筑波大農林技研 第 5 号： 1 －８，2017

総説

Developmental Control of Horticultural Plants Using Strigolactone to
Improve Marketability

Seung Won KANG1,2*
1Faculty of Life and Environmental Sciences, University of Tsukuba,
1-1-1 Tennodai, Tsukuba, Ibaraki, 305-8572, Japan

2

Agricultural and Forest Research Center, University of Tsukuba,
1-1-1 Tennodai, Tsukuba, Ibaraki, 305-8577, Japan

Abstract
Height control, branching, adventitious root formation, shelf life are very important traits that determine thequality and marketability of horticultural plants so that various type of plant growth regulators and synthetic
analogs are widely used. In particular, branching and plant height control are the key traits for cut flower andpot plant production in floricultural industry. Strigolactone is a new phytohormone known to control branching
and leaf senescence in many plants. However, commercial application related to strigolactone biosynthesis andsignaling has not been developed despite its structure and physiological role has been identified. In this review,
we suggest potentiality of commercial application related to strigolactone in order not only to control growthand development but also to improve marketability of horticultural plants.
Keywords: Branching, Cuttings, Ethylene, Height control, Ripening, Shelf lifeand development in horticultural industry.
Synthetic auxins are not only used forstimulation of pigmentation, enlargement,
ripening of fruits such as tomato, eggplant,
melon but also used for prevention offruit falling of apple and pear (Ada et al.,2013; de Jong et al., 2009; Watanabe etal., 2008). Indole-3-butyric acid (IBA) and
1-naphthaleneacetic acid (NAA), synthetic
auxins, are used in order to stimulateadventitious root formation of apple stock
for grafting and various ornamental plants(Fig. 1A and B) (Divin et al., 2011; Karakurt
et al., 2009). 2, 4-dinitrophenylhydrazine (2,Introduction
Phytohormones, also called plant growth
regulators are small chemical substances
but profoundly influence the growth anddevelopment of plants. There are ﬁve classicalplant hormones such as abscisic acid (ABA),
auxin, cytokinin, gibberellin, and ethylene.
In addition, brassinosteroids, polyamines,
salicylic acid, peptides involved in cell-tocell signaling, and strigolactones are also
considered plant hormones. Various typesof synthetic phytohormones have been
widely used in order to control plant growth*Corresponding Auther: Seung Won KANG Faculty of Life and Environmental Sciences,
University of Tsukuba, 1-1-1 Tennodai, Tsukuba, Ibaraki, 305-8572, Japan
E-mail: kang.seungwon.ga@u.tsukuba.ac.jp

─1─

筑波大農林技研 第 5 号 20174-D) is widely used for herbicide (Fig. 1C)(Egana et al., 2011; Schulz and Segobye,2016). Gibberellins are used for inductionof seedless fruits, stimulation or delayof ripening, enlargement of cucumber
fruit, flowering stimulation of azalea andcyclamen, stimulation of germination andthe growth of flowering plants (Chen andChang, 1972; de Jong et al., 2009; Martínez
et al., 1994; Vendrell 1969; Yamaguchi andKamiya, 2001). In addition, benzyl adenine(6-benzyl aminopurine, BA), thidiazuron(TDZ), forchlorfenuron (1-(2-Chloro-4pyridyl)-3-phenylurea), belong to cytokinin
family and these are used as a stimulant
for the pigmentation and enlargement offruits, fruiting, and so on (Fig. 1D, E and F).
Ethephon, converted to ethylene in plant,
accelerates fruit ripening and pigmentation ofapple, persimmon, and peach and stimulates
ﬂowering of pineapple and mango (Fig. 1G).

Roldan et al., 2008; Umehara et al., 2008; Alderet al., 2012). In 1966, it was firstly reported
and its structure was identified (Cook et al.,
1966; Cook et al., 1972; Siame et al., 1993).
Strigolactones are derived from carotenoid
pathway, terpenoid lactones (Matusova etal., 2005; Alder et al., 2012). Various types ofstrigolactones have been identified in many
plant species. However, all strigolactones
have four rings as their common structure(Fig. 2). Three rings (ring A, B, and C) form
tricyclic lactone and another is butenoic group(ring D). Of four basic rings in strigolactone,
conformation of two rings (A and B) varies
depending on side groups whereas other two
rings, C and D, are not only highly conserved inmost strigolactones but also considered playing
an important role in biological activity (Xie etal., 2010).

Fig.2 Basic structure of strigolactones.
Red dot circle shows enol-ether bridge between
tricyclic lactone (ring A, B, and C) and butenoic
group (ring D) (modiﬁed from Kohlen et al., 2013).

Fig.1 Plant hormones in plants.
A: Indole-3-butyric acid (IBA), B:
1-naphthaleneacetic acid (NAA), C: 2,
4-dinitrophenylhydrazine (2, 4-D), D: benzyl
adenine (6-benzyl aminopurine, BA), E:
thidiazuron (TDZ), F: forchlorfenuron (1-(2-Chloro4-pyridyl)-3-phenylurea), G: Ethephon, H:
Strigolactone.

Height control, branching, adventitious root
formation and shelf life are very importantfeatures which determine the quality andmarketability of horticultural plants so thatvarious type of plant growth regulators,
synthetic analogs, inhibitors have been
widely applied. This review will focus on thephysiological role of strigolactones and thepotential applicability in order not only tocontrol growth and development but also to
improve marketability of horticultural plants.

Strigolactone is a brand new phytohormone
and plays an important role in inhibitionof branching in many plants the synthetic
analog, e.g. GR24, was developed and has been
widely used in hormonal regulation researchin plants (Fig 1 H). Strigolactone is involved in
the symbiosis with fungi by acting recognition
signals in the soil and controls shoot branchingin many plants (Akiyama et al., 2005; Gomez─2─Developmental Control of Horticultural Plants Using Strigolactone to Improve Marketability

et al., 2009; Beveridge and Kyozuka, 2010).
D14 is known to involve in the downstreamof strigolactone biosynthesis as a signaling
hormone or convert strigolactone to be activeby cleaving D-ring. D14 protein group, DAD2,
AtD14 and OsD14, share the same common
structure of a hydrophobic active site that
contains Ser-His-Asp. Three amino acids areconserved in all D14 protein groups. Nocturnal
condition or senescence signal induces
expression of strigolactone synthesis genes,
MAX3 and MAX 4, in the leaves of Arabidopsis
resulting in accelerated leaf senescence byenhancing strigolactone signal pathway (Ueda
and Kusaba, 2015). In addition, MAX3 andMAX4 are induced in the leaves under drought
and salt stresses (Ha et al., 2014) by suggesting
the strong possibility of strigolactone synthesis
in plant leaves.

Biosynthesis of Strigolactone in plant
A decade ago, it was suggested thatstrigolactones might be synthesized fromcarotenoid pathway (Matusova et al., 2005).
Various genes involved in strigolactones
biosynthesis and signaling have been identiﬁed
and characterized in Arabidopsis (AtD27,
MAX3, MAX4, MAX2, AtD14, BRC1), pea(Psd27, RMS5, RMS1, RMS4, PsBRC1), rice
(D27, D17/HTD1, D10, D3, D14, FC1), andpetunia (DAD3, DAD1, PhMAX2A-B, DAD2).
Precursor of strigolactones, carlactone isproduced by three steps. All-trans-β-carotene
is isomerized to 9-cis-β-carotene catalyzed by
D27 (Alder et al., 2012). 9-cis-β-carotene iscleaved by CCD7 so that it produces β-apo10`-carotenal which is then cleaved by CCD8
to produce carlactone. D-ring, one of the basic
structures of strigolactones, is formed in thisstep (Fig. 3).

Height control and Branching
Cut flowers account for the largest market
share in floricultural industry followed bypot plants. In order to produce high quality
cut flowers, long and strong flower stalkand one big flower are required. In addition
nutrient should be concentrated only a single
stalk. On the other hand, pot plants require
short plant height with large number ofbranches with many flowers. Consideringsuch requirements for cut flower and pot
plant production, branching control canbe the main key factor for cut flowers andpot plants quality, one branch versus manybranches. Pinching has been widely used tocontrol the plant height and induce branchingfor pot plants. When the apical meristem
was removed by pinching, plants lose apicaldominance governed by auxin synthesizedfrom apical meristem. Therefore, axillarybuds start to develop resulting in formationof new branches. In addition, various dwarf
reagents such as; ancymidol, daminozide,
paclobutrazol, chlormequat chloride, ethephon,and uniconazole-P are used for controlling
plant height by inhibiting stem elongation.
Ancymidol, paclobutrazol, and uniconazole-P

Fig.3 Strigolactone biosynthesis and genes involvedin plants (modiﬁed from Czarnecki et al.,2013 and de Saint Germain et al., 2013).

MAX2 isolated from Arabidopsis and D14(DWARF14) from rice, which belong to F-box
protein and α/β-hydrolase respectively,
are considered strigolactone receptor (Arite
─3─

筑波大農林技研 第 5 号 2017

are applied by a drench, whereas daminozide isapplied by a foliar spray. Chlormequat chloride
can be applied by both methods.
Several mutants were found with increasedshoot branching phenotype, more axillary
growth (max) in Arabidopsis (Stirnberg et al.,
2002; Sorefan et al., 2003; Booker et al., 2004),
ramous (rms) in pea (Beveridge et al., 1994,
1996; Beveridge et al., 2000), and dwarf (d) andhtd genes in rice (Ishikawa et al., 2005; Zou etal., 2006; Arite et al., 2007; Jiang et al., 2013).
e9 mutant (d53) of rice, a dominant mutant,
has a phenotype of dwarf and high-tillering
even though amount of 2`-epi-5-deoxystrigol ismuch higher in the wild type suggesting thismutant is involved in strigolactone signaling(Jiang et al., 2013). In this mutant, D53
proteins are accumulated so that it blocks
strigolactone signaling resulting in dwarf andhigh tillering phenotype. High-tillering dwarf(htd) in rice (Oryza sativa) produces a lot ofaxillary buds leading increased number oftillers and also htd showed dwarf phenotype
due to reduction of internode and panicle(Zou et al., 2005). A recessive gene, htd1,
controls high-tillering and dwarf phenotype
in rice. There is a nucleotide substitution in
OsCCD7 leading an amino acid substitution
from proline to leucine resulting in htd1
gene in mutant. OsCCD7 is an orthologous
gene of MAX3/CCD7 in Arabidopsis. For
max3 mutant, a recessive gene of MAX3
exhibits shorter petiole and reduced lengthof leaf blade (Booker et al., 2004). Therefore,
these mutants imply some genes involving
in the biosynthesis of strigolactone may berelated to form tight and small rosette whichcauses various physiological disorder offlowering perennials. For chrysanthemums
in particular, rosette is the one of thephysiological dormant phenomenon andappears when chrysanthemums are exposed
to high temperature followed by the lowtemperature during the autumn. Under
these environmental conditions, internodeof chrysanthemums is not elongated androsette, as a result, is formed resulting indelayed growth of chrysanthemum andchrysanthemum cannot ﬂower even if optimum
temperature and day-length are given.
Branching is the one of the importantphenotype required to produce pot plants infloricultural practice. Once plants lose thefunction of strigolactone resulting in increasing
number of branches, it affects plant height asa result. Therefore, branching and low plantheight can be controlled without pinching andapplication of dwarf reagents. Furthermore, itwill be possible to induce desirable phenotypes,
if we can control the strigolactones at a certain
level in plants or its signaling pathway. In
addition, this may contribute not only toprotect the environment by reducing theamount of dwarf reagents but also to reduce
the labor costs.
Adventitious root formation for cuttings
Cutting is the most common techniques
widely used for proliferation of clones whose
genetic background is identical. It is not only
the easy way to maintain good genotypiccharacteristics but also guarantees the same
quality for farmers. In order to stimulateadventitious root formation from cutting, NAA
is usually treated and adventitious roots areformed from the basal part.
It has been known that strigolactones inhibit
adventitious root formation in Arabidopsis
and pea (Rasmussen et al., 2012b). It was also
reported that the formation of adventitious
root was promoted, when biosynthesis of
strigolactone was inhibited (Rasmussen etal., 2012a). Number of adventitious roots was
signiﬁcantly increased for both mutants withdefective in biosynthesis of strigolactones andin response. For three mutants defective inbiosynthesis, max1, max3, and max4, numberof adventitious root increased by 3 times at
maximum than wild type. max1, unresponsive
mutant to strigolactone signal, showed 5 times
of increase in number of adventitious root in
Arabidopsis (Rasmussen et al., 2012b). GR24,
synthetic strigolactone, inhibited adventitious
root formation of max1, max3, and max4
whereas max2 formed adventitious root even─4─

Developmental Control of Horticultural Plants Using Strigolactone to Improve Marketability

though GR24 is treated. Therefore, theseresults suggest strigolactone signaling rather
than biosynthesis pathway is involved in theadventitious root formation of cuttings.
In addition, Rasmussen et al. (2012a)
suggested inhibition of strigolactone would
improve adventitious root formation ofhorticultural plants. Fluridone, known as acarotenoid inhibitor and herbicides widelyused, was treated to five commercial plants,
Tradescantia fluminensis (Wandering Jew),
Trachelospermum jasminoides (Chinese
jasmine), Jasminum polyanthum (Pinkjasmine), Plumbago auriculata (Capeleadwort), and Pongamia pinnata (Indian
Beech Tree) (Rasmussen et al., 2012a).
Application of fluridone improved rooting
and number of adventitious roots of P.
auriculata and J. polyanthum cuttingsand optimum concentration was 100 nM.
However, application did not affect rootingof T. fluminensis and T. jasminoides. For the
Genus Tradescantia, stem cuttings is thetechnique widely used and adventitious roots
are formed very well even though synthetic
auxin for inducing rooting is not treated on thebasal part. Combined application of ﬂuridone
with commercial rooting hormone did not
improve the rooting percentage compared torooting hormone treatment, moreover, numberof adventitious roots in P. auriculata and J.
polyanthum decreased.
T. fluminensis is monocot and herbaceous
plant and T. jasminoides woody and hasvery long stems. Climbing is the common
characteristic of Both plants. In particular,
T. jasminoides is likely to restore apicaldominance when apical meristem is removed
because the very next node produces axillary
bud and then new main stem elongates. J.
polyanthum and P. auriculata are shrubs
and have climbing characteristics. However,there is lack of information about effect of
combination or fluridone only treatment on
rootings of herbaceous horticultural plants
such as chrysanthemums, carnations,poinsettias, bulbs, cacti, woody plants, etc.
These plants are used as pot plants and cut

flowers and accounts for large amount ofannual yield. Therefore, it is necessary to
identify how strigolactone signaling affects
adventitious root formation in various
horticultural plants in order to develop
practical techniques to stimulate rooting.
Ripening and Senescence process of fruits andvegetables
Quality control of postharvest fruits,
vegetables, and flowers are very importantdue to their short shelf-life (Paliyath et al.,2008). During the ripening processes of fruits
or senescence process of leafy vegetables or
ﬂowers, various physiological and biochemicalchanges occur from a cellular level to tissuelevel, by changing biochemical characteristics
such as color, flavor, texture, and taste.
Ethylene biosynthesis and signaling is oneof the key factors for fruit ripening andsenescence of leafy vegetables and cut ﬂowers.
Methionine, for the first step in ethylene
biosynthesis, is converted to S-Adenosyl
methionine (SAM) by Methionine adenosyltransferase and then, as a sequential step,
SAM is converted to 1-aminocyclopropane-1carboxylic acid (ACC) by ACC synthase. ACC
oxidase oxidizes ACC to produce ethylene(Wang et al., 2002). Leaves of Arabidopsis
mutants lack of strigolactone biosynthesis
showed delayed senescence phenotype under
the dark condition (Ueda and Kusaba, 2015),
whereas MAX3 and MAX4, genes involved inbiosynthesis of strigolactones in Arabidopsis,
were strongly expressed by ethylene treatment
under the same condition. In addition, it issuggested strigolactones is synthesized in theleaf under abiotic stresses such as drought
and salt stresses (Ha et al., 2014), although
it has been widely known that strigolactone
or its precursor are synthesized in the root
and transported through xylem (Kohlen etal., 2011). Therefore, strigolactone can be
possibly synthesized in other organs, suchas flowers, fruits, because many flowers andfruits synthesize carotenoid as a componentof pigments in the way that strigolactone is─5─筑波大農林技研 第 5 号 2017

involved in fruit ripening or ﬂower senescencevia complex hormonal responses (Zhu et al.,2010; Su et al., 2015).
Leaves of strigolactone insensitive mutant(Atd14) and signaling defective mutants(max2) kept green even though GR 24
was treated. This result implies inductionof senescence in leaves is stimulated bys trig ola c ton e sig n al in g ra t he r t ha n by
strigolactone content. Also, strigolactone
stimulates leaf senescence by enhancing
signaling pathway mediated by ethylene.
Therefore strigolactone may affect the leaf
senescence in combination with ethylene sothat crosstalk of hormonal signaling should
not be ignored in postharvest physiology.
Hormonal signaling crosstalk between
ethylene and strigolactone would induce
enhanced senescence of leafy vegetables andcut leaves under the dark condition where they
are stored during transportation. On the other
hand, hormonal relationship between ethylene
and strigolactone has not been identified
in fruits and flowers yet. In tomato, forexample, various type of strigolactones, suchas Orobanchol, 7-oxoorobanchol, solanacol,
didehydroorobanchol, has been isolated but
their physiological role in tomato is not clear(Kohlen et al., 2013).

and dedifferentiation in tissue culture andinhibitors of hormones are applied to improve
shelf life of cut flowers, fruits, etc. On theother hand, few reports can be found thatmake an attempt in order to control growthand development of horticultural plants.
Therefore, if the role of strigolactone is known
better, it will be possible to develop application
technique using strigolactones, analogs, orinhibitors in order to improve marketabilityof horticultural plants not only by controllingbranching of axillary buds but also by
inhibiting ethylene biosynthesis.
References
Ada N-L, Kagan S, Ovadia R, Forrer I, Riov J andMichal O-S (2013) Effects of temperature andauxin treatment on fruit set and pigmentation of
Dodonaea‘Dana’. Scientia Horticulturae 160:
172-176.
Akiyama K, Matsuzaki K and Hayashi H (2005)
Plant sesquiterpenes induce hyphal branching inarbuscular mycorrhizal fungi. Nature 435: 824827.
Alder A, Jamil M, Marzorati M, Bruno M, Vermathen
M, Bigler P, Ghisla S, Bouwmeester H, Beyer P
and Al-Babili S (2012) The Path from β-Carotene
to carlactone, a strigolactone-like plant hormone.
Science 335: 1348-1351.
Arite T, Iwata H, Ohshima K, Maekawa M, Nakajima
M, Kojima M, Sakakibara H and Kyozuka J(2007) DWARF10, an RMS1/MAX4/DAD1
ortholog, controls lateral bud outgrowth in rice.
Plant Journal 51: 1019-1029.
A rite T, Umehara M, Ishikawa S, Hanada A,
Maekawa M, Yamaguchi S and Kyozuka J (2009)
d14, a Strigolactone-insensitive mutant of rice,shows an accelerated outgrowth of tillers. Plant
and Cell Physiology 50: 1416-1424.
Beveridge CA and Kyozuka J (2010) New genes in the
strigolactone-related shoot branching pathway.
Current Opinion in Plant Biology 13: 34-39.
B ever i dge CA , R os s JJ and M urf et I C (1 99 4)
Branching mutant rms-2 in Pisum sativum(grafting studies and endogenous indole-3-acetic
acid levels). Plant Physiology 104: 953-959.
Beveridge CA, Symons GM, Turnbull CGN (2000)
Auxin inhibition of decapitation-induced
branching is dependent on graft-transmissible
signals regulated by genes Rms1 and Rms2. Plant
Physiology 123: 689-698.
Booker J, Auldridge M, Wills S, McCarty D, Klee H
and Leyser O (2004) MAX3/CCD7 is a carotenoid
cleavage dioxygenase required for the synthesis of
a novel plant signaling molecule. Current Biology
14: 1232-1238.
Chen SS and Chang JL (1972) Does Gibberellic

Conclusion
Strigolactones have been identified in 1960s
but their physiological role in plants havebeen receiving attention recently (Cook etal., 1966; Umehara et al., 2008). Various
physiological effects such as; crosstalk with
auxin, hyphal growth of mycorrhizal fungi,
response to soil nitrogen and phosphate,branching inhibition of axillary buds inplants, accelerated senescence in plat leaves,
have been reported in may plants including
Arabidopsis, rice, petunia and so on, andstrigolactones are considered a new plants
hormones. Analogs of plant hormones arewidely used in horticultural industry inorder to induce adventitious root formationin cuttings, differentiation, redifferentiation,
─6─

Developmental Control of Horticultural Plants Using Strigolactone to Improve Marketabilityacid stimulate seed germination via amylase
synthesis? Plant physiology 49:441-442.
Cook CE, Whichard LP, Turner B, Wall ME, and Egley
GH (1966) Germination of witchweed (Striga
lutea Lour.): isolation and properties of a potent
stimulant. Science 154: 1189–1190.
Cook CE, Whichard LP, Wall ME, Egley GH, Coggon
P, Luhan PA and McPhail AT (1972) Germination
stimulants. II. The structure of strigol– a potent
seed germination stimulant for witchweed (Striga
lutea Lour.). Journal of the American ChemistrySociety 94: 6198–6199.
de Jong M, Mariani C and Vriezen WH (2009) The
role of auxin and gibberellin in tomato fruit set.
Journal of Experimental Botany 60: 1523-1532.
Divin RS, Moghadam EG and Kiani M (2011) Rooting
response of hardwood cuttings of MM111 apple
clonal rootstock to indolebutyric acid and rooting
media. Asian Journal of Applied Sciences 4: 453458.
Egana FE, Maxwellb BD, Mortensena DA,
Ryana MR and and Smithc RG (2011)
2,4-Dichlorophenoxyacetic acid (2,4-D)–
resistant crops and the potential for evolutionof 2,4-D–resistant weeds. Proceedings of the
National Academy of Science 108: E37.
Gomez-Roldan V, Fermas S, Brewer PB, Puech-Pages
V, Dun EA, Pillot J-P, Letisse F, Matusova R,
Danoun S, Portais J-C, Bouwmeester H, Becard
G, Beveridge CA, Rameau C and Rochange
SF (2008) Strigolactone inhibition of shootbranching. Nature 455: 189-194.
Ha CV, Leyva-González MA, Osakabe Y, Tran UT,
Nishiyama R, Watanabe Y, Tanaka M, Seki M,
Yamaguchi S, Dong NV, Yamaguchi-Shinozaki K,
Shinozaki K, Herrera-Estrella L and Tran L-SP
(2014) Positive regulatory role of strigolactone
in plant responses to drought and salt stress.
Proceedings of the National Academy of Sciences
111: 851-856.
Ishikawa S, Maekawa M, Arite T, Onishi K, Takamure
I and Kyozuka J (2005) Suppression of tiller bud
activity in tillering dwarf mutants of rice. Plant
and Cell Physiology 46: 79-86.
Jiang L, Liu X, Xiong G, Liu H, Chen F, Wang L, Meng
X, Liu G, Yu H, Yuan Y, Yi W, Zhao L, Ma H, He
Y, Wu Z, Melcher K, Qian Q, Xu HE, Wang Y
and Li J (2013) DWARF 53 acts as a repressor of
strigolactone signalling in rice. Nature 504: 401405.
Karakurt H, Aslantas R, Ozkan G and Guleryuz M(2009) Effect of indole-3-butyric acid (IBA), plant
growth promoting rhizobacteria (PGPR) andcarbohydrates on rooting of hardwood cuttingof MM106 apple rootstock. African Journal of
Agricultural Research 4: 60-64.
K o h l e n W, C h a r n i k h o v a T, B o u r s R , L ó p e z Ráez JA and Bouwmeester H (2013) Tomato
strigolactones. Plant Signaling & Behavior 8:
e22785.
K o h l e n W, C h a r n i k h o v a T, L i u Q , B o u r s R ,D o m a g a l s k a M A , B e g u e r i e S , Ve r s t a p p e n
F, Leyser O, Bouwmeester H and Ruyter-

Spira C (2011) Strigolactones are transported
through the xylem and play a key role in shootarchitectural response to phosphate deficiency
in nonarbuscular mycorrhizal host Arabidopsis.
Plant Physiology 155: 974-987.
Martínez GA, Chaves AR and Añón MC (1994) Effectof gibberellic acid on ripening of strawberry fruits(Fragaria annanassa Duch.). Journal of Plant
Growth Regulation 13: 87-91.
Matusova R, Rani K, Verstappen FWA, Franssen
MCR, Beale MH and Bouwmeester HJ (2005)
The strigolactone germination stimulants ofthe plant-parasitic Striga and Orobanche spp.
are derived from the carotenoid pathway. Plant
Physiology 139: 920-934.
Paliyath G, Murr DP, Handa AK and Lurie S (2008)
Postharvest biology and technology of fruits,
vegetables. Wiley-Blackwell Publishing. Iowa.
pp1-7.
Rasmussen A, Beveridge CA and Geelen D (2012a)
Inhibition of strigolactones promotes adventitious
root formation. Plant Signal Behavior 7: 694-697.
Rasmussen A, Mason MG, De Cuyper C, Brewer
PB, Herold S, Agusti J, Geelen D, Greb T,
Goormachtig S, Beeckman T and Beveridge CA(2012b) Strigolactones suppress adventitious
rooting in Arabidopsis and pea. Plant Physiology
158: 1976-1987.
Schulz B and Segobye K (2016) 2,4-D transportand herbicide resistance in weeds. Journal of
Experimental Botany 67: 3177-3179.
Siame BP, Weerasuriya Y, Wood K, Ejeta G and Butler
LG (1993) Isolation of strigol, a germination
stimulant for Striga asiatica, from host plants.
Journal of Agricultural and Food Chemistry 41:
1486–1491.
Sorefan K, Booker J, Haurogné K, Goussot M,
Bainbridge K, Foo E, Chatfield S, Ward S,
Beveridge C, Rameau C and Leyser O (2003)
MAX4 and RMS1 are orthologous dioxygenaselike genes that regulate shoot branching in
Arabidopsis and pea. Genes & Development 17:1469-1474.
Stirnberg P, van de Sande K, Leyser HMO (2002)
MAX1 and MAX2 control shoot lateral branchingin Arabidopsis. Development 129: 1131-1141.
Su L, Diretto G, Purgatto E, Danoun S, Zouine M,
Li Z, Roustan JP, Bouzayen M, Giuliano G and
Chervin C (2015) Carotenoid accumulation during
tomato fruit ripening is modulated by the auxinethylene balance. BMC Plant Biol 15: 114.
Ueda H, Kusaba M (2015) Strigolactone regulatesleaf senescence in concert with ethylene in
Arabidopsis. Plant Physiology 169: 138-147.
Umehara M, Hanada A, Yoshida S, Akiyama K,
Arite T, Takeda-Kamiya N, Magome H, Kamiya
Y, Shirasu K, Yoneyama K, Kyozuka J andYamaguchi S (2008) Inhibition of shoot branchingby new terpenoid plant hormones. Nature 455:
195-200.
Vendrell M (1969) Acceleration and delay of ripening
in banana fruit tissue by gibberellic acid.
Australian Journal of biological Sciences 23:553─7─

筑波大農林技研 第 5 号 2017
559.
Wang KL-C, Li H and Ecker JR (2002) Ethylene
biosynthesis and signaling networks. Plant Cell
14: S131-S151.
Watanabe M, Segawa H, Murakami M, Sagawa S
and Komori S (2008) Effects of plant growth
regulators on fruit set and fruit shape ofparthenocarpic apple fruits. Journal of theJapanese Society for Horticultural Sciences 77:
350-357.
Xie X, Yoneyama K and Yoneyama K (2010) The
Strigolactone story. In VanAlfen NK, Bruening G,
Leach JE, eds, Annual review of phytopathology
48:93-117.
Yamaguchi S and Kamiya Y (2001) Gibberellins andlight-stimulated seed germination. Journal of
Plant Growth Regulation 20: 369-376.
Zhu C, Bai C, Sanahuja G, Yuan D, Farré G, Naqvi
S, Shi L, Capell T and Christou P (2010) The
regulation of carotenoid pigmentation in ﬂowers.
Archives of Biochemistry and Biophysics 504:
132-141.
Z o u J , C h e n Z , Z h a n g S , Z h a n g W, J i a n g G ,Z h a o X , Z h a i W, P a n X a n d Z h u L ( 2 0 0 5 )Characterizations and ﬁne mapping of a mutant
gene for high tillering and dwarf in rice (Oryza
sativa L.). Planta 222: 604-612.
Zou J, Zhang S, Zhang W, Li G, Chen Z, Zhai W, Zhao
X, Pan X, Xie Q and Zhu L (2006) The rice HIGHTILLERING DWARF1 encoding an orthologof Arabidopsis MAX3 is required for negative
regulation of the outgrowth of axillary buds.
Plant Journal 48: 687-698.

─8─