pruned. In many places pruning is performed almost
every year, particularly for cultivars such as sweet oranges, mandarins, lemons and grapefruit,
for which regrowth is very rapid, particularly in warm climates. This operation is very costly
and time consuming. Numerous growth regulators have been tested in an attempt to alter as-
similate partitioning in favor of fruit yield at the expense of vegetative growth with the ultimate
goal of reducing production costs and increasing grower returns. Two mini-reviews on the use
of growth retardants in citriculture have been published (Monselise and Goren, 1984; Harty
and van Staden, 1988a). Gibberellins (Iwahori and Oohata, 1981; Dhillon and Sidhu, 1992;
Deng and Zhen, 1996) and auxins (El-Otmani et al., 1996a) promote vegetative growth when
used at low concentrations. In addition, tree vigor was correlated with high IAA and low ab-
scisic acid (ABA) in citrus, whereas dwarfing is characterized by the reverse (Bertling and Lo-
vatt, 1996). Citrus trials in Florida (Phillips and Tucker, 1974) and California (Boswell et al.,
1975a) demonstrated that NAA reduced sprouting of lemon trees. Early research in the field of
vegetative growth inhibition in citrus also dealt with compounds such as maleic hydrazide,
morphactins and ammonium ethyl carbamoylphosphonate, which reduced tree growth by caus-
ing dieback of young shoots or injury to apical meristems rather than having a direct effect on
internode length (Harty and van Staden, 1988a). Other retardants included in earlier trials were
(5-hydroxycarvacryl)trimethyl chloride piperidine carboxylate (Amo-1618), (2-
chloroethyl)trimethyl ammonium chloride (CCC), and succinic acid-2,2-dimethylhydrazide
(SADH, Alar, B-9) which were believed to interfere with the growth-promoting effect of gib-
berellins in plants (Cathey, 1964). Two inhibitors of the effect of auxins on growth, benzothia-
zole-2-oxyacetate (BTOA) and triiodobenzoic acid (TIBA), were also tested on citrus. Results
from these trials were inconsistent, often contradictory, and confusing. In some instances re-
duced vegetative growth with enhanced flowering was obtained; in other situations the opposite
results were observed; and in other trials no treatment effect was obtained. Situations where a
transient inhibition of growth was followed by vigorous growth were also reported in the litera-
ture (Harty and van Staden, 1988a).
Results of several experiments demonstrated that the triazole derivatives inhibited gib-
berellin biosynthesis and, thus, vegetative growth (Dalziel and Lawrence, 1984). The triazoles
included α-(1-methylethyl)-α-[4-(trifluourmethoxy)phenyl]-5-pyrimidinemethanol (flurprimi-
dol), (E)-(RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1H-1,2,4-triazol-l-yl)pent-1-en-3-ol (uni-
conazole) and (2RS,3RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(lH-1,2,4-triazol-l-yl)pentan-3-ol
(paclobutrazol). Uniconazole and paclobutrazol have been widely tested on fruit trees including
citrus (Harty and van Staden, 1988b; Swietlik and Fucik, 1988; Delgado et al., 1996; Peng et
al., 1996). They both reduce canopy growth by a reduction in internode and shoot length and
by a reduction in the production of new shoots. However, no commercial use for either in cit-
rus production has evolved likely due to negative effects on fruit size and yield (Swietlik and
Fucik, 1988) and to the variation in their efficiacy due to the strong influence of climatic condi-
tions (Harty and van Staden, 1988b).
Winter cold hardiness is associated with cessation of shoot growth. Therefore, it should
be possible to induce cold hardiness with growth regulators. Growth regulators such as NAA
(Nauer et al., 1979b), mepiquat chloride (1,1-dimethyl piperidinium chloride) (Peynado et al.,
1981; Wilson 1983) and SADH and CCC (Young, 1971) were tried. Results were variable due
9

to shoot physiological stage at the time of application or because the plant growth regulators
had adverse effects such as prolonging arrested growth well into spring months (Yelenosky,
1985). A more recent publication (Yelenosky et al., 1995) reported that paclobutrazol can be
useful for dwarfing the trees but does not increase tolerance to flooding or freezing.
In conclusion, because of the many inconsistent results obtained from trials using
growth inhibitors on bearing trees of citrus, no commercial use of growth inhibitors is known
to us other than 1.15% NAA (TreHold®) to prevent sprouting of buds on trunks of rootstocks.
Contradictory results can be attributed to climatic factors, stage of organ development at the
time of application, the technique (soil drench, foliar spray, trunk injection) used in the ex-
periments, the concentration applied or the cultivar investigated. In addition, use of these com-
pounds can have side effects such as loss of hardiness. Similarly, we are not aware of the ex-
tensive use of plant growth regulators to specifically enhance vegetative growth of citrus as a
commercial practice. Enhanced vegetative growth by application of GA3 is a secondary effect
obtained as a result of reduced flowering due to GA3 application to inhibit flower formation
(Dhillon and Sidhu, 1992). This strategy is used to increase fruit size of ‘Satsuma’ mandarins in
New Zealand (A. Harty, personal communication; address provided in Appendix II) (Table 1).
In addition, BA is used on greenhouse-grown wase ‘Satsuma’ mandarin trees in Japan at 300
g∙ha–1 as a full tree coverage to accelerate vegetative and flower budbreak immediately follow-
ing winter heating (I. Iwagaki, personal communication; address provided in Appendix II).
This promotes earlier flowering and vegetative growth resulting in earlier fruit maturation than
in untreated trees (Table 1). A final related use is that of adding 2,4-D (isopropyl ester at 0.25
g∙l–1) to pesticide oil sprays used on citrus in California to prevent leaf drop (Coggins, 2000)
(Table 1).

V. USE OF PLANT GROWTH REGULATORS TO MAXIMIZE ORCHARD


PRODUCTIVITY (FRUIT NUMBER AND SIZE)
A. Optimization of flowering intensity
Flowering, like all other physiological processes in plants, is under the control of culti-
var genes, the environment and cultural practices. Most citrus cultivars, under optimal condi-
tions flower profusely, particularly in the spring of the «on-crop year» in alternate- or bienni-
ally-bearing cultivars, and it is common to find more than 100,000 flowers per tree (Erickson
and Brannaman, 1960; Monselise, 1977; Agustí et al., 1982). Depending on the cultivar, 90%
to 99% of the flowers abscise and final set is frequently less than 1% (Agustí et al., 1982;
El-Otmani et al., 1992). During the «off-crop year» flowers are rather scarce. Plant growth
regulators have been tested for the purpose of regulating tree production either by reducing
vegetative growth and enhancing flowering when an «off-year» is predicted or by enhancing
vegetative growth and reducing flower initiation and development when an «on-year» is antici-
pated. However, the results have not been consistent.
Since Monselise and Halevy (1964) reported that ‘Shamouti’ orange flowering can be
inhibited by GA3, this compound has been tested on most citrus cultivars with similar inhibitory
effects having been found [see reviews by Harty and van Staden (1988a) and Davenport
(1990)]. Conversely, inflorescence production increased with increased concentration of the
triazole compounds (gibberellin biosynthesis inhibitors) (Delgado et al., 1986a, 1986b; Harty
and van Staden, 1988b). Properly timed applications of GA3 can cause the reversion of floral
buds to vegetative shoots (Guardiola et al., 1982; Lord and Eckard, 1987). Applications must
10

be made prior to irreversible commitment to flowering, which corresponds approximately to


sepal formation by the apical flower (Lord and Eckard, 1987). It is noteworthy that the period
of tree responsiveness to GA3 application to inhibit flowering is wide with two peaks of sensi-
tivity (Guardiola, 1981; Guardiola et al., 1982; García-Luís et al., 1986). The first peak coin-
cides with that of flower induction, in which a flowering promoter probably is translocated
from the leaves to the buds (Ayalon and Monselise, 1960; Moss, 1976; Agustí et al., 1981a).
In subtropical climates of the northern hemisphere induction occurs during the period from No-
vember through January. For sweet oranges and some mandarins induction occurs early in this
period (Guardiola et al., 1982), whereas for ‘Satsuma’ mandarins it occurs later (Iwahori and
Oohata, 1981; Guardiola et al., 1982). The second peak of sensitivity takes place during bud-
break coinciding with the differentiation of flower primordia (Goldschmidt and Monselise,
1972; Iwahori, 1978; Guardiola et al., 1982).
Suitable concentrations for practical use are GA3 at 25 mg∙l–1 for ‘Navelate’ sweet or-
ange with the application in mid-December and for ‘Washington’ navel with the appropriate
time of application being mid-November in Spain (Guardiola et al., 1977). Optimum concentra-
tions are 100 mg∙l–1 applied late January for ‘Satsuma’ mandarin in Japan (Iwahori and Oohata,
1981), 25 mg∙l–1 applied in early winter (mid-June) for ‘Satsuma’ mandarin in New Zealand
(A. Harty, personal communication; address provided in Appendix II), and 35 mg∙l–1 applied in
mid-December on ‘Tahiti’ lime (Davenport, 1983). Reductions in flowering close to 50% in
respect to the untreated trees were obtained in most cases. Agustí et al. (1981b) found that GA3
as low as 10 mg∙l–1 applied in late November reduced flowering 40% in ‘Clementine’ mandarin
irrespective of the date of harvest (i.e., the presence of the crop was not inhibiting flowering)
(Agustí, 1980; Guardiola et al., 1982). This low GA3 concentration reduced flowering by 37%
and 70% when it was applied to ‘Satsuma’ mandarin trees in mid-December (first peak of sen-
sitivity) (García-Luís et al., 1986) and late December (peak of greatest sensitivity) (Guardiola et
al., 1982), respectively. However, the duration of the peak of greatest sensitivity is very short
and treatments have to be applied before the developing shoots are more than 1 mm in length
(Guardiola et al., 1982), which corresponds to the stage of full sepal development of the apical
flower (Lord and Eckard, 1987). Between the two peaks of response lies a period of low sensi-
tivity (Guardiola, 1981; Guardiola et al., 1982; Lord and Eckard, 1987), during which a sig-
nificant inhibition of flowering can only be obtained either by increasing the concentration ap-
plied (Guardiola et al., 1977; Cottin, 1989), or by repeated applications (Deidda and Agabbio,
1977; Guardiola et al., 1977).
Another difficulty in planning when to make the application is that the buds on a tree are
not all at the same stage of development, and therefore would not be equally affected by the
treatment. Vegetative growth stimulation as a result of this treatment has been reported even if
an effect on flowering was absent (Goldschmidt and Monselise, 1972; Agustí, 1980), which can
be beneficial on declining trees. GA3 concentration in the spray solution should be calculated in
relation to the degree of inhibition sought as shown in Table 1 for New Zealand where fruiting
trees receive less GA3 than young trees.
In addition to GA3, GA4/7 (Goldschmidt and Monselise, 1972) and 99% pure GA4
(Tamim et al., 1996) also proved inhibitory to citrus flowering. Attempts to promote flowering
using growth retardants that are reported to inhibit synthesis of gibberellins (CCC, SADH, and
paclobutrazol) have not always given conclusive results [see reviews by Harty and van Staden
(1988a) and Davenport (1990)]. More recently, the effect of paclobutrazol on flowering was
11

investigated in Israel. Greenberg et al. (1993) reported that supply of this compound by foliar
spray or soil application during fall shifts the balance of shoot types towards the pure leafless
inflorescences, which is just the reverse of GA3 which pushes the balance towards vegetative,
flowerless shoots (see above). Differences in growth regulator concentration, tissue/organ age
or stage of development, tree fruit load, cultural practices (irrigation, fertilization, pruning),
climate, and method of application are all factors that may have contributed to the confusing
and contradicting results reported earlier with the use of these compounds.
Recent research in Mexico and Cuba has attempted to shift the spring flowering to a
later date so that the resulting harvest falls in a low production period that brings better return
to the grower (Ortiz et al., 1993; Urrutia, 1993; Vargas, 1993). In particular, for ‘Washington’
navel orange, attempts were made to reduce spring (February) flowering by applying GA3
(25-40 mg∙l–1) during November through February and to enhance summer flowering using
ethephon sprays (600 mg∙l–1) in May (Ortiz et al., 1993). The inhibitory effect of GA3 was ob-
tained but the ability of the ethephon treatment to increase summer flowering was not consis-
tent. In Mexico, ‘Persian’ lime prices are best in winter. Three GA3 applications (40 mg∙l–1) in
June through August followed by an ethephon spray (100 mg∙l–1) in October results in a better
flowering and fruit set in December promising a better winter crop the following year (Vargas,
1993). ‘Persian’ lime tends to ever-bear in a tropical climate, and these treatments have the
benefit of inhibiting summer-fall flowering and promoting late fall flowering for a better winter
yield. Low ethephon concentration gave no benefit, but flowering was significantly enhanced
with 500 mg∙l–1 ethephon (Vargas and Espinoza, 1993).
The implication of auxins in the process of flowering has been reported (Zeevart, 1978).
Application of 2,4-D at 12 mg∙l–1 in mid-November to mid-December to ‘Navelate’ sweet or-
ange reduced flowering by approximately 30%, which is comparable to the level of inhibition
obtained when GA3 was used at the same period at 100 mg∙l–1 (Guardiola et al., 1977). Greater
concentrations (18 mg∙l–1) did not provide any additional benefit (Agustí, 1980) and lower rates
(7.5 mg∙l–1) had no inhibitory effect (García-Luís et al., 1986).
It should be pointed out that 2,4-D sprays are widely used to reduce mature fruit abscis-
sion (see Section VII below), particularly when the crop will be picked late in the season. Such
applications may coincide with early stages of bud development of the spring flush and ther-
fore, may cause malformed vegetative and reproductive structures (Coggins, 1992, 2000).
Abscisic acid (ABA) may also be involved in the regulation of flowering but little work
has been done to elucidate its effectiveness (Young and Cooper, 1969; García-Luís et al.,
1986).
The role of cytokinins in the regulation of flowering in citrus has received little attention
although a correlation between endogenous cytokinin levels and budbreak and growth has been
reported [see reviews by Davenport (1990) and El-Otmani et al. (1995)]. Using planted ‘Tahiti’
lime marcotts, Oslund and Davenport (1987) found that root-produced cytokinins were likely
responsible for enhanced vegetative shoot production. Davenport (1990) also argued that a lack
of flowering (off-bloom) when trees are bearing the heavy «on-year crop» for severely alternate
bearing cultivars, for example, the mandarin group, is likely a result of a lack of cytokinins,
which in turn results from a lack of carbohydrates and other nutrients necessary for active root
growth and, hence, for production of cytokinins. No practical uses of cytokinin in optimizing
flowering in citrus are known to the authors, however.
12

In conclusion, although gibberellins appear to be the only plant growth regulators that
consistently inhibit flowering, their use as a commercial strategy for reducing flower number to
increase fruit size or in an expected «on-year» to reduce alternate bearing is not widespread.
Uncertainty about climatic conditions, particularly temperature extremes that might prevail the
following spring or summer and cause poor fruit set or excessive fruit drop resulting in low
yield, is considered too great a risk by many growers. However, the use of GA3 in areas where
species such as limes are ever-bearing, in conjunction with ethephon, may lead to important
breakthroughs enabling growers to shift harvest to a time of high demand, high prices, and high
grower return.

B. Manipulation of yield
Yield (kg fruit∙tree-1) includes both fruit number and fruit size. Fruit number is a func-
tion of flower intensity and fruit set. Fruit size is a function of cell division and cell enlarge-
ment processes. The number of fruit that set and persist to harvest influences fruit size. Final
fruit size is an important component of net return to the grower. Thus, an increase in fruit
number does not always mean an increase in grower profit.
Citrus fruit growth is a continuous process. Both fruit weight and fruit diameter are rep-
resented by a single sigmoidal growth curve (Bain, 1958; El-Otmani and Coggins, 1985; Guar-
diola and Lázaro, 1987). Development of the ovary into a mature fruit proceeds through three
stages: cell division, cell enlargement, and maturation (Bain, 1958). During Stage I, cell divi-
sion predominates. Cell division is essential for the formation of the ovary during flower initia-
tion and continues past petal fall to approximately mid-June to late July (experimentally deter-
mined to be the time of maximum peel thickness for both ‘Valencia’ and navel orange in Cali-
fornia) (Lovatt, 1999). At this time cell division, which occurs in all the tissues, is completed
except in the flavedo. Thus, during the period encompassing fruit set and June drop, fruit
growth is dominated by cell division. Stage II is the period of maximum fruit growth occurring
over the following three months for the early maturing cultivars (e.g., ‘Satsuma’ mandarin) and
the next seven to ten months for the late maturing cultivars (e.g., ‘Valencia’ orange). This stage
of fruit development is dominated by cell differentiation and cell enlargement. The juice vesi-
cles and locules increase in size with the uptake of water, and the flavedo continues cell divi-
sion, whereas the cells of the albedo must expand and stretch to accommodate the growth of the
locules. Dry matter and water accumulate in the vacuoles at a high rate (Guardiola, 1992).
Fruit that persist on the tree through the June drop period are not likely to abscise in the ab-
sence of pest damage, physiological disorders such as splitting, or stress such as water deficit
and excess temperatures. Stage III is the period of fruit maturation. Fruit continue to increase in
size predominately by cell expansion, but at a slower rate. Peel thickness may increase in some
cultivars (Guardiola, 1992) and puffiness may occur, particularly in the mandarin types. Pre-
harvest drop of mature fruit may start.

1. Fruit set
In citrus the process of flower formation is short relative to deciduous trees (Monselise
and Goldschmidt, 1982), but the process of fruit development is long and takes at least six
months for early cultivars such as the ‘Marisol Clementine’ mandarin or ‘Satsuma’ mandarin
and at least 11 months for the late cultivars such as the ‘Valencia’ orange. Fruit set (early drop)
is the critical phase in the transformation of the flower into a fruit for increasing grower return
13

(Lovatt, 1999). It is governed by many factors such as carbohydrates (Bower et al., 1990;
Lovatt et al., 1992), nitrogen (Lovatt et al., 1992) and plant growth regulators (Monselise and
Goldschmidt, 1982; Bower et al., 1990; Lovatt et al., 1992). Until the fruitlets have reached
the stage of maximum peel thickness (i.e. approx. 20-25 mm in diameter for mandarins and
approximately 30-35 mm for oranges), young fruits are subject to abscission. Up to 95% to
99% of the initial flowers drop fruit within two months of initial set (Monselise and Gold-
schmidt, 1982; El-Otmani et al., 1992). Some cultivars, known as shy-bearers, naturally set an
insignificant crop. These include most of the seedless mandarins, some mandarin hybrids and
the navel oranges. For these cultivars, application of exogenous plant growth regulators is nec-
essary for the crop to be economical. In standard cultivars, shedding of reproductive organs is
controlled by a feedback mechanism (Goldschmidt and Monselise, 1977) resulting in a very
regular yield from year to year provided that flowering and fruit set are not negatively affected
by climatic factors or diseases. In alternate bearing cultivars, this competitive inhibition mecha-
nism is less active and final yields depend more on flower number and initial set (Monselise
and Goldschmidt, 1982).
Gibberellins were shown to promote fruit set many years ago (Hield et al., 1958; Krez-
dorn, 1969). GA3 was shown to enhance early fruit growth, resulting in a delay or inhibition of
early fruit drop and an increase in final fruit set (El-Otmani et al., 1992; van Rensburg et al.,
1996). It has been demonstrated that GA3 increases sink strength in ovaries during the first few
weeks after anthesis (Powell and Krezdorn, 1977; Mauk et al., 1986). Mauk et al. (1986) also
showed that GA3 promoted foliar growth and enhanced assimilate export to the developing fruit
and García-Martínez and García-Papí (1979b) reported data suggesting that foliar-applied GA3
increased translocation of mineral elements from leaves to the growing fruit.
Application of GA3 at 10 to 20 mg∙l–1 as a foliar spray at full bloom is a standard proce-
dure in many countries for cultivars that typically have low fruit set (Table 2) (Krezdorn and
Jernberg, 1977; García-Martínez and García-Papí, 1979a; de Lange et al., 1982; El-Otmani et
al., 1992; Russo, 1996). For example, in countries producing ‘Clementine’ mandarins, use of
GA3 is considered a “must” for adequate yields. In Morocco, foliar spray application of GA3 at
10 mg∙l–1 to give full tree coverage at full bloom to petal fall is necessary to reach a profitable
fruit set and final yield (El-Otmani et al., 1992). Higher concentrations did not provide further
significant increases in yield. Lower concentrations also yielded larger fruit and a higher per-
cent export yield, but absolute export yield was best with GA3 at 10 mg∙l–1. In years when
bloom takes place during a short period of time (i.e., about 10 days), it is recommended to ap-
ply GA3 at 10 to 15 mg∙l–1 as a single application, but in years when bloom occurs over a pro-
longed period of time, GA3 is applied twice at 7.5 to 10 mg∙l–1 at early to mid-bloom and at late
bloom to petal fall (El-Otmani, 1992). This practice ensures treatment of the maximum possible
number of flowers and fruitlets. Krezdorn and Jernberg (1977) reported excessive leaf drop and
some twig dieback when concentrations exceeded 15 mg∙l–1. In Spain, Fornes et al. (1992) re-
ported that optimum GA3 concentration for fruit set in ‘Clementine’ mandarin depended on the
year, the orchard, and the cultivar. They showed that under the conditions prevalent in Valen-
cia, Spain, optimum concentration of GA3 is 5 mg∙l–1 for ‘Fina’ and 3 to 6 mg∙l–1 for ‘Nules’
Clementine mandarin. Addition of a seaweed extract (Fornes et al., 1996) or a foliar fertilizer
(Russo, 1996) to the GA3 mixture enhanced the GA3 effect and reduced fruitlet abscission. In
South Africa, foliar applications of GA3 are as low as 3 to 5 mg∙l–1 (Table 2). Spray applica-
tions are done as a full tree coverage to the point of run off. Timing of the treatment is full
14

bloom to petal fall. A wetting agent is almost always added to the spray solution to improve
uptake. The pH of the spray solution is not usually controlled except in Israel and South Africa,
where a pH of 3 to 5 is often used to maximize efficacy of the treatment (Table 2).

2. Fruit thinning to increase fruit size


Large yields do not always mean high profits. Considerable revenues are lost every
year in the main citrus areas of the world because of the production of small size fruit (Monsel-
ise, 1973; Gilfillan, 1987; Robison, 1987). Consumer preference worldwide is for large size
oranges (Gilfillan, 1987; Miller and Hofman, 1988). Large ‘Valencias’, from counts 56 to 88
(15 kg carton), have produced the highest returns during the last decade in the European and
Asian markets with a sharp reduction in price for smaller fruit (counts 113 and higher) (Gilfil-
lan, 1987; Robison, 1987). Moreover, the current demand for larger fruit by the international
fresh fruit market has resulted in an increase in the proportion of undersized fruit that go to
juice which, in some years, results in a net loss of revenue to the grower (Robison, 1987). It is
important to note that, in addition to its importance for the fresh market, fruit size is the only
attribute which can be correlated directly with all juice quality constituents (Hutton, 1989).
Fruit juice acidity (percent citric acid) and juice volume are very much affected by fruit size.
Small fruit have a substantially higher percent acid than larger fruit and total soluble solids per
fruit is considerably lower in small fruit. Reducing the proportion of small fruit harvested in-
creases grower return both for fresh fruit (because there are more fruit of commercially valu-
able export size) and for juice (because of increased juice volume and total soluble solids).
Small fruit size can be brought about by many factors, including competition, which is
particularly important for cultivars that set a large number of fruit such as ‘Satsuma’ mandarin
(Hirose, 1981), ‘Wilking’ mandarin and ‘Murcott’ tangor (Monselise and Goldschmidt, 1982).
These cultivars, as well as others such as ‘Valencia’ and ‘Shamouti’ oranges, grown under less
than optimal conditions may have a considerable degree of alternate bearing, which some years
results in relatively valueless fruit. In the «off-crop year» there may be a too large, course
rough-peeled fruit, whereas in the «on-crop year» thousands of very small fruit per tree are
produced (Monselise et al., 1981; Monselise and Goldschmidt, 1982). After extremely heavy
crops, alternate bearing can result in tree decline and even collapse in such extreme cases as
‘Kinnow’ mandarin and ‘Murcott’ tangerine trees (Jones et al., 1975; Smith, 1976). Competi-
tion for carbohydrates reserves among subsequent crops and tree roots has been postulated as
the cause (Jones et al., 1975), but results are inconclusive (Monselise and Goldschmidt, 1982
and references therein).
Competition affects final fruit size in citrus. There is an inverse relationship between
fruit size and flower number and fruit number per tree (Guardiola, 1992). Consequently,
flower and fruit thinning, both manually and chemically, have been used to improve fruit size
(Hirose, 1981; Wheaton, 1981; Guardiola et al., 1982; Zaragoza et al., 1992). Hand thinning
is costly and time consuming. Thus, chemical thinning has been the desired strategy for culti-
vars that set excessively and for minimizing the effects of alternate bearing (Iwahori, 1978;
Hirose, 1981; Monselise et al., 1981; Gallasch, 1984).
A significant increase in fruit size occurs only if fruit thinning is considerable and per-
formed sufficiently early in fruit development (Zaragoza et al., 1992). Depending on the culti-
var, thinning can lead to a serious reduction in yield (kg fruit tree-1) (Monselise et al., 1981;
Gallasch, 1988).
15

The most widely tested growth regulators used for thinning include ethephon, an ethyl-
ene releasing agent (Wheaton, 1981; Gallasch, 1984, 1988), NAA (Hirose, 1981; Monselise et
al., 1981; Wheaton, 1981; Gallasch, 1988; Guardiola et al., 1988) and ethyl 5-chloro-3(1H)-
indazolylacetate (ethychlozate; Figaron) (Hirose, 1981; Noma, 1981; Guardiola et al., 1988).
Auxins such as 2,4-D (Wheaton, 1981; Guardiola et al., 1988), 2,4,5-T (Hirose, 1981; Whea-
ton, 1981; Guardiola et al., 1988), 2,4-dichlorophenoxypropionic acid (2,4-DP) (El-Otmani et
al., 1993; Agustí et al., 1994a) were also tested but thinning was either highly variable or the
compounds caused no thinning at all. NAA, ethephon and ethychlozate were more consistent as
thinning agents but their efficacy was affected by factors such as concentration, cultivar, tim-
ing, tree factors (nutrition and water-deficit stress) and environmental conditions (temperature
and relative humidity) (Monselise et al., 1981; Wheaton, 1981; Gallasch 1984).
According to published literature and to the survey we conducted (Table 3), NAA is
used commercially on citrus, particularly on mandarins, in Israel (Monselise et al., 1981), and
in Florida in the United States (Knapp, 1996) (Table 3). In Japan NAA was used to thin fruit
until the discovery of ethychlozate as a fruit thinner (Hirose et al., 1978; Hirose, 1981) and its
secondary benefit as a maturation enhancer (Tominaga et al., 1995). For the success with ethy-
chlozate, it is important to ensure that target trees are well watered and are not under any kind
of stress (Hirose, 1981; Wheaton, 1981). To avoid over-thinning, treatments should be applied
early in the morning or late in the afternoon and only during a period in which excessive tem-
peratures (>30°C) are not common (Hirose, 1981). These precautions must also be observed
with use of ethephon 200 mg.1-1 to thin common mandarins as is done in Argentina. Fruit size
is an important factor in timing applications (Table 3).
If small fruit have little value, a large reduction in fruit number can be afforded as long
as price in the market place is based on size of individual fruit. In addition to an increase in
fruit size, advantages of thinning also include prevention of tree collapse which ensures tree
survival. In this case, even a loss in immediate profit can be justified (Monselise et al., 1981).
A compound that was recently tested as a growth regulator is the auxin 3,5,6-
trichloro-2-pyridyloxyacetic acid (3,5,6-TPA) (Agustí et al., 1994b, 1995b). It was reported to
have a great thinning effect particularly when used at concentrations >20 mg∙l–1. Intensity of
thinning was dependent upon the time of application, greater thinning is achieved when the
treatment was applied during June drop compared to later applications (Agustí et al., 1995b).

3. Stimulation of fruit growth to increase fruit size


As stated above, an ability to increase fruit size is desirable because for some cultivars,
small size fruit are of little value in the fresh market. Small fruit size is particularly common in
mandarins. Fruit of unmarketable size are eliminated at the packinghouse causing serious losses
in grower profits.
Fruit size is mainly determined by the genetic make-up of the cultivar but can be af-
fected by cultural practices, factors related to the tree condition, and climatic factors. Differ-
ences in fruit size result from differences in cell number or cell size or a combination of both.
The relative contribution of each component to final fruit size is unknown, but strategically
increasing cell number first, followed by increasing cell size provides two time frames and two
means to solve the problem. The potential for increasing fruit size by enhancing cell division
(Stage I) is a relatively undeveloped approach. Treatment with plant growth regulators such as
gibberellins and cytokinins at, or shortly after, flower opening enhances early fruit growth
16

(García-Martínez and García-Papí, 1979b; El-Otmani et al., 1992; Guardiola et al., 1993) but
does not always result in an increase in fruit weight at harvest. This early increase in fruit size
is mainly due to a transient increase in cell division in the ovary wall (Guardiola et al., 1993).
Early application of auxins increases final fruit size more consistently (Guardiola et al., 1988,
1993; Agustí et al., 1992; 1996, 1998; El-Otmani et al., 1993; Aznar et al., 1995). Results
were particularly consistent with the use of 2,4-D on sweet oranges in Israel and in California,
2,4-DP on ‘Clementine’ mandarins in Morocco, Israel and Spain, and on sweet oranges in
Spain and Israel, and 3,5,6-TPA on ‘Clementine’ mandarins in Spain and Morocco (Table 4).
The auxins 3,5,6-TPA and phenothiol are also being used with success on sweet oranges in
Spain (Table 4). The rates vary with cultivar, plant growth regulator, stage of fruit develop-
ment, and climatic region. Information regarding current application parameters gathered
through the survey is shown in Table 4. For success of such treatments, it is important to en-
sure optimum tree nutrition and irrigation and to provide full tree coverage during application.
In Israel, to increase treatment efficacy, the spray solution is adjusted to pH 4 and KNO3 (5%)
is added to the spray solution, which is usually applied at the end of May to the beginning of
June [i.e., 6-8 weeks after petal fall; prior to the time of maximum peel thickness and vesicle
and locule expansion (Erner et al., 1993)]. These treatments can be utilized in all years except
those with extremely low yields in which a significant proportion of the fruit will be too large
to be commercially acceptable.
The mode of action of these plant growth regulators has been summarized elsewhere
(Agustí et al., 1996) but it is important to note that they increase vesicle size, not number
(Agustí et al., 1992, 1996; El-Otmani et al., 1993), and that their effect on vesicle size is
through an enhancement of cell enlargement, not cell division (El-Otmani et al., 1993; Aznar et
al., 1995; Agustí et al., 1996). Developing fruit are important sinks for water and photoassimi-
lates; their sink strength is increased as a result of auxin treatment, with all fruit tissues increas-
ing in proportion. Auxin application increases fruit peduncle size through an increase in phloem
cell size (Agustí et al., 1996) consistent with increased sink strength and carbohydrate move-
ment into the fruit as reported for grapes (Skene, 1971). Efficacy of these growth substances is
greatest towards the end of the June drop period. After this period, fruit sensitivity to treatment
is reduced and higher concentrations may be necessary to obtain an increase in fruit size
(Agustí et al., 1992, 1994a; El-Otmani et al., 1993; Guardiola et al., 1993). It must be noted
that early application of these compounds reduces fruitlet growth (Guardiola et al., 1988, 1993;
Agustí et al., 1995b). Reduced fruit growth rate during the early stages of fruit development
was positively correlated with fruit abscission, particularly when fruit were treated during the
June drop period (Agustí et al., 1995b). The results indicate the importance of timing in plant
growth regulator use. When fruit abscission is increased, competition among the remaining
fruit is reduced and fruit size is increased but yield may be significantly diminished.

VI. USE OF PLANT GROWTH REGULATORS TO ENHANCE FRUIT QUALITY


The rind is the fruit component that protects and gives the aesthetic quality to the fruit.
For the fresh fruit industry, rind blemishes or disorders that alter rind texture, color, and
healthy appearance inevitably reduce fruit acceptance by the consumer and reduce its commer-
cial value. In contrast, internal composition is of prime importance to the processing industry.
Good-looking fruit which also have excellent internal quality procure premium prices in the
fresh market.
17

A. Internal quality
Juice content and taste are probably as important as color and size for fresh fruit con-
sumption of citrus. Organoleptic quality is the result of sugar and acid content and the presence
of volatiles in the juice. Nutritional quality includes sugar, acid, vitamins, etc. For fruit proc-
essing, internal quality is far more important than the external appearance of the fruit. Plant
growth regulators that can be used to manipulate these parameters in citrus are almost nonexis-
tent.
Although not a plant growth regulator, lead arsenate was used for many years in Florida
to improve fruit internal quality (Knapp, 1982). This compound causes a reduction in total acid-
ity and, consequently, an increase in the sugar to acid ratio. Wilson (1983) reported that most
organic and inorganic arsenic compounds will cause a reduction in citrus fruit acidity and that
the effect can be so pronounced that fruit become insipidly sweet. No non-arsenical compounds
have been identified that have commercial possibilities (Wilson et al., 1977b). To our knowl-
edge no compounds are currently used to lower acidity and thereby hasten legal maturity of
citrus in Florida.
In South Africa, arsenic compounds have been used commercially for many years (Gil-
fillan et al., 1976). Oranges, particularly ‘Valencia’, tend to produce fruit with high acid con-
tent. Mature trees can be sprayed yearly to reduce fruit acidity with no residual effects from the
arsenic. Acidification of the spray solution to pH 4 with sulfuric acid improved the efficacy of
the treatment, which reduced the quantity of the arsenic compound required. Current practice
calls for the addition of 150 g calcium arsenate in 100 liters of spray solution. This rate can be
reduced by two thirds if pH is lowered to a value of 5 (Table 5). Calcium arsenate is registered
for use on ‘Valencia’ orange only and its use is declining with the development of improved
cultivars of ‘Valencia’ such as ‘Delta’ and ‘Midnight’ (I. Gilfillan, personal communication;
address provided in Appendix II). The sprays are made two to six weeks following full petal
fall and it is recommended that the treatment be used alone.
In Japan, a substantial number of ‘Satsuma’ mandarin plantings are in plastic green-
houses (Tominaga et al., 1995). The plantings produce early mature fruit with green rind be-
cause of the hot and humid summer months. In open field ‘Satsuma’ mandarin groves, ethy-
chlozate is used as a thinning agent (see previous sections). Besides its thinning effect, ethy-
chlozate accelerates coloration and increases °Brix of mature fruit both in the field (Tominaga
and Daito, 1981; Iwahori et al., 1986) and under greenhouse conditions (Tominaga et al.,
1995). Spray solution concentration should be between 67 and 100 mg∙l–1 and applied when the
fruit are at least 30 mm in diameter (Tominaga et al., 1995). Earlier reports showed that 100 to
200 mg∙l–1 of ethychlozate sprayed 40 to 50 days after full bloom was considered the effective
period for application to obtain a leaf to fruit ratio of 20 to 25 (Hirose, 1981). Current recom-
mendations are: two sprays of 400 g∙ha–1, one each at 50 to 60 days and 70 to 80 days past full
bloom (Table 5).
Granulation of fruit is a physiological condition in which the juice sacs become gelled
with little extractable juice (Bartholemew et al., 1935). It develops preharvest (Smoot et al.,
1971) as well as postharvest (Gilfillan and Stevenson, 1977). A reduction of the disorder by
GA3 sprays has been reported but this effect has not been consistently reproduced [see Davies
(1986) and the references therein].
18

To our knowledge, studies on growth regulators to specifically manipulate vitamin C


content of the fruit are nonexistent. The few studies on citrus fruit internal quality in response
to hormone treatments showed no growth regulator effect on this component (Coggins et al.,
1960b, 1960a; El-Zeftawi and Peggie, 1973; Shawki et al., 1978).

B. Rind quality
1. Rind disorders
For fresh fruit packers and wholesalers, fruit appearance is as important as some other
parameters such as fruit size since the aesthetic quality plays an important role in price deter-
mination of the fruit. In addition to the rind disorders associated with senescence, alterations
such as thick and rough peels (Erner et al., 1975), peel blemish (Agustí et al., 1988), creasing
(Monselise et al., 1976), puffing (Kuraoka et al., 1977), pitting (Almela et al., 1992), corky
(silvery) spots of grapefruit (Goren et al., 1976) and splitting (Almela et al., 1994; García-Luís
et al., 1994) may render the fruit valueless and cause heavy economic losses. Existing informa-
tion about the etiology and the physiology of peel puffing, splitting and creasing has been re-
viewed elsewhere in relation to growth regulators (Grierson, 1986; El-Otmani and Ait-
Oubahou, 1999b). A connection with the endogenous hormonal balance has been demonstrated
for peel roughness (Erner et al., 1976a) and growth regulators have provided an effective and
practical way to reduce incidence of rind disorders or at least delay their appearance (Coggins,
1969; Embleton et al., 1973; Erner et al., 1976b; Monselise et al., 1976; Zaragoza et al.,
1996).
Current uses of plant growth regulators as obtained from our survey are summarized in
Table 6. In Israel, GA3 is used at 10 to 20 mg∙l–1 to reduce the incidence of creasing and 2,4-D
plus KNO3 are used to reduce splitting. In South Africa and Australia, GA3 is used to reduce
creasing on navel oranges (Gilfillan et al., 1974). GA3 is applied when fruit are 30 to 40 mm in
diameter for oranges and 20 to 30 mm in diameter for mandarins. In all cases and for optimum
efficacy of the treatment, pH is maintained in the acidic range. In addition, optimum cultural
practices need to be ensured for best response of the trees to treatments, particularly since some
of these disorders can be exacerbated by water stress and nutrient deficiencies.
Although no treatment is used to control puffing directly since it is a disorder observed
in fruit past maturity, GA3 treatments that are used to delay peel senescence reduce puffing sig-
nificantly (Table 8; Kuraoka et al., 1977; Agustí et al., 1981, 1986; García-Luís et al., 1985).
Less rind puff was found on ‘Satsuma' mandarin when 2,4-DP was used to improve fruit size
(Agustí et al., 1995a). This was a result of improved peel firmness of treated fruit. Reduced
rind puffing was also reported for fruit treated with ethychlozate for thinning (Hirose, 1981).
Corky spot of grapefruit has been reported to cause economic losses to growers for the
fresh fruit market in Israel (Goren et al., 1976). Its incidence was substantially reduced by the
use of 2,4-D isopropyl ester applied two to four months following anthesis or GA3 applied three
months past anthesis.
Rind pitting of ‘Fortune’ mandarin can cause significant losses both on the tree (Almela
et al., 1992) and during low temperature storage (Martínez-Jávega et al., 1992). GA3 applied at
10 mg∙l–1 just before or at fruit color break reduced the percentage of affected fruit (Zaragoza
et al., 1996). Auxins such as 2,4-DP and 3,5,6-TPA used at the end of June drop reduced the
incidence of the rind pitting disorder (Agustí et al., 1995b).
19

Stem end rind breakdown involves collapse and subsequent darkening of the epidermal
tissues around the stem end of oranges and other citrus (see Grierson, 1986 and the references
therein). GA3 is a current treatment in commercial orchards of ‘Minneola’ tangelo in Florida to
reduce this disorder (Table 6).
Abnormal peel thickness often results in very rough fruit of little value and is believed
to result from the presence of excessive concentrations of natural endogenous hormones (Mon-
selise, 1978). SADH (1000-4000 mg∙l–1) and CCC (1500-2500 mg∙l–1) applied three to six
weeks past full bloom reduced the incidence of the problem but no commercial application of
these growth regulators on citrus is known to us.
To control creasing, it appears that besides the concentration of the plant growth regula-
tor, timing of application is also critical (Pienaar and Gilfillan, 1996). Dosage is affected by the
scion-rootstock combination (Table 6; Pienaar and Gilfillan, 1996). Timing is crucial as it can
have secondary effects on the quality of the fruit and particularly on rind color at harvest. In
fact, in South Africa when GA3 spray was applied three to four weeks prior to color break (late
March to early April) and also at color break, creasing incidence was significantly reduced at
both timings but fruit color development was adversely affected (Gilfillan, personnel communi-
cation). When GA3 was applied at mid-May, creasing was controlled but flowering of the fol-
lowing season was severely reduced. When GA3 was applied in January (when fruit were 30-40
mm in diameter), good crease reduction was obtained with a very small delay in fruit coloring.
Application in December was less efficient in controlling creasing whereas a treatment in Feb-
ruary was very efficient but color development was retarded.

2. Rind color
In citrus fruit, rind color is important for its aesthetic value as consumers of fresh fruit
prefer brightly colored fruit and are ready to pay a high price for them. Color development
results from changes in carotenoid (Eilati et al., 1969; Stewart and Wheaton, 1972; El-Zeftawi,
1978; Agustí et al., 1988) and chlorophyll levels (El-Zeftawi, 1978; Agustí et al., 1988) with
concomitant changes in plastids (Thomson et al., 1967; El-Zeftawi and Garrett, 1978; Ljubesic,
1984). Environmental, nutritional and hormonal factors play an important role in the mecha-
nisms controlling citrus fruit color development (see review by Goldschmidt, 1988).
Plant growth regulators have been used to manipulate fruit color development in two
ways: 1) by enhancing color change for early harvest, and 2) by delaying color change con-
comitant with a delay in rind softening and a reduction in rind disorders (see previous sections)
for late harvest and maintaining quality. The combination of the two strategies makes it possible
to widen the harvest window. This gives the industry better control over when and how many
fruit are harvested versus the market price during the harvest period.
Citrus fruit, particularly mandarins, often reach minimum legal maturity long before
rind color develops to a level that is acceptable to the consumer (El-Otmani, 1992). When day
and night ambient temperatures are relatively high during late summer to early fall, rind color
change is severely hampered, a situation very similar to that occurring in tropical regions where
high temperatures prevail throughout the year (Reuther and Rios-Castano, 1969). Because early
in the season citrus fruit supply is low and demand is high, growers and packers degreen the
fruit to ehance color development. Although peel color is primarily a cosmetic matter, degreen-
ing improves the fruit market value and allows for earlier harvest. When fruit are to be de-
greened, fruit picking is carefully done to avoid injuring the fruit since fruit damaged during
20

harvest will succumb to disease more rapidly during exposure to ethylene and have shorter
postharvest shelf-life. In selective picking, small fruit are left on the tree for additional growth
and only fruit that have reached minimum acceptable size and have started to break color are
harvested. Degreening takes place in rooms at 20 to 25°C and 90% to 95% relative humidity
with ethylene concentrations of 1 to 10 mg∙l–1. The duration of the degreening process varies
from three to six days according to the intensity of green color in the flavedo (Table 7). Ade-
quate air circulation [about one air change per hour (Wardowski and McCornack, 1979)] in the
degreening room must be ensured to equalize conditions of temperature, humidity, ethylene
concentration and oxygen throughout the entire atmosphere and to avoid accumulation of un-
wanted gases such as CO2 and volatiles which either tend to accumulate near the floor or the
ceiling of the room if they are heavy- or light-weight molecules, respectively (Grierson et al.
1986). In fact, CO2 at concentrations>1% was reported to inhibit ethylene action (Grierson
and Newhall, 1960). Apelbaum et al. (1976) and Young et al. (1962) reported that degreening
is markedly inhibited by 5% CO2, an effect that is dependent on the relative concentrations of
CO2 and ethylene.
Ethephon enhances color development when used as a preharvest spray on oranges and
mandarins (Young et al., 1974; Knapp, 1982; Pons et al., 1992, 1996; El-Otmani et al.,
1996b). The fruit must have achieved minimum legal maturity, including the fact that coloring
of the flavedo must have started before this treatment can be effective (Knapp, 1982). It is not
widely used commercially. In Florida for example, it used to be a recommended practice to
spray ethephon at 200 to 250 mg∙l–1 on citrus (Knapp, 1982) but the current Florida spray guide
does not show such a recommendation. Two disadvantages are often associated with this prac-
tice: erratic results and heavy leaf drop (Wilson, 1983). In fact, defoliation can reach 25% to
40% (Pons et al., 1992; El-Otmani et al., 1996b); however, the leaves that drop are mostly old
and senescing leaves. In addition, because ethylene activity, and thus color development, is
temperature dependent (Wheaton and Stewart, 1973; El-Zeftawi, 1978) use of ethephon as a
preharvest spray requires experimentation under local conditions and expertise in timing appli-
cations to take into account weather conditions for the 72 hours or so following treatment.
Ethephon is used in Corsica and South Africa and is applied two to three weeks before
harvest (Table 7). Experimental studies have also been performed in Morocco (El-Otmani et
al., 1996b), Spain (Pons et al., 1992), Australia (El-Zeftawi and Garrett, 1978), Japan (Iwahori
and Oohata, 1980), and Greece (Protopapadakis and Manseka, 1992).
Leaf drop from ethephon treatment is cultivar dependent (Protopapadakis and Manseka,
1992) and chemical concentration dependent (El-Otmani et al., 1996b). Addition of calcium
salts in the spray solution reduces leaf drop significantly (Iwahori and Oohata, 1980; Perez et
al., 1992; Pons et al., 1992) but also, in some cases, reduces ethephon effectiveness (Pons et
al., 1992). It is important that optimum irrigation and fertilization practices are ensured in
groves receiving ethephon treatments since stressed trees may lose leaves much more easily
than trees in well-managed orchards (Table 7).
Use of ethylene to degreen fruit postharvest for early harvest is a more common practice
in the Mediterranean basin and other citrus-producing areas such as California (Coggins, 1992).
Fruit maturity criteria are similar to those described above and in Table 7. It is noteworthy that
degreened fruit often lose the calyx and this makes them more susceptible to decay, dryness and
shriveling. In Morocco, fruit are dipped in 4 to 8 mg∙l–1 2,4-D solution immediately after pick-
ing which significantly reduces button abscission and keeps the calyx green following the de-
21

greening process. A green button is seen by consumers as a sign of freshness and fruit without
it are considered by the consumer to be old, less attractive fruit.
Use of 2,4-D (250-500 mg∙l–1) in storage wax is a common practice for California lem-
ons to reduce decay due to Alternaria (Coggins, 1992). Use of 2,4-D postharvest is also com-
mon practice in several countries (Table 11).
Coggins et al. (1970) demonstrated that the compound CPTA (2-(4-
chlorophenylthio)-triethylamine) dramatically increased lycopene accumulation in the flavedo of
‘Marsh’ grapefruit when applied pre- or postharvest. Jahn and Young (1975) tested CPTA on
several other cultivars at harvest and following six weeks of storage and found that it had little
effect on lycopene levels when applied at harvest, but significantly increased lycopene levels
with increasing time in storage. However, fruit tends to have an intense red color which was
not acceptable for grapefruit and lemons. Enhanced color development was also observed when
‘Satsuma’ mandarin fruit was treated with ethychlozate for fruit thinning (Hirose et al., 1978;
Tominaga et al., 1995).
In contrast, GA3 delays rind color development (see previous sections) as it delays the
loss of chlorophyll and reduces the rate of accumulation of carotenoids (El-Zeftawi, 1978;
El-Zefatwi and Garrett, 1978). Delay in color development is associated with a delay in peel
senescence (El-Otmani et al., 1990). This is beneficial since it allows for storage of fruit and
longer harvest and market seasons (Coggins, 1992). It is a technique used in many countries
and on various cultivars (Tables 7 and 8). In fact, for some cultivars such as lime, green color
is sought. For mandarins (at least in the European market) the presence of greenish areas in the
stylar end of the fruit late in the season is a sign of freshness (see previous sections). However
fruit with too much green is less attractive. It is noteworthy that excessive rates of nitrogen
fertilization or N applications close to fruit color break delay color break just like GA3.

3. Rind softening and senescence


Citrus fruit rind is physically strong and physiologically young prior to the onset of
color change. During the change from a green to a yellow or orange flavedo, rind softening
begins and continues as the fruit matures through the harvest season (Coggins and Hield, 1968;
Coggins, 1981; El-Otmani et al., 1990) and during postharvest (El-Otmani and Coggins, 1991).
This softening leads to a number of rind disorders which reduce preharvest life, length of post-
harvest storage, shelf-life and market value of the fruit. In addition, senescing fruit lose their
resistance to decay from Penicillium (Coggins, 1969) and Geotrichum (Coggins et al., 1992a).
With many citrus types such as ‘Clementine’ mandarins, navels, lemons and grapefruit,
peak harvest occurs over a short period or during a season when demand is low. Supply is then
much greater than demand and market prices fall below an economic value. As a result, the
rate of harvest slows while fruit continue to soften. To overcome the problem, growers either
pick their fruit and store them in cold rooms, a practice that is often very expensive and only
accommodates a limited quantity of fruit, or leave their fruit “stored” on the tree to wait for
better prices.
GA3 decreases the rate of softening (Coggins and Lewis, 1965) and reduces rind disor-
ders associated with senescence such as rind staining of navel orange (Coggins et al., 1963),
rind puffing (Kuraoka et al., 1977), sticky rind (Coggins, 1969), and water spot (Riehl et al.,
1966). The benefits of healthier rinds resulted in the widespread use of GA3 to maintain citrus
fruit peel at a physiologically young stage to allow for an extension of the harvest season and
22

better control over the fresh fruit marketing season. Current application rates are shown in Ta-
ble 8. They vary with the cultivar and the region but spray solution concentrations of 10 to 20
mg∙l–1 GA3 are most common with the treatment applied as full tree coverage to the point of
run-off. The optimum time to apply the treatment in the orchard for maximum benefit is at fruit
color break for most cultivars.
Instances where GA3 is applied as a postharvest dip include ‘Marsh’ grapefruit in Israel
(Table 8). In fact Greenberg and Goldschmidt (1988) indicated that GA3 is used preharvest and
postharvest in Israel. GA3 delays bronzing of the flavedo and increases storage life of the fruit,
particularly of lemon and grapefruit (Hall et al., 1973; Ferguson et al., 1982; Greenberg and
Goldschmidt, 1988).
When GA3 is applied in storage wax to lemons, the result is delayed senescence, which
maintains natural resistance to sour rot (Geotrichum candidum Link ex Pers.) and otherwise
provides for a longer storage life (Coggins et al., 1992a). When it is added to the shipping wax
which is applied to yellow lemons and other mature citrus fruit, a delay in rind senescence,
including delayed rind color changes, can often be obtained (unpublished data). The resulting
delay in postharvest coloring of regreened ‘Valencia’ oranges may be undesirable, but other
delays in coloring of citrus fruit are considered to be beneficial.
When GA3 is applied at or just prior to color break, a significant delay in the develop-
ment of mature rind color occurs (Goldschmidt and Eilati, 1970; El-Zeftawi, 1978; Coggins,
1981; El-Otmani et al., 1990). The grower cannot harvest the fruit until it reaches acceptable
color even if market prices are very attractive. When application is delayed until later stages of
color development, the net reduction in softening will be reduced because the rind will already
have softened to a certain extent prior to treatment. However, in this case the fruit will have
already attained mature color prior to treatment and can be harvested at any time, keeping in
mind that the rind is less firm than that of fruit treated earlier in the season. Therefore, if a
grove is scheduled for early harvest or for degreening (such as the case of mandarins in early
season), GA3 is typically not applied. In addition, GA3 application at fruit color break will re-
duce carotenoid accumulation (El-Zeftawi, 1978), which results in treated fruit being pale or-
ange (for oranges, mandarins and their hybrids) but with a longer shelf-life. The presence of
too much green color on the fruit can significantly increase rejection at the packhouse and re-
duce packout. However, the presence of green color at the stylar end of ‘Clementine’ mandarin
and navel orange fruit as a result of treatment is a sign of freshness in the fresh marketplace
and it can be desirable particularly when the fruit is destined to travel long distances, to be
stored in cold rooms, or to be held on the shelf for long periods. GA3-treated fruit is resistant to
environmentally adverse factors such as rain-induced disorders like water spot or wind scarring
on fruit. High temperatures during fruit coloring can cause a significant delay in color devel-
opment (Grierson et al., 1986) and this should be taken into account when planning GA3 treat-
ment in large groves.

VII. USE OF PLANT GROWTH REGULATORS TO REDUCE FRUIT DROP AND


FOR ON-TREE STORAGE
In citrus, fruit drop occurs at different stages of fruit development: first drop, early
drop, occurs during the initial period of fruit set two to three weeks after bloom during the cell
division stage of fruit development (Colt and Hodgson, 1919; Erickson and Brannaman, 1960).
Abscission during early drop is at the abscission zone between the shoot stem and peduncle of
23

the fruit (Erickson and Brannaman, 1960). The second drop is June drop, a natural phenome-
non that results from competition for photoassimilates among tree organs. At this time, citrus
and other tree crops initiate the first major root flush, which diverts resources away from grow-
ing fruit. In June drop, abscission occurs between the floral disc and the ovary. June drop is
greatly influenced by environmental conditions such as high temperature and low humidity both
in the soil and in the air (Colt and Hodgson, 1919; Levitt, 1964; Davies et al., 1981). The third
classic drop period is preharvest drop, which is discussed later in this review.
Application of GA3 at 3 to 50 mg∙l–1 during the period from full bloom to petal fall is
widespread strategy for reducing early drop of mandarins, grapefruit, tangors, tangelos, and
navels to increase fruit set and yield (Table 2). GA3 used at 20 mg∙l–1 in sufficient volume to
give full tree coverage 4 to 5 weeks after full bloom (May) reduces June drop of ‘Jaffa’ orange
in Turkey (O. Tuzcu, personal communication, address provided in the Appendix I) (Table 2).
Results were improved when treatments were applied in the morning or late afternoon and
when a surfactant was added to the spray.
de Lima et al. (1980) investigated fruit drop between June drop and preharvest drop.
They found that two other drop periods occurred for navel oranges in Florida, namely a sum-
mer and a summer-fall drop. Summer drop occurred from mid-June until mid-August and was
primarily caused by the yellowing and subsequent abscission of the secondary fruit (navel) from
the primary fruit. de Lima and Davies (1984) determined that secondary fruit (navel) abscission
was associated with an increase in cellulase at the secondary fruit abscission zone, which in
turn produced ethylene levels sufficient to cause abscission of the primary fruit. Application of
2,4-D at 10 to 20 mg 2,4-D∙l–1 to the fruit or the entire tree reduced the severity of the prob-
lem. A recommended practice in Florida is to apply 2,4-D isopropyl ester at the rate of 60 to
70 g acid equivalent per hectare six to eight weeks after bloom to control summer drop (Knapp,
1996) (Table 9). Summer-fall drop occurs from mid-August until October with fruit nearing
maturity (de Lima et al., 1980). It is associated with splitting, stylar-end decay and other un-
known disorders (de Lima and Davies, 1981). Bujanda (1984) found that GA3 plus 2,4-D ap-
plied in mid-August reduced summer-fall drop. In Florida, it is recommended to apply 2,4-D
isopropyl ester at the rate of 70 g per hectare in August-September using 2500 to 5000 l spray
solution per hectare according to tree size and orchard density to reduce summer-fall drop in
navel oranges (Knapp, 1996) (Table 9). However, periods of leaf flush should be avoided. It is
also recommended not to apply the material when fruit is to be harvested early. Leaf and fruit
drop can also occur following an oil spray. 2,4-D is recommended (using the isopropyl ester at
0.25 g∙l–1 pesticide oil) on lemons and navels in California (Coggins,2000). It is however rec-
ommended not to apply this treatment to trees less than six years of age and within seven days
of harvest (Table 1).
Another major period of fruit drop involves mature fruit abscission prior to harvest
(Stewart and Klotz, 1947). It can be a significant problem where fruit are being held on the tree
past legal maturity. The fruit progresses from mature to enter the senescence phase. Fruit de-
cay, wind and environmental stresses, and physiological disorders can also contribute to this
drop (Wager, 1941; Klotz and Stewart, 1948). Application of 2,4-D has been used successfully
for many years in most citrus-producing countries to reduce preharvest drop for on-tree fruit
storage to extend the harvest period and market season. Information from our survey regarding
use of 2,4-D to prevent preharvest drop and for on-tree fruit storage for different cultivars in
different countries is reported in Table 9. Most of the respondents indicated that 2,4-D is ap-
24

plied at or just prior to fruit color break and that sprays are applied at 10 to 20 g active ingredi-
ent per liter as a full tree coverage spray. Not only oranges but also grapefruit, lemons and
mandarins and their hybrids respond well to the treatment.
Fruit color development may be delayed somewhat by 2,4-D if used before fruit reaches
full color (El-Otmani et al., 1990), therefore, timing of treatment should be well planned. To
assure residues are well below international tolerances no application should be made within
seven days of harvest (Erner and Coggins, 1989). In addition, late applications (for example,
January for ‘Washington’ navel oranges in California) if too close to the spring flush can cause
distortion of the leaves and stems of the new growth (Coggins, 1992).
It should be noted that when used at or just before color break, 2,4-D is often used with
GA3 to delay rind senescence so that treated fruit remain physiologically young for a longer
period of time, thus improving on-tree storage to extend harvest. This strategy makes it possi-
ble to control fruit supply in relation to market demand (Coggins, 1981; El-Otmani et al.,
1990; Coggins, 1992). Autumn GA3 application to delay coloration and extend peel viability
may adversely affect flower bud differentiation, causing some loss in yield the next season.

VIII. USE OF PLANT GROWTH REGULATORS TO FACILITATE HARVEST


Several attempts have been made to develop a commercially viable and an easily imple-
mentable technique to facilitate mechanical harvesting of citrus fruit. The subject has been re-
viewed by Wilson et al. (1981) and Wilson (1983). Most of the chemicals tested caused abscis-
sion through a superficial peel burn which caused the production of wound ethylene which
moved through fruit tissues to the abscission zone to loosen the fruit. Because most of the com-
pounds caused injury to the fruit, the only commercial application known to us is that of
ethephon (Table 10).
Ethephon, an ethylene-releasing agent, is the only commercially available chemical that
causes no damage to the fruit peel that has shown promising results as a fruit-loosening agent.
This compound was registered in Florida on lemons, tangerines and tangerine hybrids at con-
centrations of 200 to 250 mg∙l–1 (Knapp, 1981). Besides its fruit-loosening effect, ethephon
enhances fruit color (Young et al., 1974). However, there are two major draw-backs: 1) activ-
ity of ethephon is significantly affected by environmental factors and the physiology of the tree;
and 2) ethephon has an erratic effect on fruit-loosening and a tendency to cause excessive leaf
drop (see Wilson (1983) and Wilson et al. (1981) and the references therein). Fruit is respon-
sive once it reaches minimum legal maturity and remains responsive as fruit maturation pro-
gresses. ‘Valencia’ orange becomes unresponsive to treatment during regreening (Wheaton et
al., 1977), a period characterized by the conversion of chromoplasts to chloroplasts in a re-
sponse to warm spring temperatures. On the other hand, for best results, daily high tempera-
tures need be at least 18.3°C or greater following treatment. Rainfall within 24 hours of
ethephon application will eliminate any beneficial result (Wilson et al., 1981). It is noteworthy
that because of these problems, use of ethephon on citrus in Florida has been discontinued.
From the survey we conducted, Cuba appears to be the only country where chemicals to
aid harvesting are used (Table 10). Ethephon is used to facilitate both manual and mechanical
harvesting. For manual harvesting, ethephon is used on ‘Eureka’ lemon to reduce the force
necessary to separate the fruit from its stem and increase picker efficiency (i.e., number of
trees harvested per day), and on ‘Dancy’ mandarin and ‘Orlando’ tangelo to facilitate separa-
tion of the fruit from the stem which results in less plugging damage to the fruit (Perez et al.,
25

1992). Other beneficial results include lower postharvest decay and better lemon color devel-
opment resulting in improved market value of the fruit. Because a fruit harvested by applying a
pulling force may lose its peduncle and calyx in the process, it is essentially destined to local
markets. For mechanical harvesting, ethephon is used on grapefruit and ‘Valencia’ orange to
reduce fruit attachment force. For best efficacy of the treatment, it is important to ensure opti-
mum irrigation and fertilization practices in the orchard. This would also reduce the risks of
excess defoliation and drop of immature fruit in cases where fruit at different developmental
stages are on the tree. Addition of calcium (as calcium hydroxide) to the spray solution is also a
common practice in Cuba in order to reduce leaf abscission (Perez et al., 1992).

IX. USE OF PLANT GROWTH REGULATORS POSTHARVEST TO MAINTAIN


FRUIT QUALITY
Prolonging citrus fruit shelf-life is the goal of researchers working in the area of post-
harvest and this subject is approached from different angles (see El-Otmani and Ait-Oubahou,
1996). Two review papers have recently been published on the subject of plant growth regula-
tors and citrus fruit quality (El-Otmani and Ait-Oubahou, 1999a, 1999b) and the reader should
refer to them for detailed information. In summary, GA3 and 2,4-D are used commercially to
maintain rind firmness and avoid loss of the calyx and thus reduce fruit decay (Table 11). Con-
versely, ethylene has a deleterious effect on the storage life of fruit. Limited information is
available on the effect of plant growth regulators on chilling injury but no significant or consis-
tent effect was observed on internal quality.

X. WAYS TO ENSURE OR INCREASE THE EFFICACY OF PLANT GROWTH


REGULATORS IN CITRICULTURE
From the survey, as well as from the literature, it appears that factors that would affect
solubilization, penetration, absorption and translocation and the molecular structure of exoge-
nously applied plant growth regulators will affect their efficacy and will thus influence the suc-
cess of their use. Use of surfactants has been shown to greatly improve wetting of the leaf sur-
face by lowering the surface tension and consequently reducing the contact angle between the
liquid and leaf surface. Complete wetting is achieved when the contact angle is zero. The criti-
cal surface tension (i.e., the liquid surface tension at which the contact angle is zero) of citrus
fruit varies slightly with the surfactant (Henning and Coggins, 1988). Infiltration of exogenous
sprays containing surfactants into the plant stomata was reported by several authors (Schonherr
and Bukovac, 1972; Greene and Bukovac, 1974). The number of stomata infiltrated was in-
versely related to the surface tension of the solution (Greene and Bukovac, 1974).
After testing seven surfactants in the laboratory, Henning and Coggins (1988) found that
the inclusion of these compounds in the treatment spray doubled the effectiveness of GA3 in
delaying the loss of chlorophyll from the rinds of green grapefruit used as the assay plant mate-
rial. Triton B-1956 provided the least benefit of the surfactants tested. Those surfactants that
had the best wetting or spreading capacity did not necessarily give greater GA3 efficacy indicat-
ing that increased efficacy, which is probably a result of enhanced penetration cannot be attrib-
uted entirely to a lowered surface tension (Sargent, 1965). In Israel, Silwet® L-77, which is an
organosilicone surfactant, enhanced the effectiveness of GA3 in prolonging the harvest season
of citrus far more that Triton B-1956 (Greenberg et al., 1987). The enhancement was a result
26

of better uptake, which resulted from changes in the epicuticular wax when L-77 was used in
the spray solution (Greenberg and Goldschmidt, 1988).
Henning and Coggins (1988) found a poor relation between surfactant concentration
versus surface tension, area wetted and biological effectiveness of GA3. They postulated that
this may be due, in part, to the inclusion of relatively ineffective ingredients as active ingredi-
ents in some commercially available surfactants that they evaluated. This can also be the reason
for the wide range in concentrations recommended by the manufacturers of these materials.
Furthermore, the addition of these materials to the spray solution can cause injury to the fruit
(Coggins and Henning, 1988) or reduce rind blemishes usually caused by the plant growth
regulator (Gilfillan et al., 1973). In a similar fashion, improved abscission was observed when
certain surfactants were included in the spray solution but other surfactants caused phytotoxicity
and excessive leaf drop (Wilson et al., 1981). Depression of abscission activity by surfactants
has also been reported (Wilson, 1983). Thus, interactions between wetting agents and plant
growth regulators should be taken into account when selecting the spreader to use. Because no
clear conclusion can be made about effectiveness of any surfactant over the others, performance
tests should be conducted with the locally available surfactants, under local environmental con-
ditions, and using local cultivars and strains.
Activity of certain plant growth regulators can be increased by acidifying the spray solu-
tion (Greenberg et al., 1984; Gilfillan, 1986; Greenberg and Goldschmidt, 1989). Highly alka-
line conditions cause decomposition and loss of activity of plant growth regulators such as GA3
(Coggins, 1981). However, a pH lower than 4 caused decomposition of GA3 (Plant Protection
Ltd., 1969). Using radioactive GA3, Greenberg et al. (1984) demonstrated that low pH en-
hances GA3 uptake and penetration into citrus fruit with uptake rates at pH 4 being 3-fold
higher that at pH 7. Enhancement of GA3 activity was independent of the acidifying agent used
(Greenberg and Goldschmidt, 1988; Greenberg et al., 1992). Results obtained in experiments
with ‘Clementine’ mandarin indicate that Silwet® L-77 as well as spray solution acidification
increase GA3 efficacy (Ben- Ismail et al., 1995). Conversely, Coggins et al. (1992b) reported
no beneficial effect from acidifying the spray solution under field conditions. They reported
that inclusion of Silwet® L-77 was more promising than lowering pH. Currently, acidification
of the spray solution is practiced in countries such as South Africa and Israel (Gilfillan, 1986;
Greenberg et al., 1992).
It is recommended that GA3 should not be combined with compounds [such as Bordeaux
mixture used to control Phytophthora or whitewash (calcium hydroxide) used to control leaf
hopper in California] that would increase pH to values greater than 8 (Coggins, 1981). The
recommendation is to apply GA3 separately from these treatments with at least three days be-
tween the two sprays (Coggins, 1981). Gilfillan and Cutting (1992) reported lower efficacy of
GA3 in sprays containing petroleum oil and found that acidification of the GA3 spray to pH 4
improved efficacy, with or without the oil spray.
In general, uptake of plant growth regulators, for example GA3, is proportional to the
GA3 concentration in the treatment solution and to the duration of uptake (Greenberg and Gold-
schmidt, 1988). Weather conditions are important factors in effectiveness of foliar-applied plant
growth regulators (Coggins et al., 1960a, 1960b; Hield et al., 1964; Fucik, 1981; Hirose,
1981; Wilson et al., 1981). Greenberg and Goldschmidt (1988) reported that GA3 uptake was
greater at high relative humidity and increased with temperature in the range of 5 to 35 °C.
Light had no effect, however. These authors also reported that uptake from the dry residue left
27

on citrus fruit surface after drying of the spray was higher when the surrounding humidity was
higher (100% versus 50%) and that at high humidities uptake still continued even as late as 48
hours after drying (Greenberg and Goldschmidt, 1990). Optimum temperature for uptake of
abscission accelerating compounds is 25°C (Evensen et al., 1981), and optimum humidity is
60% (Biggs and Kossuth, 1981). Taken together, these results indicate that the best time of day
for spray application is early morning and late afternoon when the humidity is high and the
temperature is not too warm (for summer) nor too cold (for winter) and that the nightly in-
crease in relative humidity in the field may induce renewed uptake of the compound from the
dry spray residue and increase its physiological effect provided the plant growth regulator is
stable on the plant surface. When applied in wax, Goldschmidt and Greenberg (1989) reported
that GA3 is stable and this shows the potential value of the wax for prolonged supply of GA3 to
the peel during storage. In addition, optimum temperature and humidity not only increases up-
take, but also reduces phytotoxicity and leaf and fruit drop (e.g., for ethephon or NAA sprays).
Spray application at night hours in the summer and mid-day in the winter are acceptable par-
ticularly if light has no significant effect on uptake and because in the winter mid-day tempera-
tures are suitably warm. Goldschmidt and Greenberg (1989) reported that light and low relative
humidity enhanced GA3 decomposition. Furthermore, rain within hours following foliar spray
may completely eliminate the desired treatment effect (Wilson et al., 1977a; Biggs and Kos-
suth, 1981).
Cultural practices influence the effectiveness of plant growth regulator sprays and opti-
mum irrigation, fertilization and tree health should be the rule in orchards planned to receive
hormone sprays (Gilfillan et al., 1973; Wheaton, 1981; Gallasch, 1984). This would ensure
consistent optimum results and improved performance. Monselise and Sasson (1977) found that
after prolonged storage, fruit treated with the combination of 2,4-D and GA3 from frequently
irrigated trees had stronger peels and were more resistant to postharvest handling.
Compatibility with other foliar treatments should be taken into account since combining
treatments will reduce the number of sprays and thus reduce cost of production (Gallasch,
1984). For example, efficacy of GA3 is not affected when mixed with micronutrients (Perez et
al., 1992) particularly, if the nutrient preparations are neutralized (Coggins, 1989). Ethephon
can also be mixed with micronutrients without adverse side effects and thus reduce the cost of
application (Gallasch, 1984).
Timing of application is critical for best results (see previous sections). It is also note-
worthy that the physiological stage of the target organ plays an important role in the type of
response observed as a result of a treatment. For example, regreened fruit or fruit on young
trees are less responsive to ethephon spray because these fruit are less responsive to exogenous
ethylene (Wheaton et al., 1977; Evensen et al., 1981). Timing is also critical to avoid chemical
residues in the fruit. Application of 2,4-D to control preharvest drop of mature fruit for exam-
ple should be made at least seven days prior to harvest in California (Erner and Coggins,
1989). A report from Morocco on ‘Clementine’ mandarins indicates that when 2,4-D was ap-
plied at 10 mg∙l–1 as a postharvest prestorage dip, peel residue content after four days in storage
at 4°C and 20°C was 10% and 3%, respectively (El-Otmani et al., 1998) indicating that low
temperatures slow the dissipation of the active ingredient.

XI. CONCLUSION AND FUTURE PROSPECTS


28

Plant growth regulators have been in use in citriculture for more than half a century.
Their use is dictated by the immediate objectives and needs of the citrus industries of the vari-
ous countries. Some production problems (and thus objectives) are common to many countries
and the compounds used to solve these problems are often similar but concentrations used and
amounts applied vary. A few of the compounds have been questioned with regard to their ef-
fects on human health and the environment. Most plant growth regulators are used at very low
concentrations, which result in low residues and low to no hazards to human health or the envi-
ronment. Laws regulating the use of plant growth regulators in the different citrus-producing
countries are extremely varied and beyond the scope of discussion of this review. An important
goal of researchers has been to find ways to increase or enhance the efficacy of plant growth
regulators in order to reduce the quantity applied. A long-term objective of all researchers in
this field is to be able to understand the basic mechanisms underlying citrus growth and devel-
opment in order to be able to manipulate all the key physiological processes using the right
compounds and formulations at the right concentrations and at the right time by targeting the
right organ. However, it seems that we still are far from complete knowledge of how to apply
some PGRs with maximum efficiency.
In many aspects, physiological and biochemical research may have reached a saturation
point or even a deadlock (Goren et al., 1996). Learning how PGRs work at the molecular and
cellular level may help advance our knowledge of the role of endogenous hormones to enhance
the effectiveness of exogenously applied PGRs or reduce their need if transgenic citrus plants
can be produced to meet desired production goals, provided that no health or environmental
hazard is associated with use of these plants as is the case with other plant species. Biotechnol-
ogy is only at its first steps, even once it is advanced, it may not provide all the answers or all
the solutions. It is likely that there may always be a need for plant growth regulators in citricul-
ture.

XII. ACKNOWLEDGMENT:
The authors gratefully acknowledge the contributions of all of the scientists and special-
ists who immensely enriched this article with the information they provided in responding to
our questionnaire.
29

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Wilson, W.C., Kenne, D.S., and Holm, R.E. 1977b. The Florida Department of Citrus cooperative chemical
screening programs for citrus. Proc. Intl. Soc. Citriculture 2:692–696.
Yelenosky, G. 1985. Cold hardiness in citrus. Hort. Rev. 7:201–238.
Yelenosky, G., Vu, J.C.V., and Wutscher, H.K. 1995. Influence of paclobutrazol in the soil on growth, nutrient
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710.
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39

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40

Table 1. Use of plant growth regulators on citrus to manipulate vegetative and reproductive development.

Country Cultivar⊥ Growth Time/stage of Rate of appli- Remarks Source‡


regulator application cation
used
Japan Greenhouse 6-benzyl- Immediately 300 g∙ha–1 as a accelerates vegetative and I. Iwagaki
wase Satsuma adenine after heating full tree cov- flower budbreak
mandarin the green- erage
surfactant added at
house
0.075%
New Zealand Satsuma GA3 Early winter Nonbearing reduces white blossom A.R. Harty
mandarin (mid-June) trees (≤2 years
increases vegetative shoot
old): 50 mg∙l–1
production
Bearing trees
reduces hand thinning
(>2 years
necessary to increase
old): 25 mg∙l–1
fruit size
use non-ionic surfactant
on bearing trees and an
organosilicone penetrant
on nonbearing trees
need excellent spray
coverage for good effect
USA (California) All citrus 2,4-D With pesti- Isopropyl ester may provide some degree C.W. Coggins,
cide oil of 2,4-D at of preharvest fruit drop Jr.
sprays to 0.25 g∙l–1 of control but because of the
reduce leaf pesticide oil long time span between
drop most pesticide oil sprays
and the onset of mature
fruit drop an additional
aqueous 2,4-D preharvest
spray may be needed
do not apply to trees <6
years old
do not apply within 7
days of harvest
USA (Florida) All citrus ethylester Prior to Apply to trunk excess use may result in D.P.H. Tucker
of NAA sprout as a thorough tree damage and T.A.
1.15% growth to spray or light Wheaton
(Tre- inhibit trunk brush applica-
Hold®) sprout tion
growth


Species names are provided in Appendix I.

Addresses are provided in Appendix II.
41

Table 2. Use of gibberellic acid to improve fruit set (i.e. to reduce early drop and June drop).

Country Cultivar Time/stage of Rate of Remarks Source


application application
a. Reduction of early drop
Argentina Ellendale mandarin 75-100% full 10 mg∙l–1 no pH control M.L. Ragone
bloom spray as full
no surfactant added
tree cover-
age avoid application to water-
stressed trees

Cuba Marsh grapefruit, full bloom to 42-62 g∙ha–1 surfactant added at 0.025% C. Noriega, L.
Ortanique tangor petal fall using 20 Pozo, M. del C.
spray in morning to avoid after-
(fruitlet di- mg∙l–1 spray Perez, and H.
noon showers
ameter 3-5 solution as a Oliva
mm) full tree often used in combination with
coverage urea (0.5%), Zn sulfate (0.15%)
and Mn sulfate (0.12%)

France (Corsica) Clementine manda- full bloom 10 mg∙l–1 surfactant used H. Vanniere
rin with 2500
no pH control
l∙ha–1
Israel Clementine manda- full bloom 20-25 mg∙l–1 surfactant added at 0.025% E.E. Gold-
rin with 3500 schmidt, Y.
l∙ha–1 Erner, and J.
Greenberg
Nova mandarin, full bloom 50 mg∙l–1 “
Ortanique tangor, with 3500
Minneola tangelo l∙ha–1
Italy Clementine manda- postbloom 20 mg∙l–1 E. di Martino
rin Aleppo
Morocco Clementine manda- full bloom to 10-20 mg∙l–1 rate split in two applications in M. El-Otmani
rin petal fall as full tree years of long bloom period
coverage
surfactant added at 0.02-0.03%
mandarin hybrids full bloom to 10-25 mg∙l–1 “ “
petal fall as full tree
coverage
South Africa Clementine manda- 90-100% 3-5 mg∙l–1 pH=5 when controlled. I. Gilfillan
rin petal fall spray solu-
surfactant added at 0.015-
(Oct.) tion as full
0.025%
tree cover-
age compatible with all material
except alkaline or oil sprays

Spain Clementine manda- 90-100% 18 g∙ha–1 often used in combination with M. Agustí
rin petal fall Mn and Zn
(Apr.- May)
surfactant used at 0.05%
42

no pH control
2 treatments are often required
–1
Navelate orange 90-100% 5 mg∙l as a no pH control “
petal fall full tree
often used with KNO3 (2%)
coverage
often followed by girdling 20
days later
Turkey Clementine and full bloom 5-10 g∙ha–1 surfactant used O. Tuzcu, A.I.
Satsuma mandarins (Apr.) as a full tree Ozguven, and T.
best result if applied in the
coverage Yesiloglu
morning or late afternoon

Uruguay Ellendale mandarin full bloom 15 mg∙l–1 no surfactant used A. Gravina


with 2000
pH=5.0 using H3PO4
l∙ha–1
USA (Florida) Tangelo full bloom 25-70 g∙ha–1 rates based on application in D.P.H. Tucker
2500-5000 l∙ha–1 to mature trees and T.A. Whea-
ton
can result in small fruit size and
leaf drop
b. Reduction of June drop
Turkey Jaffa orange 4-5 weeks 20 mg∙l–1 surfactant added O. Tuzcu
after full spray solu-
best results if applied early
bloom tion as a full
morning or late afternoon
(May) tree cover-
age
43

Table 3. Use of plant growth regulators to induce fruit thinning in citrus to improve fruit size.

Country Cultivar Growth regu- Time/stage of Rate of Remarks Source


lator used application application
Argentina Common man- Ethephon fruit diameter 200 mg∙l–1 severe overthinning M.L. Ragone
darins 15-20 mm and leaf drop can
occur, especially if
temperature is high
Israel Clementine and NAA fruit diameter 300 mg∙l–1, surfactant used at E.E. Gold-
Dancy mandar- 8-10 mm 3500 l∙ha–1 0.025% schmidt, Y.
ins, Murcott, Erner, and J.
tangor, Tuli- Greenberg
gold and Bo-
nanza navels
USA (Florida) Mandarins and NAA mid-May 500-1250 use lower rates on D.P.H.
Murcott tangor g∙ha–1, ‘Murcott’ Tucker and
spray at T.A. Wheaton
ammonium salt
2500-5000
formulation
l∙ha–1
activity is tempera-
ture- dependent
severe over thinning
may result from
applications to trees
of low vigor or
under water stress
44

Table 4. Use of plant growth regulators to enhance fruit growth and improve size.

Country Cultivar Growth Time/stage of Rate of Remarks Source


regulator application application
used
France (Corsica) Clementine 2,4-DP end of physio- 50 mg∙l–1 butyl glycol ester H. Vanniere
mandarin logical drop with solu-
no surfactant added
tion at 2500
1∙ha–1 trees well-watered prior
to treatment
Israel Clementine 2,4-DP end of physio- 50 mg∙l–1, surfactant used, pH=3-4 E.E. Gold-
mandarin logical drop 3500 1∙ha–1 schmidt,
KNO3 (5%) added into
Y. Erner, and
spray
J. Greenberg
Shamouti and 2,4-D “ 20 mg∙l–1, “ “
Valencia or- 3500 1∙ha–1
ange
Valencia or- 3,5,6-TPA “ 10-15 mg∙l–1 “ “
ange, Grape-
fruit, Satsuma
mandarin
Morocco Clementine 2,4-D fruit diameter 10-12 mg∙l–1 butyl glycol ester M. El-Otmani
mandarin 10-15 mm applied as
surfactant added at
full cover-
0.02-0.03%
age
may cause leaf distortion
“ 2,4-DP end of physio- 50 mg∙l–1 butyl glycol ester “
logical drop applied as
surfactant added at
full tree
0.02-0.03%
spray
“ 3,5,6-TPA end of physio- 10-15 mg∙l–1 acid formulation, surfac- “
logical drop applied as tant added at 0.02-0.03%
full tree
spray
Spain Clementine 2,4-DP end of physio- 180 g∙ha–1 butyl glycol ester formu- M. Agustí
mandarin, logical drop lation
navel, Valen-
surfactant added at
cia orange
0.05%
“ 3,5,6-TPA “ 36 g∙ha–1 acid formulation “
surfactant added at
0.05%
“ Phenothiol “ 72 g∙ha–1 surfactant added at “
0.05%
USA (California) Grapefruit 2,4-D fruit diameter 57 to 110 isopropyl ester C.W. Coggins,
6 to 25 mm g∙ha–1 ac- Jr.
requires excellent tree
cording to
coverage
fruit diame-
ter; use in also controls fruit drop
spray up to
45

5000 1∙ha–1
Valencia and 2,4-D fruit diameter 57 to 110 isopropyl ester formula- “
navel orange 5 to 19 mm g∙ha–1 ac- tion
cording to
requires excellent tree
fruit diame-
coverage
ter. Use
spray up to also controls mature fruit
5000 1∙ha–1 drop in ‘Valencia’
may decrease fruit split-
ting in navel oranges
46

Table 5. Use of plant growth regulators on citrus to improve fruit internal quality and hasten fruit maturation

Country Cultivar Growth regu- Time/stage of Rate of Remarks Source


lator used application application
Japan Satsuma mandarin Ethychlozate Two applica- results in faster sugar I. Iwagaki
tions: accumulation
The first 50-60 400 g·ha–1 additional benefits include
days after full fruit thinning and reduced
bloom (early and rind puffing
July);
surfactant added at 0.01%
the second 400 g∙ha–1
70-80 days
after full bloom
(late July)
South Valencia orange Calcium 2-6 weeks after 150 g cal- reduces acid in juice I. Gilfillan
Africa arsenate full petal fall cium arse-
earlier harvest
nate per 100
l spray solu- apply alone
tion
no surfac-
tant used
50 g cal-
cium arse-
nate per 100
l if pH of
spray solu-
tion is re-
duced to 5.0
47

Table 6. Current uses of plant growth regulators to reduce citrus rind disorders.

Country Cultivar Objective of Growth Time/stage of Rate of Remarks Source


treatment regulator application application
used
Australia Navel orange reduce GA3 late Jan.-Feb. 100-150 surfactant added P.T. Gallasch
creasing (fruit diameter g∙ha–1
pH 4.5 using buff-
30-50 mm)
ering agents
best if applied af-
ternoon for en-
hanced uptake
Israel Minneola reduce “ June-July 10 mg∙l–1; pH=3-4 E.E. Gold-
tangelo creasing 3500 1∙ha–1 schmidt,
surfactant added at
Y.Erner and
0.025%
J. Greenberg
Murcott reduce split- 2,4-D May-June 20 mg∙l–1; pH=4 “
tangor ting 3500 1∙ha–1
surfactant added at
0.025%
KNO3 (5%) added
in spray
Murcott “ 3,5,6- early July 10-15 “ “
tangor and TPA mg∙l–1;
Nova manda- 3500 1∙ha–1
rin
Nova “ 2,4-D 3 sprays from 20 mg∙l–1; “ “
mandarin end of May 3500 1∙ha–1
until mid-July
Valencia reduce GA3 July-Aug. 20 mg∙l–1; pH=3-4 “
orange creasing 3500 1∙ha–1
surfactant added at
0.025%
South Africa Navel orange reduce GA3 fruit diameter 20 mg∙l–1 surfactant used at I. Gilfillan
creasing 30-40 mm on Troyer 0.015- 0.025%
(Jan.) or trifoliate
important pH=5 when con-
rootstock
problem in trolled
and 10
saline areas,
mg∙l–1 on maintain optimum K
with heavy
rough fertilization and
crops and
lemon or optimum irrigation
with
Volkameri- to avoid leaf drop
thin-skinned
ana root-
fruit not compatible with
stock at
oil or alkaline
6000 1∙ha–1 sprays
USA (Florida) Minneola delay of GA3 early Dec. 48g∙ha–1 do not use in spray D.P.H.
tangelo stem-end with spray solution with Tucker and
rind break- at 2500- pH>8.0 T.A. Whea-
down 5000 1∙ha–1 ton
48

Table 7. Use of plant growth regulators pre- and postharvest to enhance fruit color.

Country Cultivar Growth Time/stage of Rate of Remarks Source


regulator application application
used
Argentina Satsuma and ethylene postharvest 5-10 mg∙l–1 20-25°C, 90% RH M. Ragone
Clementine
mandarin,
navel orange
France (Corsica) Clementine ethephon 25% fruit peel 360 mg∙l–1 T°>15°C but <25°C H. Vanniere
mandarin is yellow or- spray solu-
no surfactant
ange; 2-3 tion with
weeks before 1500 1∙ha–1 pH not adjusted
harvest
tree well watered prior to
treatment
Cuba All grape- ethylene postharvest 5-10 mg∙l–1 use continuous system F. Prividal,
fruit culti- (Aug.-Dec.) I. Izquierdo and
temperature=30°C
vars F. Guerra
relative humidity=95%
Israel Early navel, ethylene postharvest 5-10 mg∙l–1 - E.E. Gold-
early grape- gas schmidt, Y.
fruit Erner and J.
Greenberg
Israel Kumquat ethephon 2 weeks before 500-1000 - “
harvest mg∙l–1
Morocco Clementine ethylene postharvest 10 mg∙l–1 in temperature of 23-25 °C M. El-Otmani
mandarin gas the degreen-
humidity~95%
ing room
good air circulation and
ventilation
dip fruit in 2,4-D (4-8 mg∙l–
1
) prior to degreening
degreening time dependent
upon green color intensity
South Africa Mihowase ethephon 2 weeks before 250 mg∙l–1 surfactant used at I. Gilfillan
Satsuma and harvest 0.015-0.025%
Nules
temperatures following
Clementine
spray >25 °C for at least 3
mandarin
days
internally mature but still
green fruit
avoid water stressed trees
Spain All citrus ethylene postharvest 5-10 mg∙l–1 22-25°C, >90% RH M. Agustí
–1
USA (California) All citrus ethylene postharvest 1-10 mg∙l 21-29°C, ~95% RH C.W. Coggins,
Jr.
49

Table 8. Use of gibberellic acid to delay fruit color and rind senescence.

Country Cultivar Time/stage of Rate of application Remarks Source


application
Australia Grapefruit just before fruit use a thorough tree surfactant added P.T. Gallasch
color break coverage with 20
pH=4.5
(Mar.-Apr.) mg∙l–1 spray solu-
tion
Mandarins and their just before rind use a thorough tree surfactant used “
hybrids color break coverage with a 10
pH=4.5
(Mar.-Apr.) mg∙l–1 spray solu-
tion
Navel orange 2 weeks prior 50-75 g∙ha–1 as a surfactant added “
to fruit color thorough spray
also include 2,4-D to
break (Mar.- coverage
reduce preharvest fruit
Apr.)
drop
pH=4.5
–1
Brazil Orange (Pera, Natal, 10-20 mg∙l surfactant L-77, 0.05% P.R.C. de Cas-
Valencia, Hamlin, tro and
Westin), Bearss lime, A. Malavasi
Murcott tangor and
Ponkan mandarin
Cuba Dancy mandarin fruit color 78 g∙ha–1; 8-10 l pH = 4.0 L. Pozo, M.
break (Oct.) per tree del C. Perez,
surfactant used
C. Noriega
Persian lime postharvest 30 mg∙l–1 application in storage T. Castro-
wax Lopez
Israel Marsh grapefruit postharvest dip 20-25 mg∙l–1 surfactant used at E.E. Gold-
0.025% schmidt,
Y. Erner, and
no pH control
J. Greenberg
Navel orange, Minne- fruit color 20-25 mg∙l–1 with surfactant used at “
ola tangelo break 2-3 spray at 3500 l∙ha– 0.025%; pH=3-4
1
months prior to
harvest
Oroblanco grapefruit “ 10 mg∙l–1 with “ “
spray at 3500 l∙ha–
1

Italy Lemon and Mandarin preharvest 20 mg∙l–1 - E. di Martino


Aleppo
Morocco Clementine mandarin just prior to 10-20 mg∙l–1 spray surfactant used at 0.02- M. El-Otmani
fruit color solution as a full 0.03%
break tree coverage
color development may
be delayed further by
high temperatures dur-
ing fall particularly in
warm areas
reduces water spotting
50

and puffing
South Africa Eureka lemon 6-8 weeks be- 10 mg∙l–1 at 6000 delays harvest for I. Gilfillan
fore expected 1∙ha–1 ~3-4 weeks
start of normal
surfactant used at
harvest
0.015-0.025%
pH reduced to 5.0
–1
Grapefruit fruit color 10-20 mg∙l at extends picking period “
break 6000 1∙ha–1 de- by 1-3 weeks
pending on tree
used with 2,4-D amine
size
at 10 mg∙l–1
pH controlled to 5.0
using phosphate buffer
or sulfuric acid
surfactant added at
0.015-0.025%
not compatible with oil
or alkaline sprays
Spain Clementine mandarin 2 weeks prior 36 g∙ha–1 surfactant used at M. Agustí
to fruit color 0.05%,(NH4)2HPO4
break (Oct.) added at 1.5%
reduces rind blemishes
linked to senescence
Fortune mandarin 2 months past 36 g∙ha–1 surfactant used “
fruit color (0.05%)
break
no pH control
reduces rind blemishes
–1
Lemon postharvest 10-20 mg∙l applied in wax “
delays coloring
Navel orange 2 weeks prior 36 g∙ha–1 used with 2,4-D (15 “
to fruit color mg∙l–1)
break
surfactant used at
0.025%
no pH control
–1
Satsuma mandarin 4 weeks prior 36 g∙ha surfactant used at “
to fruit color 0.05%
break (late
(NH4)2HPO4 added at
Sept.)
1.5%
reduces rind puffing
Turkey Kutdiken lemon fruit color 10 mg∙l–1 as a full surfactant used O. Tuzcu, A.I.
break (Nov.) tree coverage Ozguven,
best results when ap-
O. Dundar
plied early morning or
late afternoon
Marsh grapefruit fruit color 10 mg∙l–1 as a full surfactant used “
51

break (Nov.) tree coverage best if applied early


morning or late after-
noon
Navel orange fruit color 10-20 mg∙l–1 as a best results if applied “
break (Oct.) full tree coverage early morning or late
afternoon
surfactant used
USA (California) Lemon (non-desert when target 25 to 50 g∙ha–1 in reduces the number of C.W. Coggins,
areas) fruit is 1/2 to spray up to 5000 small tree-ripe fruit and Jr.
3/4 full size but 1∙ha–1 delays flowering which
still green shifts following crop
towards summer
do not apply within 30
days of harvest
Lime (non-desert areas) when target up to 50 g∙ha–1 in delays fruit color “
fruit is 1/2 to spray up to 5000
3/4 full size 1∙ha–1
and still green
Minneola tangelo 2 weeks before 50-100 g∙ha–1 in reduces rind puffing “
fruit color spray up to 5000
satisfactory color is
break 1∙ha–1
attained late Jan.
do not apply in high
pH solutions
Navel orange Sept.-Nov. (2 25-100 g∙ha–1 with for groves to be har- “
weeks before spray up to 5000 vested after Feb. 15
fruit color 1∙ha–1
if delayed coloring is
break)
not tolerated, apply
after marketable color
has developed but re-
sults in less control of
aging
do not apply within 10
days of harvest or in
mixture producing a
high pH
late application (Jan.)
may reduce subsequent
production
reduces rind staining,
water spot, and sticky
rinds
Valencia orange Aug.-Sept. 100-200 g∙ha–1 in reduces creasing, de- “
spray up to 5000 lays aging and soften-
l∙ha–1 ing of the rind
later application may
cause intensive re-
greening of mature
crop if present, and
52

slower color develop-


ment in the target crop
Lemon postharvest 50-100 mg∙l–1 applied in storage wax “
–1
USA (Florida) Seedless grapefruit Nov.-Jan. 48 g∙ha with combine with 2,4-D for D.P.H. Tucker
spray at 2500-5000 fruit drop control and T.A.
1∙ha–1 Wheaton
do not use in spray
solutions above
pH=8.0
53

Table 9. Current uses of 2,4-D to reduce preharvest drop and prolong on-tree storage of fruit.

Country Cultivar Time/stage of Rate of Remarks Source


application application
Argentina Ellendale mandarin, fruit color 16 mg∙l–1 butylester, surfactant added M.L. Ragone
Navel orange, break
used in combination with GA3
Marsh grapefruit
(10 mg∙l–1)
no pH control
–1
Australia Grapefruit just before 20 mg∙l dimethylamine salt P.T. Gallasch
fruit color as a full
surfactant used
break tree cover-
(Apr.-May) age often mixed with GA3 to de-
lay rind aging
Mandarins and their just before 10 mg∙l–1 dimethylamine salt “
hybrids fruit color as a full
surfactant added
break tree cover-
(Apr.-May) age often mixed with GA3 to de-
lay rind aging
Navel orange Just before 10 mg∙l–1 dimethylamine salt “
fruit color as a full
surfactant added
break tree cover-
(Apr.-May) age often used with GA3 to delay
rind aging

Israel Marsh grapefruit, 2-3 months 20 mg∙l–1, surfactant used at 0.025% E.E. Goldschmidt,
Temple orange, before harvest 3500 1∙ha–1 Y. Erner and J.
Valencia orange, Greenberg
Minneola tangelo,
Ortanique tangor
Morocco Navel orange fruit color 10-20 not a general practice M. El-Otmani
break mg∙l–1 as a
surfactant added at
full tree
0.02-0.03%
coverage
New Zealand Grapefruit fruit color 16-20 second spray in late July to A.R. Harty
break (Apr.- mg∙l–1 early Aug. often applied (i.e.
May) before spring growth begins)
Navel orange fruit color 16-20 second spray in mid-late July “
break (Apr.- mg∙l–1 often applied
May)
South Africa Grapefruit fruit color 10 mg∙l–1 amine formulation with GA3 I. Gilfillan
break at 6000 at 10-20 mg∙l–1
1∙ha–1
pH adjusted to 5.0
surfactant added at
0.015-0.025%
Navel and Valencia just before full 10-20 compatible with all materials “
orange maturity mg∙l–1 at except alkaline or oil contain-
6000 1∙ha–1 ing spray materials
54

avoid leaf flush


no pH control
–1
Spain Fortune mandarin fruit color 16 mg∙l isopropylester M. Agustí
break (Nov.) as full tree
used with Ca(NO3)2 (2%) for
coverage
control of “peel pitting”
no pH control
surfactant used
Navel orange, Nova 2 weeks prior 54 to 75 isopropylester “
mandarin to fruit color g∙ha–1
no pH control
break
surfactant used at 0.05%
used in combination with GA3
(10 mg∙l–1)
Turkey Kutdiken lemon just before 20 mg∙l–1 used in combination with GA3 O.Tuzcu and A.I.
rind color as a full (10 mg∙l–1) Ozguven
change tree cover-
use in morning or in late
age
afternoon
avoid rainy days
compatible with most nutri-
tional and pesticide sprays.
USA (California) Grapefruit when fruit of 75 g∙ha–1 in isopropyl ester C.W. Coggins, Jr.
the following spray up to
may also result in improved
crop are at 5000 1∙ha–1
fruit size for next year’s crop
least 19 mm in to mature
diameter trees also reduces fruit stem die-
back
do not apply within 7 days of
harvest
Lemon Oct.-Dec. 37 g∙ha–1 in application to coastal lemons “
spray up to are risky because of their
5000 1∙ha–1 ever-bearing nature
to mature
use a single treatment
trees
do not apply within 7 days of
harvest
Minneola tangelo Sept.-Nov. 75 g∙ha–1 in use a single treatment “
spray up to
do not apply within 7 days of
5000 1∙ha–1
harvest
to mature
trees do not apply to trees< 6
years old
Navel orange Sept.-Dec. 37-75 g∙ha– isopropyl ester “
1
highest
do not apply within 7 days of
rates ap-
harvest
plied early
in the sea- do not apply to trees< 6
son; apply years old
solution up
55

to 5000
1∙ha–1
Valencia orange when fruit of 37 g∙ha–1 in isopropyl ester “
the following spray up to
may also result in improved
crop are at 5000 1∙ha–1
fruit size for next year’s crop
least 13 mm in to mature
diameter trees also reduces fruit stem die-
back
do not apply within 7 days of
harvest
USA (Florida) Navel orange 6-8 weeks 2,4-D at 70 reduces summer-fall drop D.P.H. Tucker
after bloom or g∙ha–1 with and T.A. Wheaton
do not apply later spray when
Aug.-Sept. for spray at
fruit is to be harvested early
fall drop con- 2500-5000
trol 1∙ha–1 avoid period of leaf flush
isopropyl ester of 2,4-D used
Temple orange, Nov.-Dec. 95 g∙ha–1 isopropyl ester of 2,4-D used “
Grapefruit with spray
avoid periods of leaf flush
at 2500-
5000 1∙ha–1
56

Table 10. Use of ethephon to aid in fruit harvesting.

Country Cultivar Time/stage of Rate of Remarks Source


application application
a. Induction of fruit loosening to facilitate manual harvesting
Cuba Dancy mandarin, one week 2.5 kg∙ha–1 facilitates separation of fruit from C. Noriega,
Orlando tangelo before ex- using an 800 stem M. del C.
pected start of mg∙l–1 solu- Perez, and
reduces plugging damage to fruit
harvest of tion as a full L. Pozo
mature fruit tree coverage
Eureka lemon one week 3-4 kg∙ha–1 ensure optimum irrigation and nutri- M. del C.
before ex- using a 1000 tion of groves Perez,
pected start of mg∙l–1 solu- L. Pozo, and
add Ca(OH)2 at 0.1% to prevent
harvest of tion as full C. Noriega
excess defoliation
mature fruit tree coverage
with 8-10 l apply in morning to avoid afternoon
per tree rain
fruit for domestic market
picker efficiency is more than dou-
bled
fruit develop good color
do not use in groves where
yields<12 t∙l–1
Valencia orange one week 3.8 kg∙ha–1 ensure optimum irrigation and fer- L. Pozo, M.
before ex- using a 1200 tilization of groves del C. Perez,
pected time of mg∙l–1 solu- C. Noriega
add Ca(OH)2 at 0.1% to prevent
harvest tion as full
excess defoliation
tree coverage
apply treatments in morning to avoid
afternoon rain
do not use in groves where yields <
12 t∙l–1
b. Induction of fruit abscission for mechanical harvesting
Cuba Marsh grapefruit preharvest of 1.3-2.6 concentration depends on fruit at- M. del C.
fully mature kg∙ha–1 tachment force Perez,
fruit using a L. Pozo, and
add Ca(OH)2 at 0.1% to avoid excess
(Oct-Dec.) 600-1200 C. Noriega
leaf abscission
mg∙l–1 solu-
tion used as ensure optimum nutrition and irriga-
a full tree tion in groves
coverage
apply in morning to avoid afternoon
rain
Valencia orange preharvest of 3.2-5.7 concentration depends on fruit at- L. Pozo, M.
fully mature kg∙ha–1 tachment force del C. Perez,
fruit 1000-1800 C. Noriega
add Ca(OH)2 at 0.1% to avoid excess
(Dec.-Feb.) mg∙l–1 solu-
leaf drop
tion as full
tree cover- ensure optimum nutrition and irriga-
age tion of groves
57

apply in morning to avoid afternoon


rain
58

Table 11. Postharvest application (aqueous dip or in wax) of 2,4-D to reduce postharvest decay and maintain fruit qual-
ity.

Country Cultivar Rate of Remarks Source


application
Australia Navel 500 mg∙l–1 can be used with fungicides such as benlate P.T. Gallasch
orange
applied as dimethylamine salt
Israel All citrus 250 mg∙l–1 Applied as Na or dimethylamine salt E.E. Goldschmidt, Y.
Erner and J. Green-
no pH control
berg
often mixed with TBZ* or IMZ*
Morocco Clementine 4-8 mg∙l–1 fruit dipped in treatment solution in the grove at M. El-Otmani
mandarin harvest
use only for fruit to be degreened
butyl glycol ester
delays button abscission and maintains green
button (calyx)
South Africa All citrus 500 mg∙l–1, Applied as Na or dimethylamine salt I. Gilfillan
dip fruit
Maintains button green, reduces Diplodia decay
for 1 min
no pH control
often mixed with IMZ* or TBZ*
Kutdiken 200 mg∙l–1 Applied in storage wax O. Tuzcu, A.I. Ozgu-
lemon ven and O. Dundar
Keeps button (calyx) alive
fruit stored for up to 8 months in cold rooms,
essentially for export
Turkey Marsh 200 mg∙l–1 Improves button (calyx) viability “
grapefruit
Reduces susceptibility to decay such as Alternaria
fruit stored for up to 6 months in cold rooms,
essentially for export
Uruguay All citrus up to 500 Isopropyl ester formulation J.C. Codina
for export mg∙l–1
USA (California) Lemon up to 500 Isopropyl ester formulation C.W. Coggins, Jr.
mg∙l–1
*
IMZ=imazalil; TBZ=thiabendazole.
59

APPENDIX I

Species Names

SCIONS

Citron (Citrus medica) (Saunt, 2000)

Clementine mandarin (Citrus reticulata) (Reuther et al., 1967; Saunt, 2000):


Includes –
Dancy
Ellendale
Fina
Fortune
Kinnow
Marisol
Nova
Nules
Ponkan
Wilking

Grapefruit (Citrus paradisi)(Reuther et al., 1967; Saunt, 2000):


Includes –
Marsh
Oroblanco

Kumquat (Fortunella species) (Reuther et al., 1967)

Lemon (Citrus limon)(Reuther et al., 1967):


Includes –
Eureka
Kutdiken

Lime (Citrus latifolia)(Reuther et al., 1967):


Includes –
Bearss
Persian
Tahiti

Satsuma mandarin (Citrus unshiu) (Reuther et al., 1967; Saunt, 2000):


wase means early group and includes Clauselline, Miho, Miyagawa, Okitsu

Sweet orange (Citrus sinensis)(Saunt, 2000):


Includes –
Bonanza navel
Hamlin
Jaffa
Natal
Navel
Navelate
Pera
Shamouti
Tuligold navel
Valencia
Washington navel
Westin
60

Tangelo (Citrus paradisi x C. reticulata) (Saunt, 2000):


Includes –
Minneola and Orlando, both hybrids of Duncan grapefruit and Dancy mandarin (tangerine)

Tangor (Citrus sinensis x C. reticulata) (Saunt, 2000):


Murcott
Ortanique

Temple orange/Temple mandarin (Citrus temple) (Saunt, 2000):

ROOTSTOCKS

Citremon 1452 (Poncirus trifoliata x C. limon) (Saunt, 2000)


Rough lemon (Citrus jambiri) (Reuther et al., 1967)
Smooth flat seville (An old Australian cultivar, generally regarded as a sweet orange and grapefruit hybrid) (Saunt, 2000)
Sour orange (Citrus aurantium) (Reuther et al., 1967; Saunt, 2000):
Trifoliate orange (P. trifoliata) (Reuther et al., 1967)
Troyer citrange (P. trifoliata x C. sinensis) (Saunt, 2000)
Volkameriana (Citrus volkameriana), Volkamer lemon (Reuther et al., 1967)
Yuma citrange (P. trifoliata x C. sp.)
61

APPENDIX II

Source Contact Information

Manuel Agustí
Universidad Politécnica
Camino del Vera, 14
Valencia 46022
Spain

Tania Castro-Lopez
Instituto de Investigación de Cítricos
Ave. 7 ma #3005 e/30y32
Mirama, Playa
La Habana
Cuba

Juan Carlos Codina


Urudór, S.A.
Misiones 1537, Esc. 502
33 Orientales 766
Montevideo 11.000
Uruguay

Charles W. Coggins, Jr.


Department of Botany and Plant Sciences
University of California
Riverside, CA 92521-0124
USA

Paulo Roberto Camargo de Castro


Botànica – ESALQ
Escola Superior de Agricultura “Luiz de Queiroz”
Caixa Postal, 9
13.418-900 Piracicaba, SP
Brazil

Emanuela de Martino Aleppo


Citrus Experimental Institute
Corso Savoia 190
95024 Acireale (Catania)
Italy

O. Dundar
Faculty of Agriculture
Department of Horticulture
Cukurova University
Balcali 01330
Adana
Turkey

Mohamed El-Otmani
Institut Agronomique et Vétérinaire Hassan II
B.P. 18/S
80000 Agadir
Morocco
62

Yair Erner
Department of Citriculture
ARO, The Volcani Center
P.O.B. 6
Bet-Dagan 50250
Israel

Peter T. Gallasch
South Australian Research and Development Institute
P.O. Box 411
Loxton, SA 5333
Australia

Ian Gilfillan
P.O. Box 27656
Greenacres
Port Elizabeth 6057
South Africa

Eliezer Goldschmidt
Herbrew University of Jerusalem
P.O. Box 12
Rehovot 76100
Israel

A. Gravina
Facultad de Agronomía
Cátedra de Fisiolgía Vegetal
Av. Garzon 780
12900 Montevideo
Uruguay

Joseph Greenberg
P.O.B. 416
Haniel 42865
Israel

F. Guerra
Instituto de Investigación de Cítricos
Ave. 7 ma #3005 e/30y32
Mirama, Playa
La Habana
Cuba

Andrew Harty
KeriFresh Group
Waipapa Road, RD2
Kerikeri
New Zealand

Isao Iwagaki
Shizuoka University
Karijuku, Fujieda
Shizuoka 426-001
Japan

I. Izquierdo
Instituto de Investigación de Cítricos
63

Ave. 7 ma #3005 e/30y32


Mirama, Playa
La Habana
Cuba

Aldo Malavasi
Departmento de Biologia
Instituto de Biociências
Universidade de São Paulo
Rua de Matão, Travessa 14, n. 321
Caixa Postal 11461
05422-970 - São Paulo
São Paulo, SP
Brazil

Caridad Noriega
Instituto de Investigación de Cítricos
Ave. 7 ma #3005 e/30y32
Mirama, Playa
La Habana
Cuba

Ahsen Isik Ozguven


Faculty of Agriculture
Department of Horticulture
Cukurova University
Balcali 01330
Adana
Turkey

Maria del Carmen Pérez


Instituto de Investigación de Cítricos
Ave. 7 ma #3005 e/30y32
Mirama, Playa
La Habana
Cuba

Francisco Pividal
Instituto de Investigación de Cítricos
Ave. 7 ma #3005 e/30y32
Mirama, Playa
La Habana
Cuba

Luís Pozo
Instituto de Investigación de Cítricos
Ave. 7 ma #3005 e/30y32
Mirama, Playa
La Habana
Cuba

Miguel Ragone
EEA-INTA-Concordia
CC34
3200 Concordia EE
Argentina

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