10.1007@s00344 015 9541 6

J Plant Growth Regul
DOI 10.1007/s00344-015-9541-6
Plant Growth Regulators: Backgrounds and Uses in Plant

Production
Wilhelm Rademacher1,2
Received: 6 May 2015 / Accepted: 9 August 2015

Ó Springer Science+Business Media New York 2015
Abstract Plant growth regulators are widely used in Introduction

modern agriculture, horticulture and viticulture. ‘‘True’’
plant growth regulators interfere directly with the plant’s Plant growth regulators (PGRs) can be defined as naturally
hormonal status. They are represented by plant hormones occurring or synthetic compounds that affect develop-
or their synthetic analogs, by inhibitors of hormone mental or metabolic processes in higher plants, mostly at
biosynthesis or translocation and by hormone receptor low dosages. They do not possess a nutritive value and,
blockers. ‘‘Atypical’’ plant growth regulators act by dis- typically, are not phytotoxic. The name ‘‘plant growth
playing a local and/or transient phytotoxic effect. substances’’ may equally be used. Newcomers might
Approximately 40 active ingredients are in use, applied associate such compounds primarily with an increase or
either as a single component or as combinations. Many reduction in longitudinal plant growth. Because a multitude
plant developmental processes can be actively regulated in of other processes can also be affected, such as flowering,
cultivated plants, for example, acceleration or delay of seed fruit formation, ripening, fruit drop, defoliation, or quality
germination, dormancy breaking in woody perennials, traits, the term ‘‘plant bioregulator’’ would be an even
stimulation or reduction of shoot elongation, induction of better choice and is used as well. Because of more common
flowering and fruiting, reduction or increase of fruit set, practice, ‘‘plant growth regulators’’ will be referred to in
acceleration or delay of senescence processes including this contribution. A major group among PGRs are com-
fruit ripening and defoliation. The achieved benefits range pounds, which reduce shoot elongation. Such substances
from facilitating crop management to increasing and are often named ‘‘growth retardants.’’ Different from
securing yield and quality of the harvested produce and PGRs, ‘‘plant biostimulants’’ comprise complex mixtures
improving its storage and shelf life. Systematic use of plant of ingredients and are often based on extracts of seaweeds
growth regulators started in the 1930s. Current global or hydrolysates of slaughterhouse leftovers. Such prepara-
annual sales are in the range of US$ 1.2 billion. tions may contain PGR-like components.
In the majority of cases, PGRs affect the balance of
Keywords Plant growth regulator Plant bioregulator plant hormones in treated plants. This can primarily be
Growth retardant Plant hormone Synthetic analog achieved by applying exogenously naturally occurring
Biosynthesis inhibitor Receptor blocker hormones or their synthetic analogs, by inhibiting the
biosynthesis of endogenous hormones or their translocation
from the site of production to the site of action and by
blocking hormone receptors. Plant hormones are involved
in many processes, which allow plants to react on internal
& Wilhelm Rademacher
[email protected]
and external stimuli. The ‘‘classical’’ groups of plant hor-
mones are the auxins, gibberellins, cytokinins, abscisic
1
Global Research Crop Protection, BASF SE, acid, and ethylene. To date, brassinosteroids and jas-
67114 Limburgerhof, Germany monates should also be regarded as having phytohormonal
2
Austrasse 1, 67117 Limburgerhof, Germany functions, whereas the classification of polyamines and
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some other compounds is still controversial. Plant hor- In the global crop protection market of approximately
mones can be formed at different cytological or morpho- US$ 56.7 billion (€ 42.7 billion) of annual sales in 2014
logical sites. They may act in the tissue of production or are (Anonymous 2015), PGRs play a relatively small role and
translocated to more remote target tissues. Their main sites account for only some 2.5 % (fungicides used as PGRs
of production are meristems and growing fruits. Plant included, herbicide-based defoliants and desiccants exclu-
hormones, in particular auxins, may cause diametrically ded) (Rademacher 2010a). However, PGRs often gain a
opposed effects on a given process at different concentra- higher interest for the cultivation of distinct plant species
tions. Synergistic or antagonistic interactions among the and/or when plants are grown under high degrees of
different hormone groups add to the complexity of the intensity. Only few companies are still engaged in finding
hormonal system in higher plants. Because of this, affect- and developing new PGRs. Investments of more than US$
ing plant performance in a desired way with PGRs can be a 240 million are required for research, development, and
difficult undertaking. registration prior to the market introduction of a new crop
PGRs are employed in agriculture, horticulture, and protectant (Anonymous 2013). Cost reductions are possible
viticulture to obtain specific advantages, such as decreased only in few cases, for instance, when the respective active
susceptibility towards biotic and abiotic stress, improved ingredient is of natural origin (for example, a non-toxic
morphological structure, facilitation of harvesting, quan- ‘‘biochemical’’ according to US legislation). Because
titative and qualitative increases in yield, and modification PGRs are significantly more complex to work with than
of plant constituents. Neljubow (1901) can be credited to other crop protectants, even higher investments have to be
be the first to discover that a distinct compound—later expected. In most cases, herbicides, fungicides, or insec-
defined a plant hormone—was biologically active in ticides appear better suited for investments. To be eco-
higher plants. He demonstrated that ethylene causes hori- nomically viable, a new PGR must have a sufficiently large
zontal growth of pea seedlings, inhibition of elongation market and be appropriately covered by patents as pre-
and radial swelling (the ‘triple response’) (Bakshi and requisites to offer sufficiently attractive profitability. The
others 2015 this issue). Also in these early days, Denny industry has to concentrate its efforts on uses in ‘‘big
(1924) demonstrated that ethylene was the active compo- crops’’, such as wheat, rice, maize, sugar cane, soybeans,
nent from kerosene stoves that promoted ripening of cotton, oilseed rape, apples, citrus fruits, grapevines, and
lemon fruits. With this and additional knowledge, one can bananas. Developing a PGR for a minor crop is, typically,
now explain that ethylene was involved in practices such not attractive for a company and will not be actively pur-
as wounding or gassing with smoke, which were used sued. To overcome this obstacle, minor uses can be made
since ancient times to accelerate fruit ripening and possible in most countries under certain regulations.
improve fruit quality (for example, Abeles and others However, a prerequisite for approaches of this type is a
1992). A systematic use of PGRs began in the 1930s, valid registration of the candidate product in at least one
when ethylene and related acetylene were used to induce major crop.
flowering and fruit formation in pineapple (Rodriguez PGRs are typically applied via foliar sprays with water
1932; Bartholomew 2014). Since then, a broad spectrum as a carrier. PGR performance does not only depend on the
of PGRs has been detected and many PGR products have active ingredient used. Proper conditions for the compound
become indispensable parts of modern plant husbandry. to reach its biochemical target have to be provided. In order
Many targets for PGRs are also goals for breeding. to penetrate the plant’s epidermis and membranes, appro-
However, in spite of the possibilities that conventional priate formulants (for example, solvents, surfactants and
breeding and genetic engineering offer, PGRs often enable water conditioners) have to be present in the commercial
faster and better solutions to many problems. Most product. Adequate consideration has also to be given to
importantly, they allow for an active regulation of plant treat plants at their right stage of development. Finally,
processes, thereby adjusting a given genotype to its par- weather conditions before, during, and after treatment can
ticular growing conditions. In other words, plant growth be critical for success.
regulators can be used highly flexibly for the ‘‘fine-tun- A broad range of chemicals with PGR activities are
ing’’ of crop plants that grow at a given location under known. Only those compounds and their derived products
largely uncontrollable and unpredictable environmental will be regarded in this contribution, which are of practical
conditions. Overviews covering different aspects of PGRs relevance in key markets such as the EU or the US. The
have, inter alia, been presented by Weaver (1972), names of active ingredients used for PGR products, like
McLaren (1982), Nickell (1983), Macgregor and others those for other crop protectants or pharmaceuticals, are
(1986), Schott and Walter (1991), Basra (2000), Petracek numerous and confusing. Only the common name
and others (2003), Rademacher (2004), and Rademacher unequivocally identifies a distinct compound, of which
and Brahm (2010). different forms (salts, isomers, and so on) may still exist.
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Common names are assigned by the International Organi- connected with many effects caused by auxins (Abeles and
zation for Standardization (ISO) and are short, distinctive, others 1992).
non-proprietary, and widely accepted. Because of this, the
common name is primarily referred to. Systematic chemical Natural and Synthetic Auxins
names, mostly based on the nomenclature of the Interna-
tional Union of Pure and Applied Chemistry (IUPAC), and Naturally occurring indol-3-ylacetic acid and other auxins
commonly used abbreviations are given in the captions for are employed in tissue cultures for plant propagation.
chemical structures. Likewise, abbreviations are used Auxins are also in use to stimulate rapid and prolific
sparingly. More information for distinct active ingredients rooting of cuttings of a wide range of trees, vines and
can be found, inter alia, in the internet on the database of shrubs, and annual and perennial ornamentals, for example,
pesticide physicochemical and ecotoxicological data pro- indol-3-ylbutyric acid (Fig. 1I), which is also occurring
vided by the University of Hertfordshire in the UK under: naturally, or 1-naphthylacetic acid (Fig. 1II). The latter
http://sitem.herts.ac.uk/aeru/iupac/ (accessed July 26, compound and 2-(1-naphthyl)acetamide (Fig. 1III) are
2015). A large number of trade names may also exist for applied at higher dosages for flower or fruitlet thinning in
products containing a given compound, which may relate to apples and pears. Lower dosages of these compounds and
different producers, different formulations or combinations of 2,4-D (2,4-dichlorophenoxyacetic acid) and triclopyr
with other active ingredients. Therefore, only the original {[(3,5,6-trichloro-2-pyridinyl)oxy]acetic acid} may be
and most widely used ones will primarily be used. Likewise, used to improve fruit setting or preharvest fruit retention.
space allows only for mentioning key producers or provi- Carbaryl (Fig. 1IV) is another synthetic auxin used for fruit
ders. Reference is also given to the scientific and industrial thinning. Auxins are also in use for improving fruit set in
‘‘roots’’ only of the more prominent PGRs. Quotations, tomatoes and other members of the nightshade family. 2,4-
particularly of older original literature, had to be restricted. D and other synthetic auxins such as mecoprop, dicamba,
The reader may find such contributions in cited review or quinclorac are primarily used as herbicides (Cobb and
articles. Many agrochemical companies have been involved Reade 2010).
in the finding and developing of PGRs. As a result of After the detection of indol-3-ylacetic acid as a natural
acquisitions and mergers, many of these companies do not auxin in the mid-1930s, plant physiologists and chemists
exist anymore. A summary of company ‘‘genealogy’’, pre- identified additional compounds with similar biological
pared and maintained primarily for US herbicide companies activities (Lüthen 2015, this issue). The first publication on
by A.P. Appleby from Oregon State University is helpful to indol-3-ylbutyric acid and 1-naphthylacetic acid is from
understand also the history of PGR companies [http://cro Zimmerman and others (1936). Numerous other chemicals
pandsoil.oregonstate.edu/herbgnl (accessed July 26, 2015)]. with auxin-like properties were synthesized in the follow-
With regard to giving credit to scientists and companies ing years. The potential of using such chemicals as PGRs
initiating work on particular PGRs, the author asks for was soon recognized, for example, by Gardner and others
pardon for any omissions and inaccuracies. (1939) and Zimmerman and Hitchcock (1944). Market
introduction of 1-naphthylacetic acid and 2-(1-naph-
thyl)acetamide was by Amchem Products, Inc. in 1949 and
‘‘True’’ Plant Growth Regulators 1953, respectively. Carbaryl was introduced in 1956 by
Union Carbide Corp. None of the companies mentioned
‘‘True’’ PGRs are compounds, which interfere directly and exists anymore, and products based on these auxins are
specifically with the natural hormone system of higher now provided by other companies. Regrettably, auxin-type
plants. herbicides/PGRs lost their innocence, when they were used
for military purposes. In particular, 2,4,5-trichlorophe-
Compounds Related to Auxins noxyacetic acid, contaminated with extremely toxic
2,3,7,8-tetrachlorodibenzo-p-dioxin has to be mentioned,
Auxins stimulate cell elongation and cell division. They are which was used as a defoliant during the Vietnam War
involved in phototropic and gravitropic reactions. Higher (Stellman and others 2003).
concentrations may inhibit such reactions and give the
opposite effect. Apical dominance is based on the fact that Inhibitor of Auxin Transport
auxin produced in the apex bud inhibits lateral buds from
growing. Auxins play also a role in attracting assimilates, Cyclanilide (Fig. 1V) acts by inhibiting the transport of
for example, for storage of starch in growing cereal seeds. natural auxins and possibly also their receptor binding
Synthetic auxins are often more persistent than the natu- (Burton and others 2008). In 1995, Rhône-Poulenc S.A.
rally occurring ones. The induction of ethylene formation is started to introduce a combination of this compound with
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Fig. 1 Auxins: I Indol-3-ylbutyric acid (IBA), II 1-naphthylacetic auxin translocation and binding: V cyclanilide = 1-[(2,4-
acid (NAA), III 2-(1-naphthyl)acetamide (NAAm), and IV car- dichlorophenyl)carbamoyl]cyclopropanecarboxylic acid
baryl = 1-naphthyl hydrogen methylcarbonimidate. Inhibitor of
ethephon as a cotton harvest-aid in the US (Fritz 1997). and Pfizer, Inc. in the US and to Takeda Chemical Indus-
Bayer Crop Science has been selling this product since tries in Japan (Martin 1983; Petracek and others 2003).
2002. A combination with mepiquat chloride for growth Syngenta Crop Protection (which now contains the agro-
regulation in cotton is also on the market. From 2009 until chemical part of ICI) is still a distributor of GA prepara-
2011, cyclanilide has been commercially available as a tions. Valent BioSciences Corp. emerged from Abbott
solo product in the US under the trade name TiberonÒ Laboratories and is now part of Sumitomo Chemical Co.,
(Bayer CropScience) for the promotion of lateral branching Ltd. In addition to selling GA-containing products, Valent
(‘‘feathering’’) in nursery trees. However, due to too small BioSciences is actively engaged in finding new uses and
a market, the product had to be discontinued. improved formulations for GAs.
G. fujikuroi is used to produce GA3 (gibberellic acid)
Gibberellins and Inhibitors of Gibberellin and a mixture of GA4 and GA7 (Fig. 2I–III) by fermenta-
Biosynthesis tion. A chemical synthesis of GAs is possible (Mander
2003), but it is too complex and expensive for any com-
Work on gibberellins (GAs) started in the 19th century in mercial exploitation. Likewise, testing other GA-producing
Japan and was initially focusing on the name-giving phy- fungi did not result in commercially viable alternatives. For
topathogenic fungus Gibberella fujikuroi (imperfect stage instance, cultures of the fungus Sphaceloma manihoticola,
Fusarium moniliforme—meanwhile corrected into F. fuji- which causes the superelongation disease of cassava, yield
kuroi), which causes abnormal shoot elongation in rice mainly GA4 but no GA1, GA3, or GA7 (Rademacher and
(‘bakanae’ disease) (Hedden and Sponsel 2015, this issue). Graebe 1979). Attempts made at ICI, Norsk Hydro ASA
Due to language barriers and times of war, the primary and BASF to produce pure GA4 with this fungus failed
work of Japanese scientists became known in other coun- because of difficulties to establish competitive large-scale
tries only around 1950. The first work on GAs in the US fermentations (Rademacher unpublished). Currently, the
was even performed at the Biological Warfare Center at majority of GAs sold globally originates from China. In
Fort Detrick, Maryland (Mitchell and Angel 1950). Com- terms of sales, GA3 is among the top-ranking PGRs. It has
panies in the UK and in the US started to work on pro- found many applications in viticulture, horticulture, and
ducing GAs by fermentation and testing them on crop agriculture (Weaver 1972; Carlson and Crovetti 1990;
plants. Research at the Akers Laboratories of Imperial Petracek and others 2003) and also in the beer industry,
Chemical Industries (ICI) in the UK on fermentation and where it is used to accelerate malting (Hornsey 2003).
chemical identification yielded the first production patents Total global annual use of GA3 is in the range of 100 tons
in 1954. Parallelly, agricultural uses for GAs were inves- with approximately three quarters of this used in plant
tigated at ICI’s Jealott’s Hill research facility. By late 1957, production, the rest for malting.
ICI had granted manufacturing and use licenses to Abbott Almost 140 different GAs are presently known to occur
Laboratories, Inc., Merck and Co., Inc., Eli Lilly and Co., in higher plants and/or gibberellin-producing fungi.
123
Fig. 2 Structures of I gibberellin A3 = gibberellic acid = GA3 = gibberellin A7 = GA7 = (3S,3aR,4S,4aR,7R,9aR,9bR,12S)-12-hy-

(3S,3aR,4S,4aS,7S,9aR,9bR,12S)-7,12-dihydroxy-3-methyl-6-methy- droxy-3-methyl-6-methylene-2-oxoperhydro-4a,7-methano-9b,3-
lene-2-oxoperhydro-4a,7-methano-9b,3-propeno[1,2-b]furan-4-carbo- propenoazuleno[1,2-b]furan-4-carboxylic acid. Note that the IUPAC
xylic acid, II gibberellin A4 = GA4 = (3S,3aR,4S,4aR,7R,9aR,9bR, nomenclature used here employs a different numbering of carbon
12S)-12-hydroxy-3-methyl-6-methylene-2-oxoperhydro-4a,7-meth- atoms than the often used ‘‘biological’’ numbering, which refers to
ano-3,9b-propanoazuleno[1,2-b]furan-4-carboxylic acid, and III ent-gibberellane
However, only a few of the GAs possess biological activity berry thinning of wine and table grapes, in improving citrus
per se, whereas the majority are precursors or catabolites. fruit quality, in increasing fruit size in pears and sweet
The main hormonal functions of GAs in higher plants are cherries, and in accelerating seed germination. The main
the promotion of longitudinal growth, the induction of use of GA4/7 is to reduce fruit russetting in apples. Com-
hydrolytic enzymes in germinating seeds, the induction of bined with the cytokinin 6-benzyladenine, GA4/7 is also
bolting in long-day plants, and the promotion of fruit set- used to improve the quality of apple fruits.
ting and development. The biosynthesis of GAs in higher
plants can be separated into three stages according to the Inhibitors of GA Biosynthesis
nature of the enzymes involved and the corresponding
localization in the cell, as follows: Terpene cyclases acting Inhibitors of GA biosynthesis (growth retardants), which
in proplastids, monooxygenases associated with the endo- reduce longitudinal shoot growth, represent the most
plasmic reticulum, and dioxygenases located in the cytosol. important group of PGRs, both in terms of commercial
These enzymes catalyze the formation of ent-kaurene from value and in terms of treated area. In intense European
D-glyceraldehyde 3-phosphate plus pyruvate, the oxidation cereal and oilseed rape production and Japanese and South
of ent-kaurene into GA12-aldehyde, and the subsequent Korean rice cultivation, they are applied to reduce the risk
hydroxylations into different GAs, respectively. Details on of lodging, which often results from severe rainfall and/or
GA biosynthesis can be found in Hedden and Thomas wind; in cotton excessive vegetative growth can be con-
(2012) and Hedden (2016). Several PGRs act by inhibiting trolled; fruit trees can be kept more compact, thereby
gibberellin biosynthesis (Rademacher 2000, 2016). reducing costs for pruning and obtaining a better ratio
between vegetative growth and fruit production; the quality
Naturally Occurring GAs of ornamental and bedding plants is generally improved by
keeping them compact, which also reduces the space in a
GA3 is the most widely used GA. Additionally, a mixture greenhouse required for production; costs for trimming
of GA4 and GA7 is commercially available. GA4 is less hedges and ornamental trees and for mowing turf grasses
persistent than GA3 and GA7 and is, therefore, better suited may also be reduced by applying inhibitors of GA
when effects lasting too long are unwanted. However, due biosynthesis.
to the close chemical similarity of GA4 with GA7, their The different types of growth retardant known to inter-
separation in fermentation extracts is very difficult. As a fere with different stages of GA biosynthesis comprise the
result, the content of GA7 in different products varies following (Rademacher 2000): (a) Chlormequat chloride
between approximately 40 % and insignificant amounts. A (Fig. 3I) and mepiquat chloride (Fig. 3II) are quaternary
preparation, particularly low in GA7, is from Fine Agro- ammonium compounds that inhibit cyclases involved in
chemicals Ltd. early stages of GA metabolism, which lead to the forma-
There are numerous uses of GAs. Valent BioSciences tion of ent-kaurene. (b) Several compounds with a nitro-
Corp., a leading developer and provider of GAs, is listing gen-containing heterocycle block cytochrome P450-
more than 40 different plant species, in which their seven dependent monooxygenases, which catalyze the oxidation
GA preparations can be utilized. GA3 has found many of ent-kaurene via ent-kaurenoic acid into GA12-aldehyde.
applications in viticulture, horticulture, and agriculture, for The pyrimidines ancymidol (Fig. 3III) and flurprimidol
example, in the production of seedless table grapes, in (Fig. 3IV) and the triazoles paclobutrazol (Fig. 3V),
123
Fig. 3 Structures of inhibitors of GA biosynthesis: I Chlormequat pentan-3-ol, VI uniconazole-P = (1E,3S)-1-(4-chlorophenyl)-4,4-

chloride = chlorocholine chloride (CCC) = (2-chloroethyl)-trime- dimethyl-2-(1H-1,2,4-triazol-1-yl)-1-penten-3-ol, VII daminozi-
thylammonium chloride, II mepiquat chloride = 1,1-dimethylpiperi- de = N-(dimethylamino)succinamic acid, VIII trinexapac-ethy-
dinium chloride, III ancymidol = cyclopropyl-(4-methoxyphenyl)-5- l = ethyl (RS)-4-cyclopropyl(hydroxy)methylene-3,5-dioxocyclohex
pyrimidinylmethanol, IV flurprimidol = 2-methyl-1-(5-pyrimidinyl)- anecarboxylate, and IX prohexadione-calcium = calcium 3-oxido-5-
1-[4-(trifluoromethoxy)phenyl]-1-propanol, V paclobutrazol = oxo-4-propionylcyclohex-3-enecarboxylate
(2RS,3RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1H-1,2,4-triazol-1-yl)
uniconazole-P (Fig. 3VI), tebuconazole [1-(4-chlor- compounds by inhibiting the conversion of 1-aminocyclo-
ophenyl)-4,4-dimethyl-3-(1H-1,2,4-triazol-1-ylmethyl)-3- propane-1-carboxylic acid (ACC) into ethylene. This
pentanol], and metconazole [5-(4-chlorobenzyl)-2,2-dime- reaction is catalyzed by ACC oxidase. These dioxygenases
thyl-1-(1H-1,2,4-triazol-1-ylmethyl)cyclopentanol are of require 2-oxoglutarate and ascorbate as their respective co-
practical relevance. Tebuconazole and metconazole are substrates and it is likely that, because of structural simi-
primarily used as fungicides, where they function as inhi- larities, the PGRs displace the co-substrates from the
bitors of P450-dependent monooxygenases involved in enzymes’ active sites. The free acid forms prohexadione
fungal ergosterol formation. In oilseed rape and some other and trinexapac are required to block the involved dioxy-
plant species, they are also efficient inhibitors of GA genases. Whereas the free acid prohexadione is readily
biosynthesis. (c) The late stages of GA biosynthesis are generated by dissolving its calcium salt in water, the ester
catalyzed by dioxygenases, which require 2-oxoglutaric trinexapac-ethyl has to be saponified by plant metabolism
acid as a co-substrate. Because of their structural rela- prior to becoming active.
tionship to 2-oxoglutaric acid, daminozide (Fig. 3VII) and Several compounds with a positively charged ammo-
the acylcyclohexanediones trinexapac-ethyl (Fig. 3VIII) nium, phosphonium, or sulphonium group are known as
and prohexadione-calcium (Fig. 3IX) inhibit these reac- plant growth retardants since the 1940s (Mitchell and
tions. However, the latter two compounds are significantly others 1949; Cathey 1964; Rademacher 2000). However,
more specific and active. In addition to blocking dioxy- only few of these were developed into practice. Although
genases involved in GA formation, prohexadione-calcium chlormequat chloride was first synthesized by Kauffmann
and trinexapac-ethyl inhibit also flavanone 3-hydroxylase, and Vorländer (1910), it took 50 years until it was
which is involved in the biosynthesis of flavonoids. Fur- described by Tolbert from Michigan State University in
thermore, ethylene formation is reduced by both East Lansing to reduce shoot length in several plant species
123
(Tolbert 1960a, b; Lindstrom and Tolbert 1960; Wittwer registrations in plants used for human or animal nutrition.
and Tolbert 1960). The commercial rights were then held Paclobutrazol was first reported as a new and very potent
by American Cyanamid Company, who obtained a regis- PGR by Lever and others (1982). Market introduction by
tration for CycocelÒ by the US Department of Agriculture ICI Agrochemicals (via Zeneca Agrochemicals now part of
in 1962. Uses were restricted to ornamental plants grown in Syngenta Crop Protection) was in 1986. The first interna-
greenhouses, since American Cyanamid did not see any tional publication on uniconazole was by Izumi and others
potential in other areas of plant production at that time. (1984). It was introduced to the market by Sumitomo
However, the significance of using chlormequat chloride as Chemical Co., Ltd. in 1985. The closely related paclobu-
an anti-lodging agent in intense European wheat produc- trazol, uniconazole (RS-racemate), and uniconazole-P
tion was soon recognized in Austria and Germany. (pure S-isomer—introduced later to replace the racemate)
Whereas Österreichische Stickstoffwerke AG developed are very persistent, with an average half-life of approxi-
the compound as a component of fertilizer granules (Linser mately 6 months, both in plants and in the soil. They are
and others 1961; Barbier and Mayr 1966), BASF concen- used particularly in countries with warmer climates to
trated its activities on foliar applications of aqueous solu- control vegetative growth of fruit trees such as avocados,
tions (Jung and Sturm 1964). The latter turned out to be the mangos, or litchis. Lodging control in rice and the pro-
better approach. Consequently, CycocelÒ was introduced duction of more compact ornamentals are further uses of
under license by BASF in Germany in 1965 and, thereafter, these compounds. In addition to blocking ergosterol
in many other countries. After 50 years, this growth biosynthesis in fungi, some triazole-type fungicides may
retardant is still the most widely used PGR in cereal pro- also block GA biosynthesis in distinct plant species. This is
duction, particularly in wheat, rye, triticale, and oats. of practical interest for using tebuconazole and metcona-
Together with uses in other cultivated plants, it is, in terms zole to control shoot growth in oilseed rape. In Germany,
of treated area, the number one PGR on a global scale. metconazole is additionally in use as a PGR for orna-
After having success with chlormequat chloride, BASF mentals. Kuck and Berg (1986) from Bayer AG reported
developed another quaternary ammonium compound, first on tebuconazole, which was commercialized as a
mepiquat chloride (Zeeh and others 1974). After its intro- fungicide starting in 1988. Metconazole was discovered by
duction in 1979, mepiquat chloride became a very suc- Kureha Chemical Industry Co., Ltd. in 1986 with the first
cessful PGR, particularly as PixÒ in cotton. Alone or in international report by Sampson and others (1992). The
combination with other PGRs, it is nowadays used in compound was jointly developed with Shell International
several crops. Mepiquat has been detected in samples of and, later, American Cyanamid Co. and BASF. Market
processed plant material, such as roasted coffee beans, introduction (first as a fungicide, then additionally as a
roasted barley seeds, crust of bread, and alfalfa pellets, PGR) was by Cyanamid Agro in France in 1994.
even if any previous use of mepiquat chloride or other Riddell and others (1962) were the first to publish on the
sources of chemical contamination could be ruled out. plant growth-regulatory activity of daminozide. Its market
Recent work involving coffee beans and barley seeds introduction by Uniroyal Chemical Co., Inc. followed in
indicates that such findings result from a degradation of 1963. Daminozide was used in several fruit tree species,
lysine driven by a Maillard reaction under dry thermal particularly in apple, to reduce vegetative growth and
conditions and in the presence of naturally occurring tri- improve fruit coloration and firmness (trade name: AlarÒ).
gonellin (Wermann and others 2014). Pipecolatebetaine, Treatment of peanut plants led to more compact plants
which is structurally close to mepiquat and which occurs at allowing for a more efficient harvesting (trade name:
relatively high concentrations in most vascular plants KylarÒ). However, due to toxicological concerns, its use on
(Rhodes and Hanson 1993) may also serve as a source of food crops was banned in 1989. The compound is still used
mepiquat under such processing conditions. as B-NineÒ in ornamentals, with Chemtura Corp. a major
Ancymidol (Tschabold and others 1970) and flurprimi- supplier. Trinexapac-ethyl was introduced to the scientific
dol (Barrett 1982) are closely related in structure. These community by Kerber and others (1990) and Adams and
pyrimidines were introduced by Elanco Products Co. (then others (1992). Ciba-Geigy AG (agro activities now with
subsidiary of Eli Lilly and Co., now part of Dow Agro- Syngenta) started with sales in 1992. Prohexadione-cal-
Sciences) in 1971 and 1989, respectively. SePRO Corp. cium originates from Kumiai Chemical Industry Co., Ltd.
has, meanwhile, become a major distributor. The com- and Ihara Chemical Industry Co. The first international
pounds are used to decrease the rate of growth in a wide publication on its properties as a growth retardant was by
range of mono- and dicotyledonous species, including Nakayama and others (1990). In 1991, BASF obtained a
perennial turf grasses, ornamental cover species, herba- license to develop this compound in Europe, North
ceous and woody ornamentals, and deciduous and conif- America, and several other countries. Kumiai launched
erous trees grown in gardens and parks. There are no prohexadione-calcium 1994 in Japan as an anti-lodging
123
agent in rice, while BASF started commercialization of the Thidiazuron originates from Schering AG (via AgrEvo
compound in combination with mepiquat chloride as a GmbH and Aventis CropScience S.A. since 2002 part of
stem stabilizer in cereals in 1998 and solo for use in pome Bayer CropScience) and was first reported on by Arndt and
fruit trees in 2000. others (1976). Since 1982 it is sold in the US as a cotton
defoliant. The first scientific publication on forchlorfenuron
Cytokinins has been by Takahashi and others (1978). Practical uses
started around 1980. Current major suppliers of products
Among other effects, cytokinins stimulate cell division in based on forchlorfenuron are Kyowa Hakko Kirin Co., Ltd.
meristematic tissues, counteract auxins in apical domi- and AlzChem AG.
nance, delay leaf senescence, and attract assimilates into
sinks. The adenine-type cytokinins, kinetin [N-(2-furyl- Abscisic Acid
methyl)-3H-purin-6-amine] and 6-benzyladenine (Fig. 4I)
are of practical relevance to produce transplants in tissue The plant hormone abscisic acid (ABA) plays a major role
cultures. 6-Benzyladenine is also used solo or in com- in protecting higher plants against abiotic stress. It leads to
bination with GA4/7 in pome fruits. The diphenyurea-type stomatal closure, when plants are under drought stress and
cytokinins, thidiazuron (Fig. 4II) and forchlorfenuron increases cold hardiness of plant tissues. Another function
(Fig. 4III) are applied as a cotton defoliant and to of ABA is its involvement in inhibiting precocious seed
improve fruit size in some species, respectively. Applied germination. In nature, ABA occurs as (?)-cis,trans-iso-
exogenously at appropriate dosages and at suitable stages mer (= S-ABA) (Fig. 5). Its synthesis is complicated and
of plant development, cytokinins are known to induce not suited for commercial purposes. The same is true for
leaf drop in cotton plants and abortion of young fruits in ABA analogs like those introduced by Grossmann and Jung
apple trees. Suttle (1985), Zubkova and others (1990), (1984) and Zaharia and others (2005). However, as with
and Grossmann (1991) have reported that cytokinin-in- GAs, several phytopathogenic fungi are known to produce
duced ethylene formation is responsible for abscission in S-ABA: Cercospora rosicola (Assante and others 1977),
cotton. Ethylene levels are also raised in apple fruitlets Botrytis cinerea (Marumo and others 1982), and other
after treatment with 6-benzyladenine (Greene and others species. Early attempts in the 1980s to produce ABA on an
1992). industrial scale by fermentation of C. rosicola failed at
The detection of kinetin and also of 6-benzyladenine in Sandoz Agro Ltd. and Kyowa Hakko. Likewise, Toray
the mid-1950s must be credited to Skoog, Miller, and Industries, Inc. pulled out of producing ABA with B.
Strong and their co-workers from the University of Wis- cinerea around 2000. S-ABA is now produced with high-
consin-Madison (cf. Skoog 1994; Amasino 2005; Kaminek yielding strains of B. cinerea by Sichuan Lomon Bio
2015, this issue). 6-Benzyladenine was evaluated for Technology Co., Ltd. Valent BioSciences sells products
commercial use in grapevines under the trade name ‘Ver- containing S-ABA since 2009.
dan’ by the Shell Development Co. in the late 1950s.
However, the onset of commercial success lasted until Compounds Affecting the Ethylene Status
1975, when the compound was registered by Kumiai
Chemical Industry Co., Ltd. for more attractive uses and The plant hormone ethylene (Fig. 6I) is involved in many
employing a more efficient synthesis. Abbott Laboratories plant processes such as stimulation or inhibition of cell
(PGRs now at Valent BioSciences) became a major pro- growth, induction of flowering, promotion of senescence of
vider of 6-benzyladenine-containing PGRs. tissues and organs, ripening of fruits, and formation of
Fig. 4 Structures of cytokinin-type PGRs: I 6-Benzyladenine (BA) = N-Benzyl-3H-purin-6-amine, II thidiazuron = 1-phenyl-3-(1,2,3-

thiadiazol-5-yl)urea, and III forchlorfenuron = 1-(2-chloro-4-pyridyl)-3-phenylurea
123
water, may also be used in this crop (Bartholomew 2014).

In contrast, ethephon (Fig. 6III) is a non-gaseous ‘‘pre-
drug’’ of ethylene and can be applied much more effec-
tively. Once absorbed by the plant and then present in a
slightly acidic environment, it slowly decomposes into
ethylene plus phosphoric and hydrochloric acid. Ethep-
Fig. 5 Structure of (S)-abscisic acid = S-ABA = (?)-cis,trans- hon, alone or in combination with other PGRs, has found
ABA = (2Z,4E)-5-[(1S)-1-hydroxy-2,6,6-trimethyl-4-oxocyclohex-2- many uses. It, too, is used for forcing in pineapples. In
en-1-yl]-3-methylpenta-2,4-dienoic acid
cereal production, particularly in barley, it reduces stem
elongation, thereby reducing the risk of lodging. In cot-
abscission layers leading to leaf and fruit fall. The ethylene ton, it acts as a boll opener and defoliant prior to har-
status of plants or plant products can be affected in dif- vesting. The flow of latex is intensified in rubber trees
ferent ways by different PGRs. under the influence of ethephon. It must also be noted that
auxin-type PGRs induce the formation of ethylene in
Ethylene, Acetylene, and Compounds Releasing treated plants, as part of their mode of action (Abeles and
or Inducing Ethylene others 1992).
Ethephon was first synthesized by Kabachnik and
Ethylene accelerates ripening in apples and other climac- Rossiiskaya (1946). The observation that it released
teric fruits. To keep this to a minimum, ethylene can be ethylene (Maynard and Swan 1963) was not exploited at
removed from storage rooms by different technical proce- that time. Later, ethephon was produced by the GAF
dures. Conversely, some stored fruits are treated with Corporation and tested for PGR activity by Amchem
ethylene as a signal for ripening in order to be ready for Products, Inc., who introduced it as EthrelÒ and FlorelÒ
consumption at a given time. The ethylene applied is either beginning in 1966. Reports in the scientific literature
from high-pressure gas cylinders or from generators, which describing the PGR effects followed, for example,
catalyze the conversion of ethanol into ethylene. Warner and Leopold (1967) and Cooke and Randall
Different from closed or even airtight storage rooms, (1968). After several changes in ownership, Amchem
application of gaseous ethylene to plants growing under Products, Inc. became part of Bayer CropScience, who
outdoor conditions is difficult and inefficient. The only is now a key producer and distributor of ethephon.
relevant exception for this type of use of ethylene is in Because of its use in many important crops, ethephon
pineapples for the induction of flowering and fruiting. has become a major commercial success with global
Instead of aqueous solutions of ethylene, acetylene annual sales at farm level estimated to be well above
(Fig. 6II), which is generated from calcium carbide in US$ 200 million.
Fig. 6 Structures of chemicals

affecting ethylene homeostasis:
I Ethylene, II acetylene, III
ethephon = (2-chloroethyl)
phosphonic acid, IV aviglycine
(AVG) = (2S,3E)-2-amino-4-
(2-aminoethoxy)-but-3-enoic
acid, (V) 1-methylcyclopropene
(1-MCP)
123
Inhibitors of Ethylene Biosynthesis and Blankenship at North Carolina State University at

Raleigh (Sisler and Blankenship 1996), in collaboration
It has been known since the 1970s that rhizobitoxin with Serek and Reid from the University of California in
(Owens and others 1971) and its ethoxy analog aviglycine Davis, who were the first to publish on this new compound
(Fig. 6IV) (Baker and others 1978) are inhibitors of ethy- (Serek and others 1994). 1-MCP is a relatively unstable gas
lene biosynthesis. Both compounds block the conversion of and, different for instance to ethylene, cannot be stored
S-adenosylmethionine to ACC, which is catalyzed by the under pressure in gas cylinders. To allow safe storage,
enzyme ACC synthase (Harpaz-Saad and others 2012). transport, and handling, 1-MCP is used as an inclusion
Interests in developing such compounds into practice were complex with a-cyclodextrin. The active gas is released by
initially dampened because of significant difficulties mixing the solid preparation with water. It can be applied
involved in their synthesis. However, a biotechnological very effectively in airtight storage facilities and transit
approach turned, again, out to be promising. Fermentations containers to slow down ripening and other senescence
of the soil microorganism Streptomyces sp. were found to processes of many types of fruit, vegetable, and cut
produce aviglycine (Pruess and others 1974) and could be flowers.
used for production. Switzerland-based F. Hoffmann-La Commercialization of 1-MCP was first undertaken by
Roche Ltd. at its US site in Nutley, New Jersey optimized Floralife, Inc. for use on ornamental crops. The product
the production technology. Initial steps in product devel- was approved by US authorities in 1999 and is sold under
opment were conducted at its subsidiary Dr. R. Maag AG the trade name EthylBlocÒ. AgroFresh, Inc., then a sub-
in Switzerland (Müller and others 1980). Later, Abbott sidiary of Rohm and Haas, acquired the sales rights on
Laboratories (now Valent BioSciences) went in and intro- 1-MCP. In 2002, it introduced it as SmartFreshÒ for use
duced aviglycine, primarily for delaying harvesting in on apples in the US and Chile. Uses in further fruits and
pome fruits, under the trade name ReTainÒ to the US vegetables and registrations in many other countries fol-
market in 1997. The product is now registered in many lowed. In 2009, AgroFresh, Inc. became a subsidiary of
countries, but not in EU member states. In addition to Dow Chemical Company, when Dow took over Rohm
blocking GA metabolism, trinexapac-ethyl and prohexa- and Haas. In 2015, Dow Chemical has sold a majority
dione-calcium (Fig. 3VIII, IX) are also inhibitors of ethy- stake in AgroFresh, Inc. to an investment company.
lene biosynthesis. In their free acid forms, they function as Formulations of 1-MCP, which can be applied as liquid
structural mimics of ascorbic acid, thereby displacing this sprays under field or orchard conditions, have been reg-
natural co-substrate from ACC oxidase, a dioxygenase istered in the US by AgroFresh and Syngenta Crop Pro-
converting ACC into ethylene. tection under the trade names HarvistaÒ and InvinsaÒ
since 2008. However, there has been no general intro-
Inhibitors of Ethylene Action duction to the market so far. The main uses are on maize,
soybeans, cotton, sunflower, wheat, and rice to enhance
Senescence processes may also be delayed with com- heat and drought resistance, on apples (for example,
pounds blocking the natural receptor of ethylene in its Watkins and others 2010), pears, and kiwifruit to delay
target tissues. Silver nitrate was found by Beyer (1976) to preharvest ripening and to improve fruit quality, and on
prevent the abscission of fruits, leaves, and flowers by tomatoes and peppers to improve fruit set under adverse
being antagonistic to ethylene. However, silver nitrate is weather conditions and to increase fruit size. It is no
not suitable for commercial use because of its relatively surprise that precise instructions have to be followed to
low mobility in plant tissues and its phytotoxicity. Silver, ensure successful application of a gas under outdoor
complexed as silver thiosulfate, turned out to be much conditions.
more useful. It significantly extends the vase life of car-
nations after application via their cut ends (Veen and van Compound Related to Jasmonic Acid
de Geijn 1978). Silver thiosulfate may only be used on
ornamentals and strict regulations have to be followed to Jasmonic acid is known to be involved in stress responses
keep toxicological and ecotoxicological risks to a of plants. Exogenously applied jasmonic acid inhibits
minimum. growth and induces senescence. However, its effect is
1-Methylcyclopropene (1-MCP) (Fig. 6V) offers a more relatively short-lived. Synthetic prohydrojasmon (Fig. 7) is
advanced and broader approach. 1-MCP is a gas, which more stable and is in use to promote fruit coloration in
mimics the chemical structure of ethylene and has a high apple by intensifying anthocyanin formation (Kondo
affinity to ethylene receptors in plant tissues. The discovery 2010). After Japan, Taiwan, and South Korea, a product
of 1-MCP as an ethylene antagonist resulted from a sys- containing this jasmonate is presently introduced to the US
tematic approach conducted in the laboratories of Sisler by Fine Americas, Inc.
123
Dikegulac (Fig. 8I) is readily available as a by-product

in the synthesis of L-ascorbic acid (vitamin C). It is used
for ‘‘pinching’’ on potted ornamentals, hedges, shrubs,
trees, and groundcovers, where it inhibits growth of the
terminal bud. As a result, there is less need for trimming,
and lateral branching is promoted, which gives plants a
Fig. 7 Structure of prohydrojasmon = propyl [(1R,2R)-3-oxo-2- denser and fuller look. These PGR-type effects have first
pentylcyclopentyl]acetate
been reported by Bocion and others (1975). Market
introduction under the trade name AtrinalÒ has been in
‘‘Atypical’’ PGRs: Compounds Rather Causing 1976 by Dr. Maag AG, then the agrochemical branch of
Phytotoxicity Hoffmann-La Roche. To date, PBI/Gordon Corp. is a
major distributor of dikegulac under the trade name
Several compounds are in use, which cannot be classified AtrimmecÒ. In 2007, any legal use of dikegulac ended in
as PGRs in a strict sense because they do not directly EU member states.
interfere with the plant hormonal system. In principle, they Hydrogen cyanamide (Fig. 8II) represents the market
rather act primarily by being phytotoxic. However, when standard for artificial breaking of bud dormancy. It is used
applied properly, there will be no visible symptoms of in deciduous plants (for example, apples and grapevines),
plant damage or intended damages will be restricted to which are grown in climates lacking a winter with suffi-
distinct plant parts. cient chilling for dormancy ending. Initially, calcium
Mefluidide [N-(2,4-dimethyl-5-{[(trifluoromethyl)sul- cyanamide, used for many years as a fertilizer and her-
fonyl]amino}phenyl)acetamide] and maleic hydrazide bicide, was found as a dormancy breaker in grapevines
(1,2-dihydro-3,6-pyridazinedione) act by inhibiting cell (Kuroi and others 1963). However, calcium cyanamide is
division in meristematic tissues. They are often referred to almost insoluble in water and is, therefore, unsuitable for
as ‘‘growth inhibitors’’. Mefluidide is mainly used in turf professional spray application in woody perennials.
grasses, where it reduces the need for mowing and elimi- Shulman and others (1983) identified hydrogen cyana-
nates seedhead formation. Maleic hydrazide and chlor- mide, a water-soluble hydrolyis product of calcium cya-
propham [isopropyl (3-chlorophenyl)carbamate] are namide, as the active principle. As a result of these and
applied in tobacco cultivation to keep resting buds from other investigations, SKW Trostberg AG, developed
forming shoots (‘‘suckers’’), when the flowering top part is hydrogen cyanamide and introduced it to the market as
removed. Maleic hydrazide and, in particular, chlor- DormexÒ in the mid-1980s, first in Israel, then in Brazil,
propham are also in use for the inhibition of sprouting of Chile, Taiwan, and, initially under special permits, in the
potatoes and onions in storage. Due to rising concerns US. The activities of SKW Trostberg AG are now pur-
about residues and toxicological risks of these compounds, sued by AlzChem AG. The biochemical mode of action
several alternatives have become available in recent years of hydrogen cyanamide and some other chemicals in
or are in the process of market introduction: Products based dormancy breaking involves a transient inhibition of
on carvone [mostly in its S-form (5S)-5-isopropenyl-2- respiration. This initiates a biochemical cascade—inter
methyl-2-cyclohexen-1-one], which is contained, for alia the induction of a-amylase—that leads to effective
instance, in caraway seeds and 3-decen-2-one, which is dormancy release (Rubio and others 2014). Very high
also a fragrance and is used as a food additive, are two rates of up to 25 kg/ha of hydrogen cyanamide have to be
examples. applied to dormant plants. Due to the compound’s toxi-
cological properties, farm workers are strictly advised to
avoid any contamination. In humans, the substance inhi-
bits aldehyde dehydrogenase and can produce the
acetaldehyde syndrome, when exposure coincides with
alcohol use. Furthermore, precautions have to be followed
to avoid drifting into neighboring crops. Severe leaf
burning (for example, in evergreen citrus plants) could be
the result. Because of such problems, alternatives for
hydrogen cyanamide are being sought. However, chemi-
cals such as thiourea, nitrate salts, or mineral oils in
combination with distinct surfactants can only partially
Fig. 8 Structures of ‘‘atypical’’ PGRs: I Dikegulac = 2,3:4,6-di-O-
isopropylidene-a-L-xylo-2-hexulofuranosonic acid, II hydrogen substitute for hydrogen cyanamide and often are also
cyanamide toxic.
123
Examples of PGR Uses of lodging can be differentiated: Stem lodging is mainly

observed when strong winds and intense rainfall exert a
Space limitation (and own knowledge) does not permit force that breaks the stem base. Stem lodging is often
giving a complete overview of PGR applications. Instead, found after a severe thunderstorm. Eyespot, caused by
examples of successful uses in major crops and some Pseudocercosporella herpotrichoides, and other foot rot
recent additions will be presented in the following part of diseases may intensify the risk of stem lodging. Root
this contribution. A compact listing is given in Tables 1 lodging is typically observed when, after several days of
and 2. Detailed information on how to use a given PGR in a rainfall, the plant’s roots find little anchorage in water-
distinct crop plant can be found in the respective labels, soaked soil and are unable to keep the stem with its heavy,
which are available from the distributing companies via the water-soaked ear upright. As a result of lodging, shading
internet. and the spreading of pathogens may drastically reduce
profitability through reduced yield and grain quality.
Increased costs for harvesting and grain drying will lead to
PGRs Used to Reduce the Risk of Lodging in Field further financial losses. If lodging occurs early (for exam-
Crops ple, shortly after anthesis), its impact on seed yield and
quality will be more intense as compared with lodging
Lodging is often observed in small grain cereals (including close to harvesting. Grain yield reductions of up to 40 %
wheat, barley, rye, triticale, oats), rice, grass grown for must be expected in years with high lodging incidence.
seed, oilseed rape, bush beans, linseed, and other field Under UK growing conditions, severe lodging in cereals
crops: Tall shoots with heavy ears, panicles, or other fruit may be expected in one out of 3–4 years (Baker and others
structures at their top can no longer stand upright and fall 2014). It is likely that the situation is similar in other
over, particularly when their leverage is increased by wind countries with high intensities of production. Assuming an
and rain. The risk of lodging is strongly affected by variety average yield of 7.5 tons per hectare and a producer price
and husbandry factors including sowing date, seed rate, of € 200.00 per ton for wheat, a reduction in yield of 20 %
drilling depth, and rate of nitrogen application. It occurs due to lodging is equivalent to € 300.00 per hectare.
primarily when plants are grown at high levels of intensity Additional financial losses are likely to result from inferior
as is typical not only in most European countries, but also grain quality and increased costs for harvesting and grain
in Japan, South Korea, Chile, New Zealand and, increas- drying.
ingly, in China. Area- and value-wise, the use of stem- After its market introduction in 1965, chlormequat
shortening PGRs in small grains, rice, and oilseed rape to chloride was the first PGR to be used on a large scale as an
reduce the risk of lodging is the main application of PGRs anti-lodging agent in European cereal production. Mean-
worldwide. Approximately 25 % of the global PGR sales while, additional PGRs have become available. Combina-
are represented by anti-lodging products. The usage of tions of chlormequat chloride or mepiquat chloride with
such products is common practice in the cultivation of trinexapac-ethyl or prohexadione-calcium, either as
small grains and oilseed rape in countries such as France, sequential sprays or by tank-mixing or by using ready-mix
Germany, and the UK. For instance, 89 % of the winter formulations, currently represent the best technical solu-
wheat, 76 % of the winter barley, 73 % of the oats, and tions for lodging control. When chlormequat chloride is
100 % of the winter rye acreage were treated with anti- applied in winter wheat at early stages of tillering, it
lodging products in the UK in 2012 (Garthwaite and others increases the number of fertile tillers in addition to
2013). Chlormequat chloride and trinexapac-ethyl rank reducing stem length. This can be of special interest after
third and seventh, respectively, in terms of treated area of winter losses of plants. Barley is less responsive to
all crop protectants in all field crops in this country chlormequat chloride and mepiquat chloride than wheat,
(Garthwaite and others 2013). In contrast, anti-lodging rye, triticale, or oats. Therefore, products containing
products for small grain cereals play only a minor role in trinexapac-ethyl, prohexadione-calcium, or the ethylene-
countries, where these crops generally have to be grown at releasing ethephon are preferentially used to reduce the risk
relatively low levels of intensity. In the US, Canada, or of lodging in this species. Additional information on
Australia, climatic factors, in particular unfavorable tem- lodging in small grains and on the use of anti-lodging
peratures and shortage of moisture, represent the main agents in cereal production can be found in Easson and
restrictions. Because sufficient arable land is available, others (1993), Berry and others (2004), Rademacher
significant surpluses for export can still be produced on a (2010b), Berry and Spink (2012), and Baker and others
national level. (2014).
Lodging in wheat, barley, rye, triticale, and oats occurs Lodging can also be a problem in intense rice pro-
mainly during the 2 months preceding harvest. Two forms duction (Moody 1986; Nishiyama 1986; Shibata and
123
Table 1 Active ingredients contained in ‘‘true’’ PGRs, mode of action, uses, product names, and producers or providers (listing not complete)
Active ingredient Physiological action Main uses Main product Main
name(s) producer(s) and/or
supplier(s)
Indol-3-ylbutyric acid Auxin Rooting of cuttings RhizoponÒ Rhizopon B.V.

AA
HormodinÒ OHP, Inc.
1-Naphthylacetic acid Apple, pear and other fruit trees: Obsthormon L. Gobbi s.r.l.
Increased fruit set; fruit 24a
thinning FruitoneÒ AMVAC
Chemical Corp.
2-(1-Naphthyl)- acetamide Diramid L. Gobbi s.r.l.
Amid-ThinÒ AMVAC
Chemical Corp.
Carbaryl SevinÒ NovaSource
Triclopyr TopperÒ Agriphar S.A.
Gibberellin A3 Gibberellin Pear: Parthenocarpic fruit ProGibbÒ Valent
formation BioSciences
Table grapes: Improved berry Corp.
quality Gibb 3 Globachem NV
Citrus: Improved fruit quality
Gibberellin A4/7 Apple: Reduced fruit russeting ProVideÒ Valent
BioSciences
Corp.
NovagibÒ Fine
Agrochemicals
Ltd.
Gibberellin A4/7 ? 6- Gibberellin ? Cytokinin Apple: Improved fruit quality PromalinÒ Valent
Benzyladenine BioSciences
Corp.
Chlormequat chloride Inhibitor GA biosynthesis Cereals: Anti-lodging CycocelÒ BASF
Ornamentals: Growth control
Chlormequat Inhibitor GA Cereals: Anti-lodging MeteorÒ BASF
chloride ? Imazaquin biosynthesis ? Herbicide
Mepiquat chloride ? Ethephon Inhibitor GA Barley and other cereals: Anti- TerpalÒ BASF
biosynthesis ? Ethylene releaser lodging
Mepiquat Inhibitors GA biosynthesis Cereals: Anti-lodging MedaxÒ BASF
chloride ? Prohexadione-
calcium
Mepiquat chloride Inhibitor GA biosynthesis Cotton: Growth control PixÒ BASF
Ò
Mepex Nufarm Ltd.
Mepiquat pentaborate Inhibitor GA biosynthesis PentiaÒ BASF
Mepiquat Inhibitor GA biosynthesis ? Auxin StanceÒ Bayer
chloride ? Cyclanilide transport inhibitor CropScience
Mepiquat chloride ? Kinetin Inhibitor GA MepexÒ Nufarm Ltd.
biosynthesis ? Cytokinin GinOut
Mepiquat Inhibitor GA Oilseed rape: Improved winter CaryxÒ BASF
chloride ? Metconazole biosynthesis ? Inhibitor GA hardiness; anti-lodging
biosynthesis and fungicide
Paclobutrazol Inhibitor GA biosynthesis Fruit trees: Growth control CultarÒ Syngenta Crop
Rice: Anti-lodging Smarect Ò Protection
Ornamentals: Growth control BonziÒ
123
Table 1 continued
supplier(s)
Paclobutrazol ? Difenoconazole Inhibitor GA biosynthesis Oilseed rape: ToprexÒ Syngenta Crop

? Fungicide Improved winter hardiness; Protection
anti-lodging
Uniconazole-P Inhibitor GA biosynthesis Fruit trees: Growth control SunnyÒ Sumitomo
Rice: Anti-lodging LomicaÒ Chemical Co.,
Ltd.
Ornamentals: Growth control SumagicÒ
Ancymidol Ornamentals: Growth control A-RestÒ SePRO Corp.
Flurprimidol TopflorÒ
Tebuconazole Inhibitor GA biosynthesis and Oilseed rape: FolicurÒ Bayer Crop
fungicide Improved winter hardiness; Science
Metconazole anti-lodging CarambaÒ BASF
Daminozide Inhibitor GA biosynthesis Ornamentals: Growth control B-NineÒ Agriphar S.A.
Trinexapac-ethyl Cereals: Anti-lodging ModdusÒ Syngenta Crop
Sugarcane: Harvest management PalisadeÒ Protection
Grass grown for seed: Anti- Primo MaxxÒ
lodging
Turf grasses: Growth control
Prohexadione-calcium Pome fruit trees: Growth control RegalisÒ BASF
Strawberries: Control runner ApogeeÒ
growth
Grass grown for seed: Anti-
lodging
Rice: Anti-lodging VivifulÒ Kumiai Chem.
Ind. Co., Ltd.
Ethephon Ethylene releaser Fruits: Ripening EthrelÒ Bayer
Rubber trees: Intensified latex Cerone Ò CropScience
flow
Pineapples: Flowering induction
Barley: Anti-lodging
Ethephon ? Cyclanilide Ethylene releaser ? Auxin Cotton: Defoliation; boll FinishÒ 6 Pro Bayer
transport inhibitor opening CropScience
Aviglycine Inhibitor ethylene biosynthesis Pome fruits: Delayed harvest ReTainÒ Valent
Pineapples: Control flower PinCorÒ BioSciences
induction Corp.
Silver thiosulfate Ethylene receptor blocker Carnations and other cut Chrysal AVB Chrysal
flowers: Improved vase life International
B.V.
1-Methylcyclopropene Ornamentals, fruits, vegetables: EthylBlocÒ AgroFresh Inc.
Improved storage SmartFresh Ò
6-Benzyladenine Cytokinin Pome fruits: Fruit thinning MaxCelÒ Valent

BioSciences
Corp.
Globaryll Globachem NV
Thidiazuron Cotton: Defoliation FreeFallÒ Nufarm Ltd.
Thidiazuron ? Diuron Cytokinin ? Herbicide Cotton: Defoliation GinstarÒ Bayer
CropScience
Forchlorfenuron Cytokinin Table grapes, kiwifruits: FulmetÒ Kyowa Hakko
Increased size of berries/fruits Kirin Co., Ltd.
SitofexÒ AlzChem AG
123
Table 1 continued
supplier(s)
(S)-Abscisic acid Abscisin Table grapes: Improved berry ProToneÒ Valent

color BioSciences
Prohydrojasmon Jasmonate Apples: Improved fruit color Jasumomeito Meiji Seika
EkizaiÒ Kaisha, Ltd.
BlushÒ Fine Americas,
Inc.
Table 2 Active ingredients contained in ‘‘atypical’’ PGRs, mode of action, uses, product names, and producers or providers (listing not
complete)
Active ingredient Mode of action Main uses Main product Main producer(s) and/or
name(s) supplier(s)
Hydrogen cyanamide Respiration inhibitor Fruit trees, grapevines: DormexÒ AlzChem AG

Dormancy breaking
Dikegulac Non-specific phytotoxicity Ornamentals: Pinching AtrimmecÒ PBI/Gordon Corp.
Mefluidide Inhibitor of cell division Turfgrasses: Growth control EmbarkÒ PBI/Gordon Corp.
Ò
Maleic hydrazide Tobacco: ‘‘Sucker’’ control Royal MH Agriphar S.A.
Chlorpropham Potatoes: Sprouting control Gro-StopÒ Certis Europe B.V.
3-Decen-2-one Non-specific phytotoxicity Potatoes: Sprouting control SmartBlockÒ AMVAC Chemical Corp.
Carvone TalentÒ ADAMA Ag. Sol. Ltd.
Takebayashi 2007; Salassi and others 2013). However, on long-lived uniconazole-P and paclobutrazol are the
modern semi-dwarf cultivars produce relatively high main anti-lodging agents in this crop. Prohexadione-cal-
yields while being largely lodging-resistant. In spite of cium, which is relatively short-lived and has to be spray-
this option, many farmers still prefer to grow tall, but applied, is only of minor importance.
lodging-susceptible varieties, which are tastier and Winter oilseed rape (Brassica napus ssp. napus) is an
achieve a higher price. For instance, Japanese consumers important oilseed crop in Europe and elsewhere. Severe
prefer rice from the traditional long-strawed cultivar yield losses due to lodging may occur, which can also be
‘Koshihikari’, which is grown on approximately 35 % of reduced by stem-shortening agents (Kightley 2001; Berry
the Japanese rice producing area. Without treatment, and Spink 2009; Baker and others 2014). The leading
‘‘Koshihikari’’ typically reaches a final shoot length of compounds used are the triazoles tebuconazole and met-
around 110 cm and is very prone to lodging (Shibata and conazole, which are marketed for this purpose in France,
Takebayashi 2007). As with cereal grains, several hus- Germany, UK, and several other European countries.
bandry methods can be employed to minimize lodging in Recent introductions are the combination of metconazole
rice. Reducing overgrowth of stem and leaves by applying with mepiquat chloride and paclobutrazol with the fungi-
PGRs at mid-tillering stage reduces lodging incidence and cide difenoconazole. Tebuconazole and metconazole are
gives a generally improved plant stature (Yim and others primarily used as fungicides in a number of crop plants
1997). It can be assumed that more than 20 % of the including oilseed rape. Their shoot growth-reducing
Japanese rice-growing area is being treated with anti- activity is restricted to oilseed rape and only few other
lodging preparations. Whereas stem stabilizers are applied species. The mentioned PGRs are also used in winter oil-
to cereal plants by spraying the foliage, granules for seed rape to keep plants from too intense growth in late
throwing into the paddy field are preferred in rice pro- autumn, thereby making the crop less vulnerable to freez-
duction, particularly in Japan and South Korea. In order ing and desiccation in winter. This is of particular interest
to be absorbed via the roots, such stem stabilizers have to in continental Europe, where harsh winters are very
be relatively persistent. Accordingly, preparations based common.
123
Trinexapac-ethyl Used in Opium Poppy different mepiquat-containing products give comparable

results.
Lodging can also constitute a problem in poppy cultivation. Treatment of cotton plants with mepiquat is especially
Trials in Tasmania, Australia, where the major portion of important when lush shoot growth is expected due to good
the global legal opium poppy production is located, were, availability of water. Application starts at the beginning of
therefore, conducted with trinexapac-ethyl and prohexa- flowering and reduces the intensity of new growth and,
dione-calcium primarily to overcome this problem. Sur- thereby, improves the sink strength of the first six to ten
prisingly, it was found that treatments with these fruiting branches. This is of great importance because the
compounds did not only improve lodging resistance but bolls retained in this part of the plant will give the highest
also changed the alkaloid spectrum in the harvested plant and earliest yields. The shifting of assimilates into the older
material in a desirable way: More thebain, which is of fruiting structures is at the expense of younger fruits, which
higher value, is formed at the expense of lower-valued the plant is continuously forming even late in the season.
oripavine (Cotterill 2005). It has been postulated that a The younger fruits are unlikely to contribute to yield when
2-oxoglutarate-dependent dioxygenase, which catalyzes plants are mechanically harvested—mostly only once per
the conversion of thebain into oripavine is blocked (Hagel season. Plants treated with mepiquat produce higher har-
and Facchini 2010; Dean 2011). Trinexapac-ethyl is now vestable yields and can, typically, be picked three to
registered in Australia for use in opium poppy. 10 days earlier than untreated plants. Earliness is of great
importance because harvesting can often be performed
PGRs Used in Cotton Cultivation prior to periods of rainfall. This would also reduce the
incidence of fungal diseases. Further benefits from short-
Cotton is a perennial plant, which, however, is cultivated in season production may result from savings in late-season
most countries in an annual cycle. In its native habitat, a irrigation and control of insect pests. Finally, decreases in
cotton plant does not die in the fall but continue to grow quantity and quality of the lint due to weathering are
until environmental conditions, in particular frost, become reduced in the oldest, first-opened cotton bolls. Detailed
too restrictive. Another growth characteristic associated background information on using mepiquat in cotton have
with its perennial nature is its indeterminate fruiting habit. been presented by Kerby and others (1986) and Gwathmey
Rather than flowering during a distinct period following and Clement (2010).
vegetative growth, cotton plants simultaneously produce Prior to harvesting, cotton plants require treatments to
vegetative and fruiting structures. Cotton fibers form mark the end of the season. Defoliation, opening of almost
around the developing seeds inside a protective capsule mature bolls, desiccation, and prevention of re-growth is
called a boll. When the seeds are mature, the boll ruptures widely practiced in modern cotton production. Defoliants
and opens, the fibers dry and unfurl and can be harvested. can be categorized as having either herbicidal or ‘‘true’’
In order to enable high yield and fiber quality and to allow PGR-type activity. Herbicides such as carfentrazone-ethyl,
efficient mechanical harvesting, an intense usage of PGRs fluthiazet-methyl, saflufenacil, or tribufos injure the plant,
and other agrochemicals has become a standard practice in thereby causing it to produce ethylene. The ethylene then
many cotton-producing countries. An actual survey on promotes abscission and leaf drop. If these herbicides are
practical aspects of cotton production, including the use of applied inadequately, they may kill the leaves too rapidly
PGRs, has been published by Edmisten and others (2014). before ethylene can be produced. This results in rapid
The growth retardant mepiquat chloride was commer- desiccation, and the dry leaves will not drop but rather
cially introduced for vegetative growth control in cotton in remain at their sites, where they will cause difficulties with
1980 in the US and has, since then, become a cornerstone mechanical harvesting. PGR-type products, containing
of modern cotton production (Kerby and others 1996). ethephon, ethephon plus cyclanilide, thidiazuron, or thidi-
Instead of or in addition to mepiquat chloride, chlormequat azuron plus the herbicide diuron are less likely to create
chloride is used for the same purpose in Australia and some such difficulties. Ethephon releases ethylene and thidi-
other countries. Mepiquat chloride is also available in azuron promotes ethylene formation. Abscission zones are
combination with cyclanilide or kinetin. Another variant is then formed in the leaf petioles leading to defoliation. A
mepiquat pentaborate, which was introduced in 2003 in the few days after treatment with PGR-type defoliants, leaves
US (Stapleton and Via 2003). The pentaborate form of drop green and fresh. Hence, the risk of contaminating
mepiquat is absorbed more rapidly and provides boron, cotton fibers with trash and stain during mechanical harvest
which, especially under boron deficiency, can be of rele- is significantly reduced. Ethephon is also used to open
vant nutritive value (Rademacher unpublished) and may, almost mature cotton bolls, thereby increasing har-
therefore, lead to better plant performance (Norton and vestable yield. None of these procedures will kill the cotton
Borrego 2006). However, in the majority of cases, the plant, and fresh re-growth may impede harvesting.
123
Therefore, treatments for defoliation and boll opening are macadamia nuts are just the main examples. It is much
often combined with a desiccant such as paraquat or more difficult to apply PGRs successfully in such woody
sodium chlorate. Another aspect to observe is the fact that perennials than in arable crops. Mistakes made in one year
the herbicide-type defoliants and desiccants work satis- often lead to problems in the years following. On the other
factorily over a wide range of temperatures and also below hand and in contrast to field crops, fruits and nuts, typi-
18 °C. However, several of these compounds are consid- cally, represent high-value crops and, hence, allow for the
ered toxicologically and environmentally questionable. In use of more elaborate and expensive products. Recent
contrast, ethephon, thidiazuron, and cyclanilide do not have overviews on different PGR uses in fruit production have
such risks, but work effectively only when applied at been presented by Petracek and others (2003), Greene
temperatures above approximately 18 °C. To obtain the (2010), and Looney and Jackson (2011).
best results possible at given situations, combinations of
different products are general practice. More details are Breaking of Bud Dormancy
given in Edmisten and others (2014).
Many deciduous plants, including apples, pears, peaches,
Prohexadione-Calcium Used in Peanuts and cherries, are genetically adapted to areas with pro-
nounced seasonal cycles. Growing in such temperate cli-
The foliage of peanut plants is still green at harvesting such mate, these plants lose their leaves in fall and start with
that excessive vine growth may reduce digging efficiency. new shoot growth and flowering after winter. Mild tem-
Prohexadione-calcium, which is registered for this use in peratures, which may occur in late autumn or early winter,
the US, retards vegetative growth and improves the visi- do not encourage shoot growth, since the plants are dor-
bility of rows, resulting in improved harvesting efficiency. mant at that time. Endodormancy is lost only when the
Pod yield and kernel quality may also be improved (Jordan plants had been exposed for sufficient time to sufficiently
and others 2001, 2008). low temperatures. This mechanism guarantees that delicate
plant parts such as leaves and flowers are not damaged by
PGRs Used in Sugarcane exposure to winter conditions. However, sufficient winter
chill is often lacking or insufficient, when temperate
Ethephon and glyphosate or other herbicides are often used perennial species are grown in subtropical or even tropical
as chemical ripeners in sugarcane production. They have to climates. Shoot growth and flowering will be very scattered
be applied via aircraft or special ground-operated booms. when a new growing season begins. Uniformity is, how-
By rapidly reducing the sink demand of young and growing ever, a major prerequisite for proper production and com-
plant parts, sucrose storage within the stalk is accelerated mercial success. The leading PGR to substitute for chilling
leading to higher harvest yields. Even better results can be is hydrogen cyanamide. Other products used for dormancy
achieved with trinexapac-ethyl (Resende and others 2000; breaking are based on thiourea, nitrate salts, or mineral
Rixon and others 2007). Trinexapac-ethyl became recently oils.
registered in Brazil, Australia, the US, and other countries
for use in this crop. Control of Vegetative Growth
Ethephon Used on Rubber Trees Proper tree growth management is of major concern in the
commercial production of tree fruits and nuts. In pome and
Latex from the rubber tree, Hevea brasiliensis, is produced stone fruits, for instance, avoiding excessive shoot growth
in laticifers, specialized cells located within the bark of the will induce flowering and fruiting in younger trees. Older
tree. On incising the tree’s bark (‘‘tapping’’), the latex trees have to be restricted to their allocated space, thereby
oozes out and is collected. Ethephon, which delays plug- reducing crowding and shading. The crowns of fruit trees
ging of the latex vessels, is widely used as a yield stimulant should be sufficiently opened to allow good light penetra-
(Abraham 1992; Lacote and others 2010). tion to the inner parts of the canopy, thereby improving
photoproductivity and fruit coloration. Additionally, effi-
PGRs Used on Fruit and Nut Trees and in Kiwifruit cient crop protection is significantly facilitated in such
Production trees. Since the beginning of professional tree fruit pro-
duction, several techniques have been employed to avoid or
A large number of different PGRs are in use for treating to control excessive shoot growth. Different types of
different fruit and nut trees and kiwifruits grown in dif- dwarfing rootstocks and scions have become available,
ferent climates. Apples, pears, cherries, peaches, kiwifruits, particularly for apples. Different cultivars may also show
citrus fruits, avocados, mangos, litchis, walnuts, pecan, and significant differences in shoot vigor. Shoot pruning
123
represents the main cultural practice for vegetative growth 3-deoxyflavonoids, such as luteoforol and luteoliflavan.
control. Other methods include root pruning, root restric- Luteoforol shows biocidal activity against several bacterial
tion, stem girdling or stem sawing, limb bending, breaking and fungal pathogens, including Erwinia amylovora and
or wounding, and restrictive fertilization and irrigation. Venturia inaequalis, the causal agents of fire blight and
However, each of these methods is cost-intensive and/or apple scab, respectively. The triggering of such phy-
bears a high risk of failure. Furthermore, part of the trees’ toalexin-like compounds explains why treated pome fruit
assimilates or potential assimilates are lost. trees are significantly less affected by a number of diseases.
Chemical regulation of tree shoot growth has been This induction of defense is of particular value to control
practiced over many years by using distinct inhibitors of shoot fire blight, a disease caused by the bacterium E.
GA biosynthesis (Greene 2010). Reduction of shoot length amylovora, which is primarily controlled by using antibi-
facilitates tree management (for example, pruning, appli- otics, a treatment that is highly controversial. (d) Apple and
cation of crop protectants, fruit picking) and more trees pear trees treated with prohexadione-calcium are also less
may be grown per unit area. Increases in fruit yield and affected by insect pests, such as aphids, psyllids, and
quality are often observed. Paclobutrazol and uniconazole- leafhoppers, which assists the action of insecticides. The
P are in use in some countries for growth control in pome underlying biochemical mode of action could also be
and stone fruits. Application is via spraying or as a soil changes in the spectrum of flavonoids, which may, for
drench. The active ingredients are highly persistent, both in instance, repel sucking and chewing insects. However, a
the soil and in treated plants, which makes precise dosag- thicker epidermis or thicker cell walls, resulting from
ing difficult and which raises concerns about residues. reduced elongation growth may also be the reason for less
Consequently, these compounds are not registered for use insect attack.
in fruit trees in the US, Canada, and most EU countries. In
several countries with warmer climates (for example, Control of Fruit Set
Australia, South Africa, India, Mexico, Chile), paclobu-
trazol and uniconazole-P are used to control excessive Most fruit species have a self-regulating mechanism,
shoot growth in avocado, mango, litchi, pecan, and maca- which results in the shedding of flowers or young fruits
damia. Here, the longevity of the compounds can be seen immediately after flowering or in the course of June drop.
as an advantage because less persistent compounds are However, the resulting fruit number is rarely optimal.
likely to be degraded too rapidly under the given climatic Achieving the right fruit set is crucial for obtaining a
conditions. Background information on shoot growth reg- good harvest, both quantitatively and qualitatively, with-
ulation in these subtropical fruit trees is given by Yeshitela out too many or too few flower buds being induced for
and others (2004), Menzel and Le Lagadec (2014), and the next season. Alternating trees producing too many
Pires and Yamanishi (2014). fruits of inferior quality in an ‘‘on’’ year, but too few and
A new option for controlling growth of fruit trees often too big fruits in an ‘‘off’’ year need to be brought to
became available with the introduction of prohexadione- a more consistent fruit production. Hence, controlling
calcium in the year 2000. This compound has a half-life in fruit set is an important cultural practice, particularly in
plants of approximately 10–14 days and of \1 day in apples, but also in other deciduous fruit trees (Dennis
microbially active soil. In fruit production, it is mainly used 2000; Wertheim 2000; Webster and Spencer 2000;
in apples, but also in pears and sweet cherries. Due to its Greene 2010; Greene and Costa 2013) and in citrus
simultaneous effects on the formation of GAs, ethylene, (Coggins and Lovatt 2014). Mechanical or chemical
and flavonoids, treated pome fruit trees benefit in several thinning with PGRs of flowers and young fruits is
ways (Rademacher and others 1999, 2004, 2006). (a) A required when there is a risk of having too high a crop
lowered production of growth-active GAs leads to less load. Conversely, the number of fruits growing on a tree
shoot growth. The labor needed for dormant and summer may become a limiting factor, when alternating trees are
pruning in treated trees is typically reduced by 30 % and in the ‘‘off’’ year or when trees have lost many flowers as
more. Furthermore, a more open canopy allows better light a result of late frost. Here, one would like to avoid any
penetration into the central parts of the crown leading to drop of intact flowers or young fruits. Within limits, this
improved fruit quality. Spray application of crop protec- can also be achieved with PGRs.
tants and foliar nutrients is also made more efficient. Hand thinning of young pome fruits shortly after June
(b) The inhibition of ethylene formation can be employed drop is very labor-intensive, but it is the only reliable
to increase fruit set. Due to the fact that there is less shoot method for obtaining optimal results. Since the 1960s,
growth, additional amounts of assimilates are available for several PGRs have been introduced for flower and fruit
fruit formation. (c) Blocking flavanone 3-hydroxylase thinning. They are all working by either releasing ethy-
activity in pome fruit shoots leads to a de novo formation of lene or by inducing the formation of this hormone.
123
Ethylene will then cause the formation of an abscission Improvement of Fruit Quality
layer and, finally, flower and/or fruit drop. None of the
PGRs available for thinning is easy to use. Especially A mixture of GA4 and GA7 is often used by apple growers
weather conditions may have a significant impact on a to reduce fruit russetting. This is a superficial disorder in
thinning agent’s performance. The worst result that can which the fruit surface is interrupted by raised corky out-
happen is severe over-thinning leading to substantial growths. The GA4 component is the primary active ingre-
financial losses. Against this background, carbaryl with dient for the control of russetting. In contrast, longer-
‘‘soft’’ auxin-like activity can be regarded as the most lasting GA7 is rather unwanted because it may reduce the
reliable thinning agent. However, carbaryl acts also as an induction of flower buds and, hence, lead to poor return
insecticide. It may eliminate bees and other beneficial bloom. Because of this, preparations low in GA7 often have
insects, particularly when blooming plants are present in an advantage (Carlson and Crovetti 1990). Combined with
the orchard’s ground cover. Not least as a result of this the cytokinin 6-benzyladenine, GA4/7 is also employed to
risk, carbaryl has been banned in all EU member states improve size and shape of apples fruits. Fruit elongation
and several other countries. Other compounds used for and development of more prominent calyx lobes, particu-
chemical thinning are the auxins 1-naphthylacetic acid larly in ‘Red Delicious’ apples, are of special interest in
and 2-(1-naphthyl)acetamide, the cytokinin 6-benzy- North America. Peel color is a highly important quality
ladenine, and the ethylene releaser ethephon. The latter is trait determining market acceptance. Coloration of red
primarily used for flower thinning. Thinning machines, apples is a complex process, which, inter alia, depends on
which ‘‘thresh’’ flowers and very young fruits from trees, low night temperatures at the end of the season. If this
have recently been introduced. This type of thinning can induction is missing or not sufficient, color development
only be applied at flowering or shortly thereafter and, can also be stimulated by treatment with ethephon. How-
therefore, bears a risk of over-thinning. Furthermore, trees ever, this treatment will also accelerate fruit ripening and
have to be specifically trimmed and mechanical wounding is, therefore, likely to reduce storage life. Prohydrojasmon,
may lead to infection with fire blight or other diseases. which does not affect ripening, could be a better alterna-
Caustic ammonium thiosulfate has also found some use tive. In sweet cherries, GA3 is used to produce brighter-
for flower thinning. Even the herbicide metamitron, a colored, firmer fruits with increased size. Forchlorfenuron
photosynthesis inhibitor, is currently introduced to cause is often used to improve the size of kiwifruits.
fruit drop in apples and pears. It is evident that improved GA3 is also of major interest to improve fruit quality in
PGR thinners or alternative methods are still needed for a most citrus species and varieties (El-Otmani and others
more reliable and acceptable approach. 2000; Coggins and Lovatt 2014). In Navel oranges, rind
Increasing fruit set, when the number of intact flowers aging may be delayed. Fruits from treated trees will also
represents a limiting factor, can also be achieved with display less rind disorders (for example, rind staining,
PGRs. Low dosages of synthetic auxins, such as 1-naph- water spotting, puffy rind, sticky rind). ‘‘Creasing’’ or
thylacetic acid and 2-(1-naphthyl)acetamide are applied for ‘‘puff and crease’’ is an important rind disorder, which is of
this purpose in apples. Even better and more consistent particular concern in Navel and Valencia oranges and in
results can be achieved with prohexadione-calcium. Its Satsuma mandarins. It occurs when the rind tissue (orange-
inhibition of ethylene formation reduces the intensity of colored flavedo layer plus epidermis) continues to stretch,
flower and fruitlet abortion. Simultaneously, there is less while the albedo layer (white tissue under the rind) has
vegetative growth and more assimilates are available for already stopped growing. As a result, some parts of the
fruit production. Plums and cherries often drop at relatively fruit surface appear inflated (‘‘puffy’’), whereas other areas
late stages of fruit growth. In such cases, prohexadione- are indented (‘‘creased’’). Puffiness occurs also in grape-
calcium has a positive effect on fruit retention (Rade- fruits. Application of GA3, together with certain cultural
macher unpublished). Fruit retention in mango can be practices, is employed to reduce the incidence of this
improved by 1-naphthylacetic acid (Hagemann and others disorder.
2014). In pears, parthenocarpic fruit formation can be
achieved with GA3. This leads to an increased fruit set and Acceleration or Delay of Fruit Ripening for Optimal
is particularly important when the generative part of the Harvesting
flower has been damaged by frost or when there has been
poor pollination. Some varieties, for example, ‘Williams’ Advancing or delaying fruit ripening may bear several
(= ‘Bartlett’) and ‘Abate Fetel’, are more responsive than advantages to growers. Harvesting can be facilitated if
others. GA3 can also be used to increase fruit set and yield fruits are more mature. Harvesting activities can also be
in Navel and Valencia oranges as well as in clementines, aligned with the availability of labor resources, while
tangelos, and tangerines. ensuring at the same time that fruits are picked at their
123
maximum level of quality. Another advantage is to harvest accelerate maturity, and promote fruit abscission as an aid
and sell fruits, when good market prices are expected. for mechanical harvesting. The latter is also of practical
Unusually high temperatures prior to harvest may cause relevance in macadamia nuts.
rapid fruit ripening and pose the risk of early fruit drop in
apples. Storage of fruits advanced in maturation may also Optimization of Fruit Storage and Ripening
be impeded. Hence, if picking crews are unable to harvest
fruits on time, severe economic losses may result. In such Ripening processes in climacteric fruits are largely regu-
situations, PGRs with a stop-drop effect are of high inter- lated by ethylene. Typically, ethylene is produced at the
est. 1-Naphthylacetic acid is used since many years for this beginning of ripening. Once induced, ethylene triggers the
purpose. It inhibits fruit abscission but the fruit continues to formation of additional amounts of this hormone, thereby
mature at an even accelerated rate. As a result, apples accelerating the ripening process. In order to keep this to
treated with this PGR against preharvest drop will not keep a minimum and to extend storage life, storage at low
well in long-term storage and should be sold soon after temperature and under controlled atmosphere (relatively
picking. In contrast, aviglycine very effectively delays fruit low levels of oxygen plus relatively high levels of CO2) is
maturation and the risk of fruit drop, thereby allowing often practiced. In addition to this, ethylene can be
regular storage after harvesting. Expanded time windows removed (‘‘scrubbed’’) in storage rooms by chemical (for
for handling and marketing of fruits are additional positive example, with potassium permanganate) or photocatalytic
effects. The use of aviglycine is of particular interest in the oxidation. With the introduction of 1-MCP, fruit storage
US and Canada, where large apple farms cultivating only is further improved at a reduced energy demand for
few varieties are relatively common. By treating trees some cooling. 1-MCP can be used very effectively in airtight
three to four weeks prior to regular harvest ripeness, the storage rooms, containers, or bags to stop ripening and/or
resulting ripening delay of one week or more allows for an maintain freshness of a number of different fruits,
improved harvesting schedule (Schupp and Greene 2004). including apples, pears, plums, kiwifruit, bananas, per-
Attention has to be paid to using aviglycine at the right simmons, avocados, and mangos. 1-MCP has to be
timing and dosaging and to appropriate storage condition applied by trained personnel because different fruit spe-
following harvest. Otherwise, inadequate maturation pro- cies and even varieties need specific treatment strategies
cesses may lead to fruits of inferior quality, due, for to guarantee optimal quality, when the fruits are sold for
instance, to lack of aroma formation (Fan and others 1998). consumption.
The need of aviglycine is much less pronounced on smaller Conversely to delaying maturation, some fruits require a
farms or when several apple cultivars are grown, which signal for ripening in order to be ready for consumption at a
need to be harvested in sequence over a longer period of given time. The most prominent example for this is the
time. Aviglycine is not registered in EU member states, ripening of bananas. In order to gain maximum life, banana
where such types of fruit farms are more common than, for bunches are harvested green and then shipped under
instance, in North America. Spray-applicable 1-MCP, sophisticated cooling at 13 °C. On arrival at the destina-
which is likely to act faster than aviglycine, is currently tion, the bananas are held at approximately 17 °C and
introduced to the market as an additional stop-drop agent treated with a low concentration of ethylene. After a few
(Greene 2010). days, the fruits have begun to ripen and are distributed for
GA3 is used in sweet and sour cherries to delay fruit retail sale.
ripening by several days and thus extend the picking per-
iod. Wet weather conditions imposing a risk of rain cracks PGRs Used in Grapevine Production
might be avoided. This treatment also increases fruit size,
firmness, and resistance to postharvest disorders. GA3 can In areas lacking sufficient winter chill, it may be necessary
also be employed in most citrus species and varieties to to break the vine’s dormancy. Hydrogen cyanamide is the
delay harvesting (El-Otmani and others 2000; Coggins and most commonly used active ingredient for this purpose.
Lovatt 2014). Growers can spray part of their groves to Control of vegetative growth may also be of interest in the
allow sequential harvesting after picking fruits from non- production of table and wine grapes. Excessive cane
treated blocks early. A delayed harvest may also be used to growth is particularly observed in warmer climates, when
‘‘store’’ citrus fruits (which are non-climacteric) on the tree there is ample supply of water. As a result, shading and
until a better market window opens. Conversely, ethephon insufficient air circulation often lead to reductions in yield
can be used in sour cherry orchards or coffee plantations to and quality. Additionally, berry production will severely
promote maturity and its uniformity, fruit loosening and, as suffer from the shoot’s more intense competition for
a result of both, optimize mechanical harvest. Ethephon assimilates. Mepiquat chloride is in use for vegetative
may also be used in walnuts to speed up hull dehiscence, growth control, for instance, in Spain, Japan, and South
123
Korea (Lim and others 2004). Chlormequat chloride is used poor induction of shoot and flower buds for the next
in India. season and, therefore, are not suitable for GA3 treatment
When GA3 had become available to researchers in the (Petgen 2005; Molitor and others 2012). Recently, an
late 1950s, the first major practical uses of this new PGR alternative strategy was developed through the application
were found in table and wine grapes (Weaver 1958; of prohexadione-calcium. Mid flower opening, when the
Weaver and others 1962). As far as can be judged, the main caps on 20–80 % of the flowers have abscised, is rec-
use of GA3 on a global scale is still in grapevines, partic- ommended as an ideal timing for prohexadione-calcium
ularly in seedless table grapes. Table grapes without seeds application, which overlaps with the time window for
are attractive to consumers. However, their small fruit size GA3. At first glance, it may appear paradoxical that an
represents a problem for commercialization. GA3, applied inhibitor of GA biosynthesis is giving effects equivalent
at the right time and dosage for a given variety, is used to to those of an active GA. So far, attempts to find out the
initially create space for large berries and, thereafter, underlying mechanisms remained inconclusive (Böll and
increase the size of the remaining berries by substituting others 2009). However, it appears likely that prohexa-
for GAs, which are normally produced by the seeds. In dione-calcium blocks the metabolic inactivation of active
general, treatment with GA3 at approximately 20 mm of GAs present at the time of treatment. Increased levels of
cluster length is used to ‘‘stretch’’ the rachis, application at GA1 or other active GAs are likely to be the cause of
30–80 % cap (calyptra) fall reduces berry set and later berry thinning. Different from applying relatively persis-
treatments increase the size of the remaining berries. Under tent GA3, this effect will be comparatively transitory and
advanced production conditions (for example, in Califor- will not cause negative effects for the following season.
nia, Italy, or Chile), seedless varieties are grown on more This explanation corresponds well with the detailed ana-
than 80 % of the area and are virtually all treated with lytical data on the presence of different GAs in devel-
GA3. For additional reading, see Dokoozlian and Peacock oping grape berries (Giacomelli and others 2013). GA1
(2001) and Casanova and others (2009). Ethephon is a and GA4, which are both biologically active, give a clear
further important tool in the production of table grapes. It peak at anthesis and decline sharply thereafter. Prohexa-
advances maturation and, in red-pigmented cultivars, also dione-calcium is likely to inhibit the hydroxylation of
fruit coloration. Increase in profitability results not only these GAs via GA 2-oxidase into inactive GA8 and GA34,
from earliness and better color, but also from a reduced respectively. A GA-like effect resulting from treatment
number of picks necessary to complete harvest. Intensified with prohexadione-calcium has been reported for
pigmentation in red table grape varieties can also be Matthiola incana. The authors suggest that inactivation of
achieved with S-ABA, which may be combined with existing GAs by GA 2-oxidase is blocked by prohexa-
ethephon. Another PGR used to increase berry size in dione-calcium as the underlying mechanism (Hisamatsu
table grapes is forchlorfenuron (Peppi and Fidelibus 2008). and others 1998).
Several cultivars of (seeded) wine grapes tend to form
very dense clusters. At veraison, the pressure exerted by PGRs Used in Berry Production
the ripening and expanding adjacent berries causes
wounding and leakage of juice, which may lead to bunch GA3 is often used in professional blueberry production to
rot (caused by B. cinerea) and also sour rot (caused by increase fruit set. Foliar sprays of ethephon a few days
different bacteria and yeasts). This is of particular concern prior to expected harvesting concentrate maturity of
to vintners, when rainfall keeps the clusters moist, thereby blackberries and blueberries for easier and more efficient
facilitating the spreading of diseases. As in seedless mechanical harvesting. Recently, prohexadione-calcium
table grapes, GA3 is of interest to ‘‘stretch’’ the rachis and has been registered in the US and Canada for use in
reduce berry set in order to increase berry separation. The strawberries. In areas with pronounced annual shifts in day
window for application lies between a cluster length of length, short-day cultivars are mainly grown. In these
approximately 7.5 cm and the end of flower opening. cultivars, day length has a significant impact on the dif-
Earlier treatments lead primarily to a stretching of the ferentiation of the axillary buds. Runners develop under
inflorescence, whereas later timings cause berry thinning long-day conditions, whereas branch crowns are formed in
or the formation of non-seeded ‘‘shot’’ berries. Berry short days. Flowers for the coming season develop from
abortion results from excessively high GA levels, a branch crowns. Prohexadione-calcium controls runner
combination of endogenous hormones produced by the formation, which favors the number of branch crowns. As a
developing seeds and that applied externally. Many vari- result, significant increases in berry yield are achieved
eties, for instance of the Pinot family and Chardonnay, (Hytönen and others 2008; Handley and others 2009).
respond relatively well to treatment with GA3. However, Prohexadione-calcium can also be used to produce high-
Riesling, Sauvignon Blanc and other varieties suffer from quality strawberry transplants (Reekie and others 2005).
123
PGRs Used in the Production of Vegetables and fruit induction. For more details on flowering regula-
tion in pineapples, see Van de Poel and others (2009) and
Growth retardants are used in several countries to keep Bartholomew (2014). GA3 may be used prior to harvesting
vegetable transplants from overgrowing. In the US, uni- to increase fruit size.
conazole-P is registered for such use in tomatoes, egg-
plants, and other species. Mepiquat chloride (for example, PGRs Used on Ornamental Plants
in Spain and Turkey) and chlormequat chloride (for
example, in India) are applied in onions and garlic to Propagation of orchids and many other ornamental species
improve productivity and bulb quality. Fruit yield in field- by tissue culture is nowadays a standard technique for
grown tomatoes and other solanaceous species can be producing large numbers of cloned plants. Auxins and
increased with synthetic auxins. This is of particular cytokinins are indispensable components of the media
interest when fertilization and proper fruit set is impeded used. 2,4-D is widely used for callus induction, whereas
by too low or too high temperatures. Applied at the end of indol-3-ylacetic acid, indol-3-ylbutyric acid, and 1-naph-
the season, ethephon accelerates and concentrates ripening thylacetic acid are used for root formation. Cytokinins such
in field-grown tomatoes, thereby increasing recoverable as kinetin, 6-benzyladenine, and thidiazuron promote cell
fruit yield from a once-over harvest. GA3 may be used on division, shoot proliferation, and shoot morphogenesis
artichokes to advance maturation for earlier harvesting. (Smith 2013). Auxins, in particular indol-3-ylbutyric acid,
are used to root cuttings. GA3 has found some use in the
PGRs Used in Pineapple Production production of herbaceous ornamentals, when longer stems
or peduncles are desirable. Increased flowering is also
The ‘‘production cycle’’ of pineapple plants requires on induced by GA3 in certain species. However, reducing
average 18 to 22 months from planting to harvesting. In shoot elongation and promoting lateral branching and
modern large farms, the crop is planted and harvested flowering in ornamental and bedding plants is of much
weekly year-round. This would be impossible without the greater relevance. Compact and dense plants require less
use of PGRs. Using appropriate PGRs for floral induction space in a greenhouse, they need less water for irrigation,
and fruiting is known for a long time. Such harmonizing they have an increased shelf life, but, above all, they sell
and timing of fruit development and harvesting offer a better because of their dark green leaves, which is gener-
number of benefits to the planter such as (a) rational use of ally associated with better quality. A wide assortment of
land and more efficient crop management, (b) fructification growth retardants is currently available to ornamental
uniformity and harvest concentration leading to cost growers in the US. These products are based on chlorme-
reduction, and (c) regular and constant supply of fruits for quat chloride, daminozide, ancymidol, flurprimidol,
the fresh fruit market and for the canning and juice paclobutrazol, and uniconazole-P (Whipker and Latimer
industry. Currently, pineapple plants on large plantations 2013). Additionally, dikegulac may be used to increase
typically are forced to flower with ethephon or aqueous branching. Detailed use recommendations for a large
solutions containing ethylene. On smaller farms, applica- number of ornamental species and bedding plants raised
tion of acetylene dissolved in water is more common. under greenhouse conditions are given by Whipker (2013).
Aqueous solutions of the gases, often containing activated The spectrum of active ingredients available in EU member
charcoal for a delayed gas release, are applied over the countries consists primarily of chlormequat chloride,
whole plant or into the plant’s ‘‘cup’’, which is formed by paclobutrazol, and daminozide. In Germany, metconazole
the terminal whorl of leaves. Ethephon is preferentially and prohexadione-calcium are additionally allowed for use
applied over the entire plant. in ornamentals. The latter compound should not be used in
Pineapples are commonly planted within approximately plants with red or blue flowers because of its interference
30 °N to 30 °S latitude. Particularly in areas more distant with anthocyanin formation (Rademacher 2000). 1-MCP is
from the equator, natural induction of flowering in all or used in cut flowers, potted flowers, and bedding, nursery
parts of the plants, which results from lower temperature, and foliage plants to prevent premature wilting, leaf yel-
shortened photoperiod, reduced intensity of solar radiation, lowing, premature flower opening, and premature death
or other environmental factors, is often experienced and when plants are in storage or transport (Serek and others
impedes efficient year-round production as described 1994). Additionally, vase life of treated cut flowers is often
above. Evidence is available that natural induction of improved. The same is achieved with silver thiosulfate,
flowering involves the formation of ethylene. Aviglycine is which is especially applied to cut carnations.
nowadays used in many pineapple plantations to minimize Inhibitors of GA biosynthesis, in particular flurprimidol
this unwanted process, thereby upgrading the following use and paclobutrazol, are also being used in the US and
of ethephon, ethylene, or acetylene for an optimized flower some other countries to reduce the growth of woody
123
ornamentals in gardens or parks. Paclobutrazol and flur- PGRs Used in Hybrid Seed Production
primidol often serve as tools to arborists to limit the size
and growth of trees and shrubs in power line and utility Hybrid cultivars have become common in maize, sugar
rights-of-ways corridors. Tree growth regulation is regu- beet, and several other crop plants. However, no cost-ef-
larly applied in high visibility locations such as parks, ficient hybrid seed production systems exist so far for
historic downtowns, residential areas, and other places, several important species. A prerequisite for hybrid
where trees have a cultural value and where pruning and breeding is a tight pollination control, which avoids self-
trimming is difficult to conduct or unwanted (Chaney fertilization and provides viable pollen from the ‘‘male’’
2005). While paclobutrazol is applied via soil injection or plant at the right time and at the right place to fertilize the
soil drenching, flurprimidol is typically administered via ‘‘female’’ plant. GA3 is used by breeders in several plant
stem injection. ‘‘Nuisance fruit’’ produced by ornamental species to coordinate the development of the ‘‘male’’ (male
trees or shrubs can be a concern for homeowners, land- fertile) and the ‘‘female’’ (male sterile) plant for crossing.
scapers, and park and city officials. The fruits and seeds For instance in rice, GA3 increases the emergence of the
of some trees and shrubs, such as buckthorn, mulberry, ‘‘female’’ panicles from the leaf sheath, thereby improving
persimmon, sweetgum, and ginkgo are unsightly, smelly, the ability to accept pollen from the ‘‘male’’ plant (Ja-
may stain cars and walkways, or can be dangerous gadeeswari and others 2004; Gavino and others 2008). S-
because they create a falling hazard. Ethephon and ABA may be used to facilitate maize hybrid seed produc-
1-naphthylacetic acid are registered in the US to avoid tion. Applied as a seed dressing, it delays germination of
fruit set in such trees or shrubs. Typically, licensed ‘‘male’’ inbred lines, which can be useful for widening the
applicators with special equipment have to be employed pollen shed window.
for tree and shrub treatment.
PGRs Used in Pasture and Turf Grasses Conclusions and Outlook: PGRs Between Rocket
and in Grass Grown for Seed Science and Hangover?
GA3 has found some use in the US and other countries to 1-MCP is, indeed, structurally related to certain cyclo-
stimulate shoot growth in pasture grasses (Matthew and propanes, which have been used for many years as rocket
others 2009). However, much more interest is directed fuel in the former Soviet Union. Likewise, cyanamide is
towards reducing shoot elongation. Here, inhibitors of GA not only applied to break dormancy in crop plants. It has
biosynthesis are important in high-intensity fine turf, also found use as a pharmaceutical for the treatment of
particularly on golf courses. A main reason is to reduce alcohol misuse. After administration, cyanamide inhibits
vertical leaf growth, which leads to smoother and more the enzyme aldehyde dehydrogenase, which is involved in
uniform playing surfaces. Darker leaf color, intensified ethanol metabolism. Any consumption of alcohol would
root growth, reduced water consumption, seed head sup- then result in an intense hangover because of increases in
pression of unwanted annual bluegrass (Poa annua), and, blood acetaldehyde levels.
not least, less need for mowing are additional benefits. Now, where are we with PGRs and what is likely to
Trinexapac-ethyl, paclobutrazol, flurprimidol, and meflu- occur with this type of agrochemical in the future? In the
idide are the main PGRs used for this purpose in the US. 1970s, PGRs had become very ‘‘fashionable’’ in pure and
Similar products for growth regulation in fine turf grasses applied plant sciences. It was generally expected that
are available in several other countries. So far, prohexa- almost all problems in plant production could be overcome
dione-calcium is available as a PGR for use on turf by PGRs or related chemicals and most agrochemical
grasses only in Germany. A recent survey of growth companies spent lots of resources in finding and developing
regulators in turfgrasses is available from March and new PGRs. Similarly, plant hormones and PGRs and their
others (2013). applied aspects were of major interest in academia.
As in cereal grain production, lodging may also be a Meanwhile, things have drastically changed and become
problem when grasses are grown for seed production. more realistic—if not to say sobering. The industry is
Trinexapac-ethyl and prohexadione-calcium are the main putting much more emphasis on solving agronomic prob-
active ingredients used to reduce this risk, particularly in lems by introducing improved fungicides, herbicides, or
the US state of Oregon, where grass grown for seed is a insecticides, because such products require less investment,
major business. Prohexadione-calcium has a small advan- are less ‘‘complicated’’, and have a larger market. Breeding
tage in performance. Most likely due to its more immediate new and better varieties with advanced technologies ranks
action after application, it inhibits seedhead formation in also high on the agenda of several companies. In contrast,
unwanted P. annua (Beam and Askew 2005). PGRs represent just a niche market. Likewise, it is also
123
obvious that there is less work on PGR applications in Edinburgh (quoted in: Annual Report 2014/2015, German Crop
academic research. Protection Association, Frankfurt)
Arndt FR, Rusch R, Stillfried HV, Hanisch B, Martin WC (1976) SN
In spite of the mentioned reduction of interest, PGRs 49537, a new cotton defoliant. Plant Physiol 57:S-99
are well established in agriculture, horticulture, and viti- Assante G, Merlini L, Nasini G (1977) (?)-Abscisic acid, a
culture (Tables 1, 2). The products existing on the market metabolite of the fungus Cercospora rosicola. Experientia
are currently embedded in a fairly stable economic 33:1556–1557
Baker JE, Lieberman M, Anderson JD (1978) Inhibition of ethylene
environment. At least for the time being, it is difficult to production in fruit slices by a rhizobitoxine analog and free
imagine alternatives for PGRs, which, for instance, steer radical scavengers. Plant Physiol 61:886–888
fruit formation in pineapples, optimize fruit set in pome Baker CJ, Sterling M, Berry P (2014) A generalised model of crop
fruits, stimulate latex flow in rubber plants, or adjust lodging. J Theoret Biol 363:1–12
Bakshi A, Shemansky JM, Chang C, Binder B (2015) History of
cotton plants to an annual production schedule. Because research on the plant hormone ethylene. J Plant Growth Regul
of the enormous costs involved and the low degree of 34. doi:10.1007/s00344-015-9522-9
interest currently existing at most agrochemical compa- Barbier S, Mayr HH (1966) Untersuchungen zur Wechselwirkung
nies, the probability for the introduction of new PGRs zwischen Stickstoff und Chlorcholinchlorid (CCC) bei Winter-
weizen im Gefäszversuch. Plant Soil 24:167–177
appears to be relatively low. However, one may expect Barrett JE (1982) Chrysanthemum height control by ancymidol,
that additional markets or additional uses for the estab- PP333, and EL-500 dependent on medium composition [growth
lished PGRs will be found. Improved formulations and regulators, pine bark, daminozide]. HortScience 17:896–897
new combinations of registered compounds will also be of Bartholomew DP (2014) History and perspectives on the role of
ethylene in pineapple flowering. Acta Hort 1042:269–283
interest. Also, one should not exclude the possibility that Basra AS (2000) Plant Growth Regulators in Agriculture and
some active ingredients lose registration due to toxico- Horticulture. Food Products Press, New York
logical concerns. In such cases, there would be a demand Beam JB, Askew SD (2005) Prohexadione calcium effects on
to fill the gap with equivalent alternatives. Enabling a bermudagrass, Kentucky bluegrass, perennial ryegrass, and
zoysiagrass. Internat Turfgrass Soc Res J 10:286–295
simple, safe, and reliable spray application of 1-MCP Berry PM, Spink JH (2009) Understanding the effect of a triazole
would be of enormous value. 3-Cyclopropyl-1-enyl-pro- with anti-gibberellin activity on the growth and yield of oilseed
panoic acid sodium salt is a pre-drug of a highly specific rape (Brassica napus). J Agric Sci 147:273–285
ethylene receptor blocker (Huberman and others 2014). Berry PM, Spink J (2012) Predicting yield losses caused by lodging in
wheat. Field Crops Res 137:19–26
This non-gaseous compound is well-suited for spray Berry PM, Sterling M, Spink JH, Baker CJ, Sylvester-Bradley R,
application and deserves thorough evaluation for practical Mooney SJ, Tams AR, Ennos AR (2004) Understanding and
use. All in all, one can judge that PGRs not only have a reducing lodging in cereals. Adv Agron 84:217–271
history—they also have a future! Beyer EM (1976) A potent inhibitor of ethylene action in plants. Plant
Physiol 58:268–271
Bocion PF, de Silva WH, Hüppi GA, Szkrybalo W (1975) Group of new
chemicals with plant growth regulatory activity. Nature 258:142–144
Böll S, Hofmann H, Schwappach P (2009) Rebsortenspezifische
References Unterschiede in Fertilitätsparametern und ihre Bedeutung für die
Anwendung der Wachstumsregulatoren GA3 und Prohexadion-
Abeles FB, Morgan PW, Saltveit ME (1992) Introduction and Calcium. Mitt Klosterneuburg 59:209–216
historical perspectives. In: Abeles FB, Morgan PW, Saltveit ME Burton JD, Pedersen MK, Coble HD (2008) Effect of cyclanilide on
(eds) Ethylene in plant biology, 2nd edn. Academic Press Inc, auxin activity. J Plant Growth Regul 27:342–352
San Diego, pp 1–13 Carlson RD, Crovetti AJ (1990) Commercial uses of gibberellins and
Abraham PD (1992) Tapping of Hevea brasiliensis. In: Sethuraj MR, cytokinins and new areas of applied research. In: Pharis RP,
Mathew NT (eds) Natural rubber: biology, cultivation and Rood SB (eds) Plant growth substances 1988. Proceedings of the
technology. Elsevier Science Publishers B.V., Amsterdam, 13th international conference on plant growth substances,
pp 263–281 Calgary, 17–26 July, 1988, Springer-Verlag, Berlin, pp 604–610
Adams R, Kerber E, Pfister K, Weiler EW (1992) Studies on the Casanova L, Casanova R, Moret A, Agusti M (2009) The application
action of the new growth retardant CGA 163’935 (cimectacarb). of gibberellic acid increases berry size of ‘Emperatriz’ seedless
In: Karssen CM, van Loon LC, Vreugdenhil D (eds) Progress in grape. Span J Agric Res 7:919–927
plant growth regulation. Proceedings of the 14th international Cathey HM (1964) Physiology of growth retarding chemicals. Ann
conference on plant growth substances, Amsterdam, 21–26 July, Rev Plant Physiol 15:271–302
1991, Kluwer Academic Publishers, Dordrecht, pp 818–827 Chaney WR (2005) Growth retardants: A promising tool for
Amasino R (2005) 1955: Kinetin arrives. The 50th anniversary of a managing urban trees. Publication FNR-252-W, Purdue exten-
new plant hormone. Plant Physiol 138:1177–1184 sion, Purdue University, Purdue. pp 1–5. https://www.extension.
Anonymous (2013) R&D trends for chemical crop protection purdue.edu/extmedia/fnr/fnr-252-w.pdf. Accessed 26 July 2015
products and the position of the European Market. Phillips Cobb AH, Reade JPH (2010) Herbicides and plant physiology, 2nd
McDougall, Crop Protection & Biotechnology Consultants, edn. Wiley-Blackwell, Chichester
Edinburgh. http://www.ecpa.eu/files/attachments/R_and_D_ Coggins CW, Lovatt CJ (2014) Plant growth regulators. In: Ferguson
study_2013_v1.8_webVersion_Final.pdf. Accessed 26 July 2015 L, Grafton-Cardwell EE (eds) Citrus production manual: The
Anonymous (2015) Global Crop Protection Market 2014, Phillips Regents of the University of California. Agriculture and Natural
McDougall, Crop Protection & Biotechnology Consultants, Resources Communication Services, Richmond, pp 215–226
123
Cooke AR, Randall DI (1968) 2-Haloethanephosphonic acids as Hagel JM, Facchini PJ (2010) Dioxygenases catalyze the O-demethy-
ethylene releasing agents for the induction of flowering in lation steps of morphine biosynthesis in opium poppy. Nat Chem
pineapples. Nature 218:96–97 Biol 6:273–275
Cotterill, P. (Nov 17, 2005) Method of altering the alkaloid composition Hagemann MH, Roemer MG, Kofler J, Hegele M, Wünsche JN
in poppy plants. Internat Pat Appl WO/2005/107436 (2014) A new approach for analyzing and interpreting data on
Dean GJ (2011) Modifying poppy growth and alkaloid yield with fruit drop in mango. HortScience 49:1498–1505
plant growth regulators. Master Thesis, University of Tasmania, Handley DT, Dill JF, Moran RE (2009) Prohexadione-calcium
Hobart applications to suppress runner growth in strawberries grown
Dennis FG (2000) The history of fruit thinning. Plant Growth Regul in a plasticulture system. Acta Hort 842:801–804
31:1–16 Harpaz-Saad S, Yoon GM, Mattoo AK, Kieber JJ (2012) The
Denny FE (1924) Hastening the coloration of lemons. J Agric Res formation of ACC and competition between polyamines and
27:757–769 ethylene for SAM 53. In: McManus MT (ed) Ann Plant Rev, vol
Dokoozlian NK, Peacock WL (2001) Gibberellic acid applied at 44. The plant hormone ethylene, Wiley-Blackwell, Hoboken,
bloom reduces fruit set and improves size of ‘Crimson Seedless’ pp 53–81
table grapes. HortScience 34:706–709 Hedden P (2016) Gibberellin biosynthesis in higher plants. In:
Easson DL, White EM, Pickles SJ (1993) The effects of weather, seed Hedden P (ed) Annual plant reviews, vol 49, The Gibberellins.
rate and cultivar on lodging and yield in winter wheat. J Agric Wiley-Blackwell, Hoboken
Sci 121:145–156 Hedden P, Sponsel VM (2015) A century of gibberellin research.
Edmisten KL, Yelverton FH, Bacheler JS, Koening SR, Crozier J Plant Growth Regul 34. doi:10.1007/s00344-015-9546-1
CR, Meijer AD, York WNR, Hardy DH, Cleveland BR (2014) Hedden P, Thomas SG (2012) Gibberellin biosynthesis and its
2014 Cotton Information, North Carolina State University, regulation. Biochem J 444:11–25
Raleigh. Available at http://cotton.ces.ncsu.edu/wp-content/ Hisamatsu T, Koshioka M, Kubota S, King RW (1998) Effect of
uploads/2014/02/2014-Cotton-Information.pdf. Accessed 26 gibberellin A4 and GA biosynthesis inhibitors on growth and
July 2015 flowering of stock [Matthiola incana (L.) R. Br.]. J Jpn Soc Hort
El-Otmani M, Coggins CW, Agusti M, Lovatt CJ (2000) Plant growth Sci 67:537–543
regulators in citriculture: world current uses. Crit Rev Plant Sci Hornsey IS (2003) A history of beer and brewing. The Royal Society
19:395–447 of Chemistry, Cambridge
Fan X, Mattheis JP, Buchanan D (1998) Continuous requirement of Huberman M, Riov J, Goldschmidt EE, Apelbaum A, Goren R (2014)
ethylene for apple volatile synthesis. J Agric Food Chem The novel ethylene antagonist, 3-cyclopropyl-1-enyl-propanoic
46:1959–1963 acid sodium salt (CPAS), increases grain yield in wheat by
Fritz CD (1997) FinishÒ cotton harvest-aid EUP results. In: Davis TD delaying leaf senescence. Plant Growth Regul 73:249–255
(ed) Proceedings of the 23rd Annual Meeting of the Plant Hytönen T, Mouhu K, Koivu I, Junttila O (2008) Prohexadione-
Growth Regulation Society of America, Plant Growth Regula- calcium enhances the cropping potential and yield of strawberry.
tion Society of America, Research Triangle Park, pp 224–229 Eur J Hort Sci 73:210–215
Gardner FE, Marth PC, Batjer LP (1939) Spraying with plant growth Izumi K, Yamaguchi I, Wada A, Oshio H, Takahashi N (1984) Effects of
substances to prevent apple fruit dropping. Science 90:208–209 a new plant growth retardant (E)-1-(4-chlorophenyl)-4,4-dimethyl-
Garthwaite DG, Hudson S, Barker I, Parrish G, Smith L, Pietravalle 2-(1,2,4-triaxol-1-yl)-penten-3-ol (S-3307) on the growth and
S (2013) Arable crops in the United Kingdom 2012. Pesticide gibberellin content of rice plants. Plant Cell Physiol 25:611–617
Usage Survey Report No. 250. Food & Environment Research Jagadeeswari P, Sharma SP, Dadlani M (2004) Effect of different
Agency, York. https://secure.fera.defra.gov.uk/pusstats/surveys/ chemicals on traits favouring outcrossing and optimization of
documents/arable2012.pdf. Accessed 26 July 2015 GA3 for seed production of cytoplasmic male sterile line in
Gavino RB, Pi Y, Abon CC (2008) Application of gibberellic acid hybrid rice. Seed Sci Technol 32:473–483
(GA3) in dosage for three hybrid rice seed production in the Jordan DL, Beam JB, Johnson PD, Spears JF (2001) Peanut response
Philippines. J Agric Technol 4:183–192 to prohexadione calcium in three seeding rate-row pattern
Giacomelli L, Rota-Stabelli O, Masuero D, Kwame Acheampong planting systems. Agron J 93:232–236
A, Moretto M, Caputi L, Vrhovsek U, Moser C (2013) Jordan DL, Nuti RC, Beam JB, Lancaster SH, Lanier JE, Lassiter BR,
Gibberellin metabolism in Vitis vinifera L. during bloom and Johnson PD (2008) Peanut (Arachis hypogaea L.) cultivar
fruit set: functional characterization and evolution of grapevine response to prohexadione calcium. Peanut Sci 35:101–107
gibberellin oxidases. J Exp Bot 64:4403–4419 Jung J, Sturm H (1964) Wachstumsregulierende Wirkung von
Greene DW (2010) The development and use of plant bioregulators in Chlorcholinchlorid (CCC). Landwirtschaftl Forsch 17:1–9
tree fruit production. Acta Hort 884:31–40 Kabachnik MI, Rossiiskaya PA (1946) Organophosphorus com-
Greene DW, Costa G (2013) Fruit thinning in pome- and stone-fruit: pounds. I. Reaction of ethylene oxide with phosphorous
state of the art. Acta Hort 998:93–102 trichloride. Izv Akad Nauk SSSR Khim Nauk 406:295–304 (in
Greene DW, Autio WR, Erf JA, Mao ZY (1992) Mode of action of Russian, quoted from Chem Abstr 41:7241)
benzyladenine when used as a chemical thinner on apples. J Am Kaminek M (2015) Tracking the story of cytokinin research. J Plant
Soc Hort Sci 117:775–779 Growth Regul 34. doi:10.1007/s00344-015-9543-4
Grossmann K (1991) Induction of leaf abscission in cotton is a Kauffmann M, Vorländer D (1910) Über den Nachweis des Cholins,
common effect of urea- and adenine-type cytokinins. Plant nebst Beiträgen zur Kenntnis des Trimethylamins. Ber Dtsch
Physiol 95:234–237 Chem Ges 43:2735–2743
Grossmann K, Jung J (1984) The influence of new terpenoid Kerber E, Leypoldt G, Seiler A (1990) CGA 163’935, a new plant
analogues of abscisic acid on stomatal movement and leaf growth regulator for small grain cereals, rape and turf. Proceed-
senescence. J Agron Crop Sci 153:14–22 ings Brighton Crop Protection Conference: Weeds, 1989. The
Gwathmey CO, Clement JD (2010) Alteration of cotton source-sink British Crop Protection Council, Farnhem, pp 83–88
relations with plant population density and mepiquat chloride. Kerby TA, Hake K, Keeley M (1986) Cotton fruiting modification
Field Crops Res 116:101–107 with mepiquat chloride. Agron J 78:907–912
123
Kerby TA, Weir BL, Keeley MP (1996) The uses of PixTM. In: Hake Molitor D, Behr M, Hoffmann L, Evers D (2012) Research note:
SJ, Kerby TA, Hake KD (eds) Cotton production manual, benefits and drawbacks for pre-bloom applications of gibberellic
Publication No. 3352. University of California, Oakland, acid (GA3) for stem elongation in Sauvignon blanc. S Afric J
pp 294–304 Enol Viticult 33:198–202
Kightley S (2001) Lodging control in winter rape. Plant Var Seeds Moody K (1986) Role of plant growth regulators in tropical rice
14:171–180 cultivation. In: Macgregor P, Saito Y, Wang R (eds) Plant
Kondo S (2010) Roles of jasmonates in fruit ripening and environ- growth regulators in agriculture. Food and Fertilizer Technology
mental stress. Acta Hort 884:711–716 Center for the Asian and Pacific Region, Taipei, pp 140–147
Kuck KH, Berg D (1986) Systemische Eigenschaften und biochemis- Müller R, Bocion PF, de Silva WH (1980) Die physiologische
cher Wirkungsmechanismus eines neuen Azolfungizids. Mitt Aktivität von Aminoäthoxyvinylglycine auf die Seneszenz von
Biol Bundesanst Land- Forstwirtschaft Berlin-Dahlem 232:196 Früchten und Schnittblumen. In: Bangerth F (ed) Anwen-
Kuroi I, Shiraishi Y, Imano S (1963) Studies on breaking the dungsmöglichkeiten von Phytohormonen und Wachstumsregu-
dormancy of grapevines. J Jpn Soc Hortic Sci 32:175–180 latoren in der Pflanzenproduktion. Eugen Ulmer, Stuttgart,
Lacote R, Gabla O, Obouayeba S, Eschbach JM, Rivano F, Dian K, pp 175–180
Gohet E (2010) Long-term effect of ethylene stimulation on the Nakayama I, Miyazawa T, Kobayashi M, Kamiya Y, Abe H, Sakurai
yield of rubber trees is linked to latex cell biochemistry. Field A (1990) Effects of a new plant growth regulator prohexadione
Crops Res 115:94–98 calcium (BX-112) on shoot elongation caused by exogenously
Lever BG, Shearing SJ, Batch JJ (1982) PP 333-a new broad spectrum applied gibberellins in rice (Oryza sativa L.) seedlings. Plant
growth retardant. Brighton crop protection conference—Weeds Cell Physiol 31:195–200
1982, vol 1. British Crop Protection Council, Croydon, pp 3–10 Neljubow D (1901) Über die horizontale Nutation der Stengel von
Lim SC, Kim SK, Kim YH, Youn CK, Yoon T (2004) Vine growth Pisum sativum und einiger anderer Pflanzen. Beih Bot Centralbl
and fruit quality of ‘Kyoho’ grapes as affected by mepiquat 10:128–139
chloride and GA. Acta Hort 653:145–149 Nickell LG (1983) Plant Growth Regulating Chemicals, vols 1 & 2.
Lindstrom RS, Tolbert NE (1960) (2-Chloroethyl)trimethylammo- CRC Press, Boca Raton
nium chloride and related compounds as plant growth sub- Nishiyama I (1986) Lodging of rice plants and countermeasures. In:
stances. IV. Effect on chrysanthemums and poinsettias. Macgregor P, Saito Y, Wang R (eds) Plant growth regulators in
Michigan Agric Exp Stn Quart Bull 42:917–928 agriculture. Food and Fertilizer Technology Center for the Asian
Linser H, Mayer HH, Bodo G (1961) Über die Wirkung von and Pacific Region, Taipei, pp 152–163
Chlorcholinchlorid auf Sommerweizen. Bodenkultur Norton ER, Borrego HJ (2006) Evaluation of plant growth regulator
12:279–280 formulations in Arizona cotton production systems. 2006 Cotton
Looney N, Jackson D (2011) Use of bioregulators in fruit production. Report (Series P-145). The University of Arizona, Tucson,
In: Jackson D, Looney NE, Morley-Bunker M, Thiele G (eds) pp 143–152
Temperate & Subtropical Fruit Production, 3rd edn. CABI, Owens LD, Lieberman M, Kunishi A (1971) Inhibition of ethylene
Wallingford, pp 92–100 production by rhizobitoxin. Plant Physiol 48:1–4
Lüthen H (2015) What we can learn from old auxinology. J Plant Peppi MC, Fidelibus MW (2008) Effects of forchlorfenuron and
Growth Regul 34. doi:10.1007/s00344-015-9527-4 abscisic acid on the quality of ‘Flame Seedless’ grapes.
Macgregor P, Saito Y, Wong CM (1986) Plant growth regulators in HortScience 43:173–176
agriculture. FFTC Book Series No. 34, Food and Fertilizer Petgen M (2005) Gibberellin-Einsatz zur Qualitätsregulierung. Sch-
Technology Center for the Asia and Pacific Region, Taipei weizer Z Obst- Weinbau 2005(7):6–9
Mander L (2003) Twenty years of gibberellin research. Nat Prod Rep Petracek PD, Silverman FP, Greene DW (2003) A history of
20:49–69 commercial plant growth regulators in apple production. Hort-
March SR, Martins D, McElroy JS (2013) Growth inhibitors in Science 38:937–942
turfgrass. Planta Daninha 31:733–747 Pires MC, Yamanishi OK (2014) Girdling combined with paclobu-
Martin GC (1983) Commercial uses of gibberellins. In: Crozier A (ed) trazol boosted yield of ‘Bengal’ lychee in Brazil. Acta Hort
The Biochemistry and Physiology of Gibberellins. Praeger 1042:189–195
Publishers, New York, pp 395–444 Pruess DL, Scannell JP, Kellett M, Ax HA, Janecek J, Williams TH,
Marumo S, Katayama M, Komori E, Ozaki Y, Natsume M, Kondo S Stempel A, Berger J (1974) Antimetabolites produced by
(1982) Microbial production of abscisic acid by Botrytis cinerea. microorganisms. X L-2-Amino-4-(2-aminoethoxy)-trans-3-bute-
Agric Biol Chem 46:1967–1968 noic acid. J Antibiot 27:229–233
Matthew C, Hofmann WA, Osborne MA (2009) Pasture response to Rademacher W (2000) Growth retardants: effects on gibberellin
gibberellins: a review and recommendations. N Z J Agric Res biosynthesis and other metabolic pathways. Ann Rev Plant
52:213–225 Physiol Plant Mol Biol 51:501–531
Maynard JA, Swan JM (1963) Organophosphorus compounds. I. Rademacher W (2004) Recent situation and trends in global plant
2-Chloroalkylphosphonic acids as phosphorylating agents. Aus J bioregulator utilization. Regul Plant Growth Devel 39:142–151
Chem 16:596–608 Rademacher W (2010a) Dealing with plant bioregulators: an indus-
McLaren JS (1982) Chemical manipulation of crop growth and trial view. Acta Hort 884:717–724
development. Butterworth Scientific, London Rademacher W (2010b) Control of lodging in intense European cereal
Menzel CM, Le Lagadec MD (2014) Increasing the productivity of production. In: Whipker B (ed) Proceedings of the 36th Annual
avocado orchards using high-density plantings: a review. Sci Meeting of the Plant Growth Regulation Society of America,
Hort 177:21–36 August 2–6, 2009, Asheville, NC, USA, The Plant Growth
Mitchell JE, Angel CR (1950) Plant-growth-regulating substances Regulation Society of America, LaGrange, pp 61–69
obtained from cultures of Fusarium moniliforme. Phytopathol Rademacher W (2016) Chemical regulators of gibberellin status and
40:872 their application in plant production. In: Hedden P (ed) Annual
Mitchell JW, Wirwille JW, Weil L (1949) Plant growth-regulating plant reviews, vol 49, The Gibberellins. Wiley-Blackwell,
properties of some nicotinium compounds. Science 110:252–254 Hoboken
123
Rademacher W, Brahm L (2010) Plant growth regulators. In: Shibata H, Takebayashi Y (2007) Development of a forefront one-
Ullmann’s encyclopedia of industrial chemistry, Wiley-VCH, shot basal application fertilizer ‘‘RakuichiÒ’’ for paddy rice.
Weinheim, doi:10.1002/14356007.a20_415.pub2 Sumitomo Kagaku 2007-I, 1–13 (English translation)
Rademacher W, Graebe JE (1979) Gibberellin A4 produced by Shulman Y, Nir G, Fanberstein L, Lavee S (1983) The effect of
Sphaceloma manihoticola, the cause of the superelongation cyanamide on the release from dormancy of grapevine buds. Sci
disease of cassava (Manihot esculenta). Biochem Biophys Res Hort 19:97–105
Commun 91:35–40 Sisler EC, Blankenship SM (May 21, 1996). Methods of counteract-
Rademacher W, Speakman JB, Evans RR, Evans JR, Römmelt S, ing an ethylene response in plants. US Patent Number 5,518,988
Michalek S, Lux-Endrich A, Treutter D, Iturriagagoitia-Bueno T, Skoog F (1994) A personal history of cytokinin and plant hormone
John P (1999) Prohexadione-Ca—a new plant growth regulator research. In: Mok DWS, Mok MC (eds) Cytokinins: Chemistry,
for apple with interesting biochemical features. In: Shafer WE activity and functions. CRC Press Inc, Boca Raton, pp 1–14
(ed) Proceedings of the 25th Annual Meeting of The Plant Smith RH (2013) Media components and preparation. In: Smith RH
Growth Regulation Society of America, July 7–10, 1998, (ed) Plant Tissue Culture. Academic Press, London, pp 31–43
Chicago, IL, USA, The Plant Growth Regulation Society of Stapleton GS, Via JD (2003) PentiaTM (BAS 130): next generation plant
America, LaGrange, pp 113–118 growth regulator for enhanced growth control, boll retention and
Rademacher W, van Saarloos K, Garuz Porte JA, Riera Forcades F, cotton lint yield. In: Dugger P, Richter D (eds) Proceedings of the
Senechal Y, Andreotti C, Spinelli F, Costa G (2004) Impact of Beltwide Cotton Conference, 6–10 January 2003. Nashville, TN,
prohexadione-Ca on the vegetative and reproductive perfor- National Cotton Council, Memphis, TN, pp 1792–1793
mance of apple and pear trees. Europ J Hort Sci 69:221–228 Stellman JM, Stellman SD, Christian R, Weber T, Tomasallo C
Rademacher W, Spinelli F, Costa G (2006) Prohexadione-Ca: modes (2003) The extent and patterns of usage of Agent Orange and
of action of a multifunctional plant bioregulator for fruit trees. other herbicides in Vietnam. Nature 422:681–687
Acta Hort 727:97–106 Suttle JC (1985) Involvement of ethylene in the action of the cotton
Reekie JY, Hicklenton PR, Duval JR, Chandler CK, Struik PC (2005) defoliant thidiazuron. Plant Physiol 78:272–276
Leaf removal and prohexadione-calcium can modify Camarosa Takahashi S, Shudo K, Okamoto T, Yamada K, Isogai Y (1978)
strawberry nursery plant morphology for plasticulture fruit Cytokinin activity of N-phenyl-N0 -(4-pyridyl)urea derivatives.
production. Can J Plant Sci 85:665–670 Phytochemistry 17:1201–1207
Resende PAP, Soares JE, Hudetz M (2000) MODDUSÒ, a plant Tolbert NE (1960a) (2-Chloroethyl)-trimethylammoniumchloride and
growth regulator and management tool for sugarcane production related compounds as plant growth substances. I. Chemical
in Brazil. Internat Sugar J 102:5–9 structure and bioassay. J Biol Chem 235:475–479
Rhodes D, Hanson AD (1993) Quaternary ammonium and tertiary Tolbert NE (1960b) (2-Chloroethyl)-trimethylammoniumchloride and
sulfonium compounds in higher plants. Ann Rev Plant Physiol related compounds as plant growth substances. II. Effect on
Plant Mol Biol 44:357–384 growth of wheat. Plant Physiol 35:380–385
Riddell JA, Hageman HA, J’Anthony CM, Hubbard WL (1962) Tschabold EE, Taylor HM, Davenport JD, Hackler RE, Krunkalns
Retardation of plant growth by a new group of chemicals. EV, Meredith WS (1970) A new plant growth regulator. Plant
Science 136:391 Physiol 46:S-19
Rixon CM, Di Bella LP, Kingston G, Dorahy K, Davies B, Wood AW Van de Poel B, Ceusters J, De Proft MP (2009) Determination of
(2007) Moddus—a sugar enhancer. Proc Austral Soc Sugar Cane pineapple (Ananas comosus, MD-2 hybrid cultivar) plant
Technol 29:318–327 maturity, the efficiency of flowering induction agents and the
Rodriguez AB (1932) Influence of smoke and ethylene on the fruiting use of activated carbon. Sci Hort 120:58–63
of pineapple (Ananas sativus Schult.). J Dept Agric Puerto Rico Veen H, van de Geijn SC (1978) Mobility and ionic form of silver as
16:5–18 related to longevity of cut carnations. Planta 140:93–96
Rubio S, Donoso A, Perez FJ (2014) The dormancy-breaking stimuli Warner HL, Leopold AC (1967) Plant growth regulation by
‘‘chilling, hypoxia and cyanamide exposure’’ up-regulate the stimulation of ethylene production. Bioscience 17:722
expression of a-amylase genes in grapevine buds. J Plant Physiol Watkins CB, James H, Nock JF, Reed N, Oakes RL (2010) Preharvest
171:373–381 application of 1-methylcyclopropene (1-MCP) to control fruit
Salassi ME, Deliberto MA, Linscombe SD, Wilson CE, Walker TW, drop of apples, and its effect on postharvest quality. Acta Hort.
McCauley GN, Blouin DC (2013) Impact of harvest lodging on 877:365–374
rough rice milling yield and market price. Agron J Weaver RJ (1958) Effect of gibberellic acid on fruit set and berry
105:1860–1867 enlargement in seedless grapes of Vitis vinifera. Nature
Sampson AJ, Cazenave A, Laffranque JP, Jones RG, Kumazawa S, 181:851–852
Chida T (1992) Metconazole, an advance in disease control in Weaver RJ (1972) Plant Growth Substances in Agriculture. W. H.
cereals and other crops. Brighton crop protection conference: Freeman and Company, San Francisco
pests and diseases, vol 1. The British Crop Protection Council, Weaver RJ, Kasimatis AN, McCune SB (1962) Studies with
Farnhem, pp 419–426 gibberellin on wine grapes to decrease bunch rot. Am J Enol
Schott PE, Walter H (1991) Bioregulators: present and future fields of Viticult 13:78–82
application. In: Gausman H (ed) Plant Biochemical Regulators. Webster AD, Spencer JE (2000) Fruit thinning plums and apricots.
Marcel Dekker, New York, pp 247–321 Plant Growth Regul 31:101–112
Schupp JR, Greene DW (2004) Effect of aminoethoxyvinylglycine Wermann S, Theurillat V, Verzegnassi L, Hofmann J, Kuchenbecker
(AVG) on preharvest drop, fruit quality, and maturation of R, Constable A, Delatour T, Stadler RH (2014) N,
‘McIntosh’ apples. I. Concentration and timing of dilute N-dimethylpiperidinium (mepiquat) Part 2. Formation in roasted
applications of AVG. HortScience 39:1030–1035 coffee and barley during thermal processing. Food Add Contam:
Serek M, Sisler EC, Reid MS (1994) Novel gaseous ethylene binding Part A 31:234–241
inhibitor prevents ethylene effects in potted flowering plants. Wertheim SJ (2000) Developments in the chemical thinning of apple
J Am Soc Hort Sci 119:1230–1233 and pear. Plant Growth Regul 31:85–100
123
Whipker BE (2013) Growth regulators for floriculture crops in Zaharia LI, Walker-Simmon MK, Rodrı́guez CN, Abrams SR (2005)
greenhouses. Grower Talks, 2013 Plant Growth Regulator Chemistry of abscisic acid, abscisic acid catabolites and analogs.
Guide. Ball Publishing, West Chicago, pp 8–37 J Plant Growth Regul 24:274–284
Whipker BE, Latimer J (2013) Wide assortment of available PGRs. Zeeh B, König KH, Jung J (1974) Development of a new plant growth
Grower Talks, 2013 Plant Growth Regulator Guide. Ball regulator with biological activity related to CCC. Kemia
Publishing, West Chicago, pp 2–4 (Helsinki) 9:621–623
Wittwer SH, Tolbert NE (1960) (2-Chloroethyl)-trimethylammoni- Zimmerman PW, Hitchcock AE (1944) Substances effective for
umchloride and related compounds as plant growth substances. increasing fruit set and inducing seedless tomatoes. Proc Am Soc
III. Effect on growth and flowering of the tomato. Am J Bot Hort Sci 45:353–361
47:560–565 Zimmerman PW, Hitchcock AE, Wilcoxon F (1936) Several esters as
Yeshitela T, Robbertse PJ, Stassen PJC (2004) Effects of various plant hormones. Contrib Boyce Thompson Inst 8:105–112
inductive periods and chemicals on flowering and vegetative Zubkova NF, Bukashkina ZV, Kulaeva ON (1990) A study of
growth of ‘Tommy Atkins’ and ‘Keitt’ mango (Mangifera defoliating effects of cytokinins. Fiziol Rast 37:535–541 (in
indica). N Z J Crop Hort Sci 32:209–215 Russian)
Yim KO, Kwon YW, Bayer DE (1997) Growth responses and
allocation of assimilates of rice seedlings by paclobutrazol and
gibberellin treatment. J Plant Growth Regul 16:35–41
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J Plant Growth Regul

Plant Growth Regulators: Backgrounds and Uses in Plant

Received: 6 May 2015 / Accepted: 9 August 2015

Abstract Plant growth regulators are widely used in Introduction

Fig. 2 Structures of I gibberellin A3 = gibberellic acid = GA3 = gibberellin A7 = GA7 = (3S,3aR,4S,4aR,7R,9aR,9bR,12S)-12-hy-

Fig. 3 Structures of inhibitors of GA biosynthesis: I Chlormequat pentan-3-ol, VI uniconazole-P = (1E,3S)-1-(4-chlorophenyl)-4,4-

Fig. 4 Structures of cytokinin-type PGRs: I 6-Benzyladenine (BA) = N-Benzyl-3H-purin-6-amine, II thidiazuron = 1-phenyl-3-(1,2,3-

water, may also be used in this crop (Bartholomew 2014).

Fig. 6 Structures of chemicals

Inhibitors of Ethylene Biosynthesis and Blankenship at North Carolina State University at

Dikegulac (Fig. 8I) is readily available as a by-product

Examples of PGR Uses of lodging can be differentiated: Stem lodging is mainly

Indol-3-ylbutyric acid Auxin Rooting of cuttings RhizoponÒ Rhizopon B.V.

Paclobutrazol ? Difenoconazole Inhibitor GA biosynthesis Oilseed rape: ToprexÒ Syngenta Crop

6-Benzyladenine Cytokinin Pome fruits: Fruit thinning MaxCelÒ Valent

(S)-Abscisic acid Abscisin Table grapes: Improved berry ProToneÒ Valent

Hydrogen cyanamide Respiration inhibitor Fruit trees, grapevines: DormexÒ AlzChem AG

Trinexapac-ethyl Used in Opium Poppy different mepiquat-containing products give comparable

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