A c t a A g r o n o m i c a H u n g a r i c ' t , L ' o i .' 1 i f 1 - 2 ) , p p / 8 J - . t 9 7 ( 1 9 9 4 - 9 5 )
AUXTNAND CYTOKININBIOASSAYS:A SHORTO\TERVIEW
G. Gvulru andL. E. Hr,szxY
DEPARTMENT OF GENETICS AND PLANT BITEEDING,
LNIVERSITY OF AGRICLTLTURAI. SCIENCES.
H-2103coDol-t-o. HuNcRRv.
(Received:9 February,1994;.accepted:6 October, 1994)
Introduction
Plant Growth Regulators (PGRs). especialll' the auxins and c-tokinins, pla)' a key
role in plant development(Kamalay and Goldberg, 1980; Goldberg, 1988: Gray and
Purohit, l99l). By means of bioassays,a compound can be testedto determine whether it
exerts auxin or c1'tokinin activity. In this paper, auxin and citokinin bioassays are
summarized and briefly discussedfrom Darwin's (1896) assumption (Table 1) to recent
achievements.
In general. there are five groups of classical PGRs: basic cy'tokinins, natural
ethylene, acidic auxins, abscisic acid, and gibberellins @onner 1933; Letham, 1963,
l91I; Stadenand Davey, 1979). There are also severalgroups of compoundswith growth
regulating activity: (l) qynthetic hormone analogues(Mornet et al., 1979; Horgan 198?;
Horgan and Scott, 1987), (2) hormone antagonists(Leopold and Klein, 1952: Christou
and Barton, 1989; Nagy and Tabi, 1982), (3) hormone inhibitors (Varga and Kdves,
1959; K6ves and Sirokman, 1973; Liirsen and Reiser, 1987). (4) hormone herbicides
(Corbett, 1974), (5) antidotesor safeners(l(6mives and Dutka, 1989; Halmann 1990),
Table 1
The history of auxins and cytokrnins
( D o b y 1 9 6 5 :M o o r e 1 9 7 9 ;M a c M i l l a n1 9 8 0 )
I
"Downward moving sap"
2 "Organ-formin g substances
"
3 "L,ateral li ght-inductive factor"
4
5 SyntheticIAA
6 " Transporlable factor"
8
9
l0
lt
t2
l3
Growth promoting substances
a-. b- Auxins**
Heteroauxin
3-[ndoleaceticacid
References
C.)'tokrmrus
Du Hamel(1758)*
"Upward moving sap"
Sachs(l 880)
Darwrn (1896)
Wiesner( 1892)
Ellinger( 1904)*
P a , i l( 1 9 1 9 )
Haberlan&( l92l )
W e n t( 1 9 2 8 )
Kogl et al. (1934a)
Kogl et al. (1934b)
llaagen-Smitet al. (1946)
Miller et al.(19-56)*
S k o o ge t a l . ( 1 9 6 5 )
" Organ-forming substances"
Cell division factor
"Wound horrnone"
Kinin, phytokinin
Cytokinin
* in Moore (1979). ** cyclopentenederivatives,they were provedto be artifacts.
0 2 l E 0 l 6 1 i 9 4 9 5 1 $5 0 0 O l 9 q 5 l f t a d d m i a i K i a d 6 , B u d a p e s t
r86
REVIEWS
Table 2
The eight d(ferent typesof auxins and a*rin metabolites
Natru-al auxrns
Synthetic awjns
Phenyl denvaties
lr,dole
aux[ls
Naphtalene acids Chlorophenoxy
acids
Benzoicacids
Pyridines
Amlno acid precurson
PHI]
'ruYR
Benzo hrazoles
Dthio carbamates
Benzoisotlua(oxa) mles
TRY
PAA PHAA IAA
T-C.A,
ChIAA
N3IAA
IBA
IPRA
NAA
NOX.A
NOAC
2,4-D
2,5-DBA 3,5,6-TPA
2,4-DP 2,3,6-T8A
2'4'5-T 3'6-DMBA
2,4,5-TP
4-CPA
BTR
BIA
BOA
CDDC
Auxin - gluco/ribosidesltides
Auxin - amino acid conjugates
BIA'= 1.2-benzoisothiazole-3-aceticaci4 BTR:beuotf,iazole, BOA:1,2-benzisoxazole-3-acetic aci{ tCA:trzurs-curnamic acid, ChlAA:4-chloro-indoleacetic acid, CDDC:carboxydimethyl-dithiocarbamate, 4-CPA:4chlorophenoxy-aceticacid 2,4-D:2,4-dictrlorophenory-aceticacid, 2,5-DBA:3-amino-2,5-dichlorobenzoicacid"
(CHLORAMBEN), 3,6-DMBA:3,6-dichloro-2-methoxybqzoic acid (DICAN{BA), 2,4-DP:2-(2,4-dichlorophenoxy)-propionic acid (DICHLORPROP), IAA:-indole-3-acetic acid, IBA:indole-3-butlric acid, IPRA:indole3-propionic eci{ NAA==l-napthacetic aci4 N3lAA:azidoindoleacetic aci4 NOAC:methyl-2-naphthory-acetate,
NOXA:2-naphthoxyacetic aci4 PAA:phenylacetic acid, PHAA:para-hydrory-phenylacetic aci4 PHE:phenylalanine, 2,4,5-\':2,4,-5-trichlorophenoryacetic acid 2,3,6-TB{:2,3,5-trichorobeuric
acid, 2,4,5-TP:2(2.4,5-trichlorphenoxy)-propionic acid (FENOPROP), 3,5,6-TPA:4 amino-3,5,6,-trichloropicolinic acid
(PICLORAM) TRY:try?tophan, TYR:tyrosine.
Table 3
The three dffirent types of cytokinins and cytokinin metabolites
Free bases
PtiRINE CYTOKJNINS
Metabo[tes
ISOPENTENYL
ztP.Z
AROIT4ATIC
BA
HETEROCYCLIC
KIN
TIREACYTOKTNINS
PU, DPU, CPPU,TDZ
BENZIMID AZOLE
DEzuVATIITS
BACN
3-,7-,9-N-GLUCO
and
0-GLUCO
and
RIBOSIDES/TIDES RIBOSIDES/TIDES
DBR
8.4:6-benzylaminopurine,
BACN:benz),midaznle
acetonitrile,
CPPU:N-[2-chloro-4-pyridyl]-N"
phenylurea- DtsR:5,6-dichlorobenzimidazole-l-b-D-riboside, DPU:1,3-diphenylurea" 2iP:N-6-[2-isopentenyl]
adenine, KlN:6-fi,trfurylaminopurine, TDZnhidiazuro4 N-phenyl-N'-1,2,3-thiadiazol-5-yl urea. Z:6-[4-hydroxy3-methyl-but-2-enylamino]purine, PU:phenylurea
Acta Agronomica
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REVIT]WS
187
and (6) plant alkaloids with hormone t)?e activiry-@uta anclh^alinski. lg8tt)" In addrtion
(7) a number of miscellaneousgpes of non-classicalPGRs are to bc found: brassinosteroids, elicitors, oligosaccharides,
glycoproteinsand polianrines{Yopp et ai.- l{)81: Corbett, 1974; Bruinsma, 1980).Besides,(8) hormonemetabolitesfi.aloue and Fcthe. 1982)
also have biologrcal activity. The loose connectionsbehveenthe molccular structure and
hormone activities underline the importance of bioassaysin classifr,rnsancl determinrng
the newbioactive [email protected]; Zhao,et al. 1992;Kanbc et ai.. 1993).
Intensive researchhas been focusedon finding netv grorl'th-regulatingcompounds.
such as modified auxins like azido-auxins [Lee et al., 19tt4), modified cytokinins iike
thidiazuron (Thomas and Katterman, 1986; Visser et al." 1992)"CPPU ([{almann. 199(,)
and new types of PGRs like S-(cerboxymethyl)-dimethyldrthrcarbamare
fcrr agronomical
purposes (Kerk et al., 1957), benzimidazole @erson et al., 1957). picloram @eyl and
Sharma, 1983), dicamba (Conger et al., 1983), benzothrazole(BIA) and bennznxal Lole
@ranca et al., 1990). No general theory exists which enablesus to predict the hormone
activity from the molecular structure. Wlule the molecular structure requirements for
PGR-activity are evident in the casesof abscisic acid. gibberelins and ethvlene, in the
case of auxins, except for certain types (Porter and Thimann, 1965), and in the case of
qtokinins, it is still an open question, becauseof their various molecular structures
(Tables 2. 3). Auxins comprise over ten types of compounds with different molecular
structures,but with similar biological activity (Table 2). Cvtokinins include three main
groups of compounds(Table 3).
The discovery of the correlation between the electronic and steric properties of
PGR molecules led to the formulation of QSAR (Quantitative Structure-Activit-v
Relationship). In this way, the new PGR molecules can be designed by computer
progralnmes (Halmann, 1990). Obviously, bioassaysare necessaryto veri$ bioactivity.
Bioassays
Different plants of monocots and dicots are used for bioassays(Tables 4, 5). The
value of any bioassay ultimately dependson the specificity and selectiviqvof the observed
biological responseson test objects(Nissen,1985; lgSS).
When the responses are compared. bioassays for auxins and cytokinins can be
classified as: (l) Cell elongation, (2) cell division, (3) cell function, and (4) cell differentiation (Tables 4, 5).
From the observationof Darwin (1896) on Phalaris coleoptile, severalassayswere
developed on the coleoptile of different monocot plant species"especiall-_v
of Avena
l9l3: Pa6l, l9l9: Stark, l92l). In Avena coleoptiletests,both cun'ature
@oysen-Jensen,
and straight growth were used (Bentlev, 1950; Kefford, et al., L962). This technique rvas
extendedto other plant species,e.g. ,Ageratum (Bottelier, 195,1).cuculnber (Katsumr el
al., 1965)and sunflower@rauner, i966).
Severalother effectsof auxins (Table 4) were also usedfor bioassays,e.g. initiation
of tomato parthenocarpy (Zimmennann and Hitchcock, 1940). Lotus hairy root test
(Shen et al., 1988) and in the de novo root initiation assavin vitra (Glulai et al.. lggL..
1993).
Acla
Agronomica
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REVIEWS
Table 4
Bioassyasfor auxins
t. Cell elongation assays
Atxin induced coleopttle curvature assays
'fest
(Boysen-Jensenl9l3; Pa.il l9l9; Stark 1921)
1. Avena Coleoptile Section
2. Avena Coleoptile Cun'ature Test (Went 1928; Went and Thimann 1937)
3. Avena DeseededColeoptile Test (Skoog 1937)
4. Avena GeocurvatureTest (Kaldewey et al. 1969)
-5. Avena Coleoptile Split Test (Rayle 1973)
5. Phalans EtiolatedCoieoptile PhototropicTest (Darwin 1896)
Auxin tnduced cc.leoptilestraight grow,thassays
7 Atena Coleoptile StraightCnowth Test (Bonner 1933,1949; Schneider1938; Bentley 1950; Sirois 1966)
8. Avena PeeledColeoptile Test (Bonner 1934; Nitsch and Nitsch 1956)
9 Avena Micro Slraight Growth Test (fuetserna 1949)
10. Corn ColeoptileTest (Bridges and Wilkins 1973)
ll. Rice Coleoptile Test (Yamada 1954)
12. Sorghum Coleptile Test (Ramana et al., l97l)
13. I'l/heatColeoptile Test Q.Jitschand Nitsch 1956; Hancocket al. 1964)
Auxin induced dilferent curvature assays
14. Ageratum PetioleCun-ing Test (Bottelier 1954)
15. Bean lnternade Curving Tests (Meudt and Bannett 1978; Strand and Kaminek 1985)
16. Citnts Petal Curving Test (Goldschmidtand Monselise1966; Goldschmidt1968)
17. Pea Stem Split Curving Test (Went 19341vanOverbeekand Went 1937)
18. Pea Quarlered Stem Split Curving Test (Thimann and Schneider 1939 Ockerse and Galston 1967)
19. Tomato Petiole Curving Test (Kazerni and Kefford 1974)
Auxin induced d{ferent elongation assays
20. Avena First lntemode Test (Nitsch and Nitsch 1956; Crosbyet al. 1961)
21. Cabbage Hypocotyl ElongationTest (Andersenand Muir 1966)
22. Cucumber Hypocotyl ElongationTest (Katsumi et al., 1965; Sakuraiet al. 1974)
23. Mung Bean Hypocotyl ElongationTest (Goldberg 1980)
24. Prne Hlpocotyl ElongationTest (Wodzicki and Wodzicki 1973)
25. Pea Epicotvl Elongation Test (Christiansenand Thimann 1950)
26. LlunjlowerHypocotyl ElongationTest (Brauner 1966)
II. Cell division assays
27. Soybean CotyledonCallus Test (Miller 1963)
28. Tobacco Callus Test (Murashigeand Skoog 1962)
I II. Cell function assays
Bean t€af Inhibition Test (Brown and Weintraub 1950)
Cichorrum Root Disk Water Uptake Test (Rutherford et al. 1966)
Coleus kaf Abscissiontest (Luckwill 1956.1
Tomato ParthenocarpyInduction Test (Zimmermann rul Hitchcock 1940)
IV. Cell d{ferentiation assays
Auxin controlled rooting assays
33. Avenq Root Inhibition Test (Bonner and Koepfli 1939)
34. Bean Root Test (Luckwill 1956)
35. CornRoot Test (Swanson1946)
36. Cucumber Cotyledon Rooting Assay (Zhao et al. 1992)
37. Cress Root SectionTest (Moewus 1949)
38. Lettuce Rool Test (Ready and Grant 1948)
39. Lotus Hairy Roots Test (Shar et al., 1988)
40. Pea Root Test (Aberg 1950t Audus and Thresh 1953)
Auxin induced morphogenesis assays
41. Tobacco Root RegenerationTest (Skoog and Miller 1956; Murashigeand Skoog 1962)
42. Tohacco Test With Auxin Selectivitv(Gwlai et al. 1992. 1995).
29.
30.
31.
32.
Acla Agronomica Hungarica 43, 1994-95
189
REVIEWS
Table 5
B i o a ssay sfor cytoki ni n s
I. Cell elongation assays
Cytokinin induced coleoptile elongation assays
1.
Avena Coleoptile ElongationTest (Shrank 1957; 1958)
2.
Weat Coleoptile ElongationTest (Rothwell and Wright 1967)
Cytokinin i nduced expansion assays
3.
4.
5.
6.
7.
8.
9.
10.
Bean l naf Disc ExpansionTest (Miller 1956)
Cucumber Cotyledon Expansion Test (Green and Muir 19781,Zhao et al. 1992)
Lemna Frond Growth Test (Hillrnan 1957; tneffler and VanOverbeek 1964)
Lettuce Cotyledon Expansion Test (Ikuma and Thimann 1963)
Radish Cotyledon Enlargernent Test (Kuraishi 1959)
Radish CotyledonExpansionTest (L,etham1971)
Spirodel Frond Growth Test (ktham 1967a)
Xanthium CotyledonExpansionTest (Esashiand tropold 1969)
II. Cell division assays
Cytohnin induced callus prolferation assays
I l. Bean PodParanchyma Test (Wehnelt 1927;English and Bonner 1937)
12. Carrot Callus Test (Caplin and Steward 1949; Stewardand Shantz1955)
13. Carrot Root Callus Test (I-etham 1963, 1967a)
14. Soybean CotyledonCallus Test (Miller 1960, 1963)
15. SoybeanHypocotylSectionTest(ManosandGoldthwaitel9T6;Newtonetal.1980)
16. Tobacco SternSlab Test (Skoog and Miller 1956)
'l'obacco
17.
Pith Callus Test (Murashigeand Skoog 1962)
t' c ell fun cti o n a ssays
c'y tokrn i n i nduc ed p igme nt fo rm ati or r, or,!t
I 8. Amaranthus retroflexus Betacyanin Test (Bamberger and Mayer I 960, K6hlsr and Conrad 1966)
-lhomas
19. Amaranthus caudatusBetacyaninTest (Bigot 19681Biddington and
1973; Redaand Rasmussen
1 e7s )
20. Bean Etiolated kaf-Disc Test (Millewr 1963)
21. Cucumber Chlorophyll FormationTest (Fletcherand McCulla$t l97l; Fletcheret al. 1982)
22. Soybean DeorysoflavoneTest (Miller 1969)
Cytohnin included leaf senescencedelaying assays
23. Avena kaf Chlorophyll Retantion Test (Thimann and Sach 1966; Varga and Bruinsma 1973)
24. Barley lraf Chlorophyll Retention Test (Kende 1964, 1965;'lrtham 1967b; Engelbrecht l97l)
25" Bean lruaf Chlorophyll Retsntion Test (Goldthwaite and I"-aetsch1967)
26. Radish kaf Chlrophyll Retention Test (Kefford et al. 1968)
27. Rumex Leaf Chlorophyll Retention Test (Goldthwaite 1972; Manos and Goldthwaite 1975)
28. Xanthium kaf Chlorophyll RetentionTest (Richmondand tang I957; Osbomeand McCalla 1961)
29. Wheat t eaf Chlorophyll Retention Test (Person etal. 1957)
Cytokinin stimulated transpiration assays
30.
Avena t eaf Transpiration Test (Luke and Freernan 1967;Tetley and Thimann 1974)
M. Cell d{ferentiation qssays
Cytokinin induced morphogenesisassays
31. Barley Root Inhibiton Test (VanOnckelen and Verbeek 1972)
32. Impatiens I-ateral Shoot lnitiation Test (Bozsik 1983)
33. Funaria Moss Bud Formation Test (Szweykowska 1962;Halnand Bopp 1968; Brandes and Kende 1968)
34. Leftuce Seed Germination Test (Skinner and Shive 1956; Skirurer et al. 1957).
35. Pea Etiolated Stern Inhibiton Test (Sommer 1961)
36. Pea l.a;tqal Bud Induction Test (Thimann and Sachs1966)
37. Pohlia Moss Bud FormationTest (Mitra and Allsopp 1959)
38. Tobacco Shoot ReganerationTest (Skoog and Miller 1956; Murashigeand Skoog 1962)
39. Tobacco Test With Cytokinin Selectivit-v(Glulai et al. 1992, 1995)
40. Tortella Moss Bud Formation Test (Gorton et al. 1957)
Acta Agronomica
Hungarica
43, 1994-95
I9 O
R EV IEW S
The first c,vtokinin-dependentcell division testswere basedon tobaccopith (Skoog
and Tsui, 1951; Patauet al., 1957'"Skoogand Miller, 1956 Das et al., I958; Murashige
and Skoog, L962). The s.vstem was widened to carrot (Steward and Shantz, 1955;
Letham, 1963; 1966), and to soybean (Miller, 1963 Manos and Goldthwaite, 1976;
Newton et al., 1980). The cy'tokinin-initiated leaf expansion-effectwas used first in a
bean leaf test (Miller 1956); later, other dicot plants were found to be responsive,e.g.
radislr cotyledon (Kuraishi 1959; Letham I97I), Spirodela frond (Letham, 1967b) and
cucumbercotyledon (Greend and Muir, 1978).
A special part of the cytokinin bioassayswere based on the initiation of pigment
formations like betacyanin in Amaranthus @amberger and Mayer, 1960; Elliot, 1979),
chlorophyll in cucumber (Fletcher and McCullagh, I97I) and deoxyisoflavone in
soybean(lr4iller, I 969).
From the obscn'ation of the c1'tokirrin-inducable chlorophyll retention effect
(Richmond and Lang. 1975; Person et al", 1957 Reilly, 1986), several assays were
developedto barley (Kende, 1964, 1965).Avena (Thimann and Sachs, 1966) and radish
(Keffiord et al., 1968). As a result of the key role of cytokinins on plant morphogenesis,
different morphoregulatory assayswere developed for moss bud formation (Skinner and
Shieve. 1956: Gorton et al.. 1957'.Hahn and Bopp, 1968), for tobaccoplant regeneration
(Skoog and Miller, 1956) and for selectivetobacco shoot initiation (Gyulat et al., 1992,
1995).
Limitations
Different bioassays have various limitations and disadvantages. The seed
germination assays(Skinner and Shive, 1956',Skinner et al., 195'7)can be influenced not
only by qtokinins but by many other factors such as red light, gibberellins and thiourea
(Miller, 1963).
The kinetin-retarded senescenceassaysare responsivenot only to rytokinins but
also to benzimidazole or large amount of sugar, as was observed in the tests of Xanthium
(Osborn and McCalla" 196l) and Rumex (Goldthwrite 1972). Therefore, their uses are
very limited (Varga and Bruinsma, 1973).
The leaf expansion rytokinin tests (Miller, 1956; Kuraishi, 1959) are limited by
the side effect of cobalt ions, red light, and the cross effect of gibberellins (Miller, 1963).
In the q'tokinin tests of tobacco and carrot, cross effects of IAA and myo-inositol were
found (Miller. 1963; Letham, 1967a) and there was a tendency of soybeancotyledons to
becomeindependentof cJtokinins (Newton et al., 1980).
Assays based on callus or suspensioncultures involved limitations of habituation
(Gautheret,1934; 1938, 1960:K6ves and Szabo,1987;Christou, 1988).
Coleoptile test were considered to be highly specific for auxins, until the
observation of Shrank (1957, 1958) and Rothwell and Wright (1967) demonstratedthat
there was not only auxin but cytokinin activity in coleoptile tests (Table 5).
Leaf expansion and pigment production assaysfor qrtokinins showed difficulties of
standardizationand hormone specificity (Letham, I9'71',Lad6 et al., 1975).
Acta Agronomica Hungarica 43, 1994-95
191
REVIEWS
Betacyanin accumulation in Amaranthus assays(Table 5) exhibited cross activity,
being enhanced not only by cy'tokinins but also by fusicoccin, KCl, NaNO3, KNO. and
phosphateions @lliot, 1979).
In bioassaysbased on leaf senescenceretardation, the protein synthesisinhibitors
could imitate the rytokinin-effect (Tefley and Thimann, 1974).
Facilities of tissue culture techniques made it feasible to increase hormone
specificity (Linsmayer and Skoog, 1965; Gamborg et al., 1968, Malone and Dix. 1986;
Schenk and Hildebrandt, 1972), eliminating the s-vnerging effects of PGRs
@hattacharayaet al., 1978; Hasensteinand Evans, 1986). In Xanthium cotyledon tests
the gibberellin cross activity was eliminated successfi.rllyby the application of mannitol
in the test media @sashi and Leopold, 1969).
A new selective bioassayfor auxins and cytokinins was reported recently (Gyular et
al., 1993, 1995). It was based on the ability of cytokinins to induce adventitious shoots
and the auxins to induce adventitious roots applied alone on tobacco leaf discs cultures,
giving a so-called "all-or-none" (Hahn and Bopp, 1968) reaction. T"he new bioassay
eliminated the synerging effects of auxins and cytokinins. Abscisic acids and gibberellins
did not show activity. The method was patented in Hungary (Gyulai et al., 1992).
The new physicochemical (PHLC, GC-MS etc.) and immuno assays(ELISAI RIA
etc.) opened a new dimension in PGR science(Letham, 1.966',Ernst et al., 1983; Sutter
and Cohen, 1992). Nevertheless,bioassaytestsare always neededto prove bioactivity.
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Brauner, L. (1966): Versuche zur Analyse der geotropischen Perzeption. V. Ober den Einfluss des
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