Genes of Auxin Biosynthesis and Auxin-regulated Genes

8/13/2019 Genes of Auxin Biosynthesis and Auxin-regulated Genes

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The article was published in Journal: Biopolymers and cell, 2005, V.21, N2,

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1Victoriya +. Tsy8an9oa, 1arysa +. (al9ina, 2udmila %. usaten9o,2;onstantyn . 3ytni9

1Institute of Bioorganic Chemistry and Petrochemistry, National Academy of Sciences of the

Ukraine

Murmanska Street, Kie !"!#$, Ukraine

"Kholodny Institute of Botany, National Academy of Sciences of the Ukraine

%ereshenkiska Street, Kie !1!!$, Ukraine

In the review a spectrum of enzymes genes determining different ways of indole-3-acetic acid

(IAA) biosynthesis identified at Arabidopsis is given: the !"# gene of antranilat

phosphoribosyltransferase#$ !"3 gene of tryptophan synthase and family %I genes of

nitrilase$ catalysing tryptophan-independent way of IAA biosynthesis from precursor indole-3-

acetonitrile& '"*+ and '",3+# genes (members of family cytochromes "./ genes)$

controlling IAA biosynthesis from tryptophan& the enzymes genes catalysing of the IAA

biosynthesis from indole-3-butyric acid: the "0A# and "10# genes of pero2isomal membrane

proteins - the A+'-A"as family members$ the "10. and "10 genes of cytoplasmic protein

receptors$ genes of pero2isomal matri2 proteins-enzymes (ac23 gene of acyl-'oA o2idase$

aim# gene of multifunctional protein and ped# gene of thiolase)& enzymes genes catalysing

synthesis of IAA-conugates and their hydrolysis - the genes of IA45c syntase$ IAInos

transferase$ serin carbo2ypeptidase-li6e IAInos acyltransferase and IA!3 gene of IAA-Ala

hydrolase7 he nomenclature and classification of the au2in-regulated genes responsible for

the cell division are presented: cyclin genes and cyclin-dependent protein 6inase genes$ as well

as genes of numerous family of mitogen-activated protein 6inases7 he au2in-induced genes of

enzymes participating in biosynthesis and hydrolysis of polysaccharides components of plant

cell walls in the period of their growth by e2tension are considered in detail: the 1I gene ofendo-#$3:#$--8-glucanase and 109II gene of e2o--8-glucanase$ numerous families 01

genes of 2yloglucan endotransglycosylases$ e10" genes of e2pansins$ At;-fucosyltransferases and glycosyltransferases$ '?5 genes

of 2yloglucan glucan synthases and -#$-mannan synthases$ @

galactosyltransferases as well as the At0# gene and homologous At4=- genes 2yloglucan

2ylosyltransferases at Arabidopsis& the 0?# gene of 2ylan synthase at rice$ and also 4?# gene

of glucan synthase at corn7 A possible role of cell wall protein -e2tensin (encoded by the au2in-

regulated !4" gene) in the plants defence from pathogens and unfavourable factors ofe2ternal environment is discussed 7

Key words: genes of indole-3-acetic acid (IAA) biosynthesis& au2in-regulated genes

responsible for plant cell division and e2tension&

1


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%ntroduction. Information concerning the hormonal regulatory system in

'lants (as first 'u)lished more than 1!! years ago& In the first 'u)lications the

suggestions a)out the e*istence of endogenous mechanisms controlling 'lant

gro(th and deelo'ment, )ased only on em'iric, fragmentary data regardingfunctional correlatie interrelations )et(een arious 'lant 'arts, (ere made +1&

%hose early 'u)lications 'resented facts 'roing these interrelations, a)out many

'hysiological 'rocesses such as stimulation of the lateral )uds gro(th during

se'aration of the main a'ical )ud +", increasing seeds germination )ecause of

their se'aration from fruits +-, termination of the coleo'tiles tro'ism (hen their

a'ical .ones are deca'itated +$, termination of starch degradation after germs

remoal from grains +/, iolation of )uds gro(th occurring (ith a 'artial

defoliation +0, retardation of leaes and stems senescence (hen 'lant

re'roductie organs are remoed +& 2enceforth, these o)serations ena)led t3

su''ose that interactions )et(een se'arate 'lant organs may )e mediated )ychemical com'ounds called later )y 'hytohormones&

A significant 'rogress in this field has )een made as a result of the organic

chemistry deelo'ment& %he first au*in 4 indole5-5acetic acid 6IAA7, ado'ted )y

a gro(th hormone, (as isolated and identified )y an outstanding Ukrainian

)iologist, the founder of the 'hytohormone theory 5 Kholodny in 1#"8 +8, #&

Almost simultaneously )ut inde'endently, similar e*'erimental studies (ere

conducted and theoretically grounded )y a 9utch 'hysiologist :ent& As a result,

the general hormonal theory of tro'ism, kno(n from the scientific literature and

manuals as the Kholodny5:ent theory, (as formulated& A classical conce'tionof the 'hytohormone as a chemical messenger that is synthesi.ed in one 'art of

the 'lant, transferred into the other and affects arious 'hysiological 'rocesses,

(as first e*'ounded )y :ent and %imann in 1#- +1!; later, in 1#0!, %imann

terminologically su''lemented the conce'tion of 'hytohormones as organic

su)stances acting in small amounts +11&

It should )e noted that since the time (hen 9ar(in discoered the

'henomenon of tro'ism and for the follo(ing decades, IAA has )een considered

the main regulatory hormone at all ste's of the 'lant ontogenesis, (hile the

e*'erimental data giing eidence of the e*istence of other 'hysiologically actiecom'ounds, regulating the cell 'roliferation and differentiation, (ere de)ata)le

u' to /!th years of the last century& In the su)se


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'lant gro(th su)stances, (hich may )e eleated to hormone status=

olygosaccharides, >asmonates, salicilates, 'olyamines +1- 5 1/, turgorines,

stiroles +10; hormone5like su)stances of a dou)le au*in5cytokinin action 6they

include )rassinosteroids +1/ and fusicoccine found in flo(er 'lants +1; in>ury

hormones 6necrohormone, traumatine7 4 com'ounds formed on in>ured and

(ounded surfaces and facilitating their recoery 6it is )elieed that in addition t3the (ell5kno(n )ut insufficiently s'read traumatic acid the (ound hormone

function is also 'erformed )y au*in and cytokinins7 +18; flo(ering hormones 5

ernaline and florigene&

%he florigene e*istence (as first suggested )y an outstanding ?ussian

)iologist Chailakhian in 1#- +18& Su)se


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'rocesses has )een inestigated in details= cell diision and elongation 6for

e*am'le, there (ere identified 'hytohormone5regulated genes and their 'roducts

4 en.ymes inoled in hydrolysis and )iosynthesis cell (all com'onents during

cell elongation as (ell as in the regulation of the 'lant cell mitotic cycle7& %here

(as disclosed the role of 'hytohormones in 'hotomor'hogenetic 'rocesses as

(ell as in 'lant resistance t3 enironmental unfaoura)le factors and 'athogens&Promising fields of 'ractical a''lications of 'hytohormonology achieements are

as follo(s=

17 creation of ne( 'lant mutant lines (ith a distur)ed or reduced

)iosynthesis of some 'hytohormone or, ice ersa, o)taining transgenic 'lants

hy'er5e*'ressing some 'hytohormone t3 im'art agriculturally useful 'ro'erties t3

'lants 6these techni


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- stimulation the cell elongation in coleo'tiles and shoots 6au*in

concentrations normal for stimulation of cell gro(th are 1! 584 1!50 moll7& At the

molecular leel the cell elongation stimulation is associated (ith an increased

transfer of 'rotein5en.ymes from the cyto'lasm into the cell (all resulting in its

)reaking u' and elongation under the turgor influence;

-actiation the cam)ium cell diision 6there is a direct interrelation )et(een

the )ud o'ening and cam)ium actiity in trees= as a result of )ud remoal a

secondary (idth gro(th is a)sent7& In grassy dicotyledons the secondary (idth

gro(th is induced )y au*in transferred t3 the stem from young leaes and a'ical

)uds;

- a'ical dominance= au*ins, trans'orted do(n(ards from the a'ical )ud,

inhi)it the lateral )ud gro(th; au*ins, synthesi.ed in the root a'e*, stimulate the

lateral )ud formation;

- 'romotion the fruit setting 6the most actie au*in 'roducers in a

deelo'ing fruit are seed5)uds7;- 'reention the 'rocess of leaf se'aration= au*ins, transferred from the

'etiole, inhi)it the actiity of 'olysacharide hydrolases 6endo'olygalacturonase

and cellulase 5 51,$5glucanase7, synthesi.ed in a se'arating tissue of the 'etiole

)ase at a high ethylene concentration; o''osite effect of induction of this 'rocess

is o)sered (hen au*in is trans'orted from the a'ical shoot& A se'arating tissue

res'onse de'ends on the gradient of au*in concentration )et(een 'etiole tissues

and a'ical shoot&(enes o< au=in biosynthesis. Itis kno(n that s'ecific tumors cro(n gallsD

as (ell as )earded rootsD in 'lant tissues, infected )y Agrobacteriumtumefaciens and A7 rhyzogenes, containing i and !i 'lasmides, res'ectiely,

result from the e*'ression of 'lasmide genes encoding IAA )iosynthesis +"=

tms genes ofA7 tumefaciensand au2 genes ofA7 rhizogenes& %hese genes encode

en.ymes try'to'han5"5monoo*igenase and indole5-5acetamidehydrolase,

normally a)sent in higher 'lants and cataly.ing IAA )iosynthesis from

try'to'han +"/& In 'lant cells, transformed )yA7 tumefaciens, IAA is synthesi.ed

s'ecifically= firstly try'to'han conerts into indole5-5acetamide (hich is then

used t3 IAA synthesis& :hen 'lants are transformed 6for e*am'le, to)acco,

tomato, au)ergine7 )y other )acteria 6"seudomonas syringae or Antirrhinummaus 9ef2#7, there is o)sered a s'ecific hy'er5e*'ression of iaa@ or iaa

genes corres'ondingly manifesting itself in some intensification of hy'ocotyl

elongation and in 'romoted deelo'ment of fruits from non5'ollinated seed5)uds

as a result of su''ly t3 them of a great amount of IAA +"8& %he gene iaa@ (as

found t3 encode try'to'hanmonoo*igenase 4 en.yme inoled in the try'to'han

transformation in indole5-5acetamide, )iotransforming later in 'lants in IAA&

Arabidopsis and to)acco shoots, transformed )y the iaa5 gene of )acteria

"seudomonas savastonoi 6encoding the IAA en.yme lysine synthase7, (ere

sho(n t3 hae short hy'ocotyls )ecause of some reduction in free IAA ugates (ith lysine +"#&

/


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%here are arious (ays of the natural IAA )iosynthesis ty'ical for 'lants;

some of them inole try'to'han as a deriatie from indole or early 'recursors&

A genetic analysis of the IAA )iosynthesis in Arabidopsisand corn has sho(n

that IAA is an intermediate com'ound in the antranilate5try'to'han 'ath(ay of

)iosynthesis +-!, -1& It is suggested that there are different 'ath(ays of IAA

)iosynthesis from try'to'han ia indole5-5'yruic acid, indole5-5)utyric acid,try'tamine indole5-5acetonitrile +"/, -1, -"&

Studies of try'to'han5au*otro'hic 'lant mutants hae sho(n that IAA

)iosynthesis may also occur along a try'to'han5inde'endent (ay inoling

'recursor IAN, (hich (as firstly identified in 'ruciferae family& It (as, for

e*am'le, esta)lished that mutant trp3-#and trp=-#try'to'han5au*otro'hic 'lants

of Arabidopsisand corn accumulated high leels of indole5-5glycero'hos'hate

and IAN (hile the free IAA leel remained normal +-"& @n the )asis of this data

as (ell as results of studies on the cell5free system from an immature corn

endos'erm, e*hi)iting a transformation of radio5actiely la)elled indole in IAA, atry'to'han5inde'endent (ay of IAA )iosynthesis from indole or indole5-5

glycero'hos'hate has )een suggested +"/, -1 5 --&

At 'resent, genes of )asic en.ymes, inoled in IAA )iosynthesis in

try'to'han5au*otro'hes, hae )een determined= !"# gene of antranilate5

'hos'hori)osyltransferase, conerting antranilate in /5'hos'hori)osylantranilate,

and !"3 gene of try'to'han syntase E, cataly.ing transformation of indole5-7

glycero'hos'hate in indole +-!, $!& IAN, formed at the su)se


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e*hi)it an increased a'ical dominance and hy'er5elongation of hy'ocotyls at the

'lants 6)ecause the high leels of free IAA concentrations and lo( ones of

con>ugated IAA are accumulated7 +$!& %he 'henoty'e, o''osite t3 rnt#-#

mutants, is o)sered in Arabidopsis 'lants (ith an ecto'ic hy'er5e*'ression of

'",3+# k9NA under 3.?5'romoter (hich are 'henoty'ically similar t3 a2r#

mutant 'lants (ith a reduced a'ical dominance, short hy'ocotyls and decreasedfertility that results from a reduced sensitiity of those 'lants t3 au*in +$1&

In normal conditions the '",3+#gene e*'ression (as esta)lished t3 occur

in roots, leaes, stems, flo(ers and stamens +$" (hile in stress conditions

6caused )y dehydration or damages7 and under influence of au*in a '",3+#

gene e*'ression intensification is o)sered mainly in roots +$-& %he '",3+#

gene 'romoter (as sho(n t3 hae four au*in5res'onse cis5elements that confirms

the au*in regulation of '",3+#gene e*'ression +$$&

?ecently, there (as 'u)lished information that in the regulation of IAA

cata)olism a considera)le role is 'layed )y 'ero*isomes 5 im'ortant organellesinoled in a li'id meta)olism, nitrogen fi*ation, 'hotores'iration and o*ygen

decom'osition; they also sere as sites for the late stages of >asmonic acid and

IAA )iosynthesis in 'lants +$/, $0& Pero*isomes are actie throughout the 'lant

ontogenesis= from em)ryogenesis, seed germination and flo(er deelo'ment till

senescence 'hase& Gast studies hae ena)led t3 demonstrate a leading role of

'ero*isomes in 'hotomor'hogenesis and their 'rotectie functions in a)iotic

stress conditions; 'ero*isomal en.ymes take 'art in the formation and

decom'osition of reactie o*ygen com'ounds 6?@S7 5 signal molecules

regulating the nuclear gene e*'ression and 'roducing a damaging effect on the

cell in case of their a)undant formation +$&

Shoots and aging 'lant tissues (ere found t3 contain s'eciali.ed 'ero*isomes 4

glyo*ysomes synthesi.ing more than "! en.ymes for H5o*idation of fatty acids

6FA7 and amino acids& Acetyl5coen.yme A 6acetyl5CoA7, formed as a result of

en.ymic s'litting of FA, is conerted into succinate and trans'orted t3

mitochondria (here it enters the cycle of tricar)o*ylic acid reactions (ith

su)se


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Mutant 'lants e*hi)it some retardation of germination and su)se


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Mutant 'lants, resistant t3 an inhi)iting effect of IBA or its synthetic analogue

",$59B 6meta)oli.ed during H5o*idation in 'ero*isomes of au*in ",$597, are

kno(n t3 )e defectie in the )iosynthesis of arious PMP such, as acyl5CoA

o*idase 6mutation ac237 +/, multi5functional 'rotein 6mutation aim#7 +/8 and

thiolase 6mutationped#7 +/#& It has also )een found that since the H5o*idation of

FA in 'lants occurs only in 'ero*isomes 6in animals 4 in 'ero*isomes andmitochondria7, mutations of 'ero*isomal mem)rane 'roteins 6through (hich FA

are trans'orted into 'ero*isomes7 as (ell as 'roteins5rece'tors 6trans'orting PMP

into 'ero*isomes7 may distur) the H5o*idation of FA and IBA& For e*am'le,

pe2. mutants, defectie in the synthesis of cyto'lasmatic rece'tor PQ/, are

insensitie t3 IBA +/1 (hile ",$59B5resistantped= mutants are defectie in the

synthesis of 'ero*isomal mem)rane 'rotein PQ1$ +0!&

@n the )asis of the studies on IBA5 and ",$59B5resistant mutations there (as

suggested a hy'othetic model of the 'ero*isomal trasformation IBA into IAA,

occurring similarly the FA H5o*idation 6Fig&"7& According t3 this diagram thePA1 'rotein trans'orts acyl5CoA esters of FA into the 'ero*isome (here they,

during H5o*idation, are transformed into acetyl5CoA (hich is, as a result of

fermentatie disintegration, meta)oli.ed into succinate ia the glyo*ylate cycle

+$0, $8& Since p2a# mutants are resistant t3 IBA and its analogue ",$59B, it

might )e that the PA1 'rotein e*'orts acyl5CoA esters of IBA and ",$59B into

'ero*isomes t3 )e o*idi.ed into IAA5CoA and ",$595CoA, res'ectiely +/1&

%hese com'ounds are hydroly.ed and in the form of IAA and ",$59 are e*'orted

from 'ero*isomes causing su)seugates

formation as (ell as their disintegration indicates that a com'le* homeostatic

mechanism controlling this 'hytohormone content e*ists& IAA is 'resent in 'lant

tissues mainly as con>ugates (ith amino acids, 'e'tides or car)ohydrates (hich

can )e hydroly.ed into free IAA moreoer, egetatie organs are characteri.ed )y

a considera)ly lo(er amount of free IAA, (hile in generatie organs 6es'eciallyin mature seeds7 con>ugated forms, )eing )iologically inactie resere forms of

IAA and maintaining the hormone homeostasis 7, are 'redominant +"/, "0, 01&

%he analysis of IAA cata)olites in corn grains, for e*am'le, ena)led t3 identify

mostly con>ugates containing an ester )inding= IAA5glucose, IAA5myo5inositol,

IAA5myo5inosytol5glycosides as (ell as cellulosic glucan con>ugates 5 all

together amounting t3 # 5 ## O of the total 'ool of IAA in the seed endos'erm

+0"& QsterifiedIAA is also a 'redominant con>ugate in rice grains 6containing 0"

4 ! O of ester5)ound con>ugates7 +0-, in a li


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many 'lants= for e*am'le, at the early stages of the )ean seed deelo'ment, the

ma*imum leel of esterified IAA is -/ O, that of free IAA 4 around $! O of the

total IAA 'ool +0/; then during the seed ri'ening, the leels of esterified and free

IAA reduce do(n t3 1- O and 0 O, res'ectiely, from total IAA 'ool; at the

com'lete maturation stage the con>ugates amide5)ound (ith 'oli'e'tides and

'roteins 68! O of the total IAA 'ool7, 'reail +00& Unlike )eans, in mature soyseeds IAA con>ugates (ith amino acids 6as'artates and glutamates7, haing a

lo(er m&m&, are 'redominant +0&

Ras chromatogra'hy and mass5s'ectrometry techniugates and cata)olites during the (hole egetatie gro(th of many 'lants

includingArabidopsis, corn, to)acco, tomato, )ean, soy, rice, oat, chestnut, 'ine,

'o'lar +"0, 08& For e*am'le, the analysis of arious Arabidospsis tissues

inoling the a''lication of la)elled standards= cata)olites of "5o*indole5-5acetic

acid and also IAA con>ugates (ith amino acids 6as'artates, glutamates,

com'le*es of IAA (ith alanine and leucine7, hae sho(n that elongating leaesand roots e*hi)it relatiely high concentrations of free IAA and the highest

concentrations of IAA5as'artate, IAA5glutamate and "5o*indole5-5acetic acid

(hile IAA con>ugates (ith leucine are concentrated in roots and IAA con>ugates

(ith alanine 4 in aerotissues +08& A 'ercentage of the leel of esterified IAA

con>ugates inArabidopsis makes u' 8 4 1! O and leel of IAA con>ugates (ith

amino acids 4 a)out " 4 - O relatie t3 the total IAA 'ool&

%he genetic and molecular5)iological methods ena)le to identify and clone the

genes of en.ymes, inoled )oth in the formation of arious IAA con>ugates and

their hydrolytic disintegration& In 'articular, in immature seeds of 'ea and corn

endos'erm the genes of en.ymes cataly.ing the formation of esterified IAA

con>ugates= IARGc synthase 'artici'ating in the formation of a com'le* IAA

ester (ith glucose 5 15@5indole5-5acetyl5H595glucose +08, IAA5myo5

inosytoltransferase 6IAInos transferase7, cataly.ing a su)se


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elongation of these ne(ly5formed cells +"/, $& %he mechanisms of au*in

regulation of these key cell differentiation 'rocesses hae no( )een esta)lished at

the molecular leel&

%he mitotic cycle is regulated )y au*ins through heterodimeric 'rotein

com'le*es com'osed of catalytic su)unit 5 cyclin5de'endent kinase 6C9K7 and

regulatory su)unit 4 cyclin& Cyclins (ere first identified in sea urchin eggs as'roteins (hose


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high e*tent of se


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inoled in the M% desta)ili.ation during the cell transition from one 'hase of the

cell cycle t3 another, in 'articular, uniersal for all organisms cytosol

'hos'ho'rotein @'18stathmin +1!-; M% translocators, 'artici'ating in the

mitotic s'indle formation, and motor KGPs 'roteins 6kinesin5like 'roteins7,

inoled in maintaining its )i'olar structure +1!$ and also uniersal for all 'lants

negatiely charged motor KCBP 'rotein 6kinesin5like calmodulin )inding7,)inding (ith M% through a calciumcalmodulin com'le* +1!/; 'ositiely

charged motor 'roteins, for e*am'le, identified in to)acco cells %K?P1"/ 'rotein

6to)acco kinesin5related 'oly'e'tide7 (ith m&m& of 1"/ k9a (hose function is t3

maintain fragmo'last )i'olarity +1!0; E5tu)ulin and associated (ith it 'roteins,

centrosome 'roteins +1! such, as centrin +1!8; 'rotein 4 homologue of the

elongation factor QF15E +1!# and other 'roteins directly inoled in the s'atial

formation of M% into microtu)e organi.ed centers 6M%@Cs7 (hich are

centrosomes +11!; 'roteins 4 myosins, dynein5related 'oly'e'tides, 'roteins5

homologues of kinesin and illingelsolin'roteins family, associated (ith actinmicrofilaments and inoled in the actin microfilaments organi.ation control& All

these motor 'roteins hae )een found t3 interact (ith calmodulin and their

actiity is regulated in a Ca"T5de'endent manner +111 5 110&

%he C9Kcyclin com'le* is regulated at the arious leels= it is actiated

through 'hos'horylation, inoling C9K5actiating kinase CAK1At res'onsi)le

for the 'hos'horylation of the %hr510! end of C9K, found in arious organisms

+#1, 111 5 11#; this com'le* inhi)ition occurs in case of )inding CKI 4 negatie

regulators of C9K, and is follo(ed )y the cyclin su)units 'roteolysis +8$&

%here are numerous facts 'roing that the cell cycle is controlled )y au*in

through the regulation of e*'ression of genes that encode arious classes of C9K

and cyclins& For e*am'le, the results of o)seration of C9K and cyclin gene

transcri'tional actiity sho( that in au*in5treated cells and organs of arious

'lants the synthesis of C9C"aAt of 'rotein kinase, classified as A5class C9K, is

induced; a su)se


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leels of m?NA encoding '-$cdc"5like 'roteins in to)acco root cells& %he au*in

actiation of the cell cycle has )een esta)lished t3 re


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%hree ty'es of s'ecific functionally interrelated 'rotein kinases 6MAPK,

MAPKK, MAPKKK7 form a )asic module of the MAPK 'ath(ay& @ther

MAPKKK kinases 6MAPKKKK7 or R 'roteins such, as ?as 'roteins or their

heterotrimeric com'le*es, function as mediators )et(een sensitie t3 intra5

cellular signals and located in the 'lasmatic mem)rane 'rotein5rece'tors and

MAPK module +1$, 1$8&Biochemical and genetic studies +1$#, 1/! hae confirmed the e*istence of the

MAPK5cascade mechanism in mediating signals of au*ins and other

'hytohormones +1-", 1-8, 1/1 5 1/$& %he genetic analysis of au*in5sensitie

yeasts mutants sho(s that the I!I5gene encoded F5)o* 'roteins 6(hich are a 'art

of the Q- u)i


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'rocesses, for e*am'le, cell diision and elongation, NPK1 and MAPK5like

'rotein, actiating in mitosis, may 'reent au*in signals transmission in a

diiding cell&

No( one more regulatory com'onent of the MAPK5signal 'ath(ay of au*in 4

MAPK 'hos'hotase, regulating the MAPK actiity, has )een identified and an

im'ortant role of this en.yme in the au*in5induced gro(th of Arabidopsis has)een esta)lished +101& In 'articular, studies on au*in5resistant mutants hae

ena)led t3 isolate the ibr. mutant line of 'lants e*hi)iting an insignificant

sensitiity t3 IBA and 'ractically insensitie t3 inhi)iting concentrations of IAA,

synthetic au*ins ",$59, ",$59B and NAA, t3 inhi)itors of the au*in trans'ort 7

15na'hthyl'hthalamic and ",-,/5triiodo)en.oic acids as (ell as t3 a)scisic acid

6ABA7& %he 'henoty'e of ibr. mutants is similar t3 that of au*in5resistant

mutants, for e*am'le, a2r#mutants gro(n in light and haing long roots and

short hy'ocotyls& Moreoer, it has )een reealed that as a result of these

mutations a normal deelo'ment of the ascular system is distur)ed, leafindentation increases and accumulation of au*in5induced carriers decreases&

By the immune )lotting and 'romotor5re'orter genetic analysis it has )een

esta)lished that theI+!. gene e*'ression is o)sered in all tissues and organs of

(ild flo(ering 'lants and gymnos'erms& I+!. has )een sho(n t3 encode the

'rotein com'osed of "/ amino acid residues 6a& o&7 'ossessing a catalytic domain

6$# 4 18" a&o&7, -/O 5 identical (ith that 'resent in human MAPK 'hos'hotases

6MKP1 and PAC17 +10"&

%he studies hae ena)led t3 find out that I+!. gene5encoded MAPK5

'hos'hotase e*hi)its a de5'hos'horylating actiity to(ard signal com'onents of

)oth au*in and ABA that allo(s t3 make conclusion a)out a dual s'ecificity of

this MAPK, modeling signal (ays of au*ins and ABA&

Further detail studies on mutants (ith a distur)ed res'onse t3 the au*in

influence (ill facilitate the formation of a more correct idea concerning a

stimulating or inhi)iting influence of these 'hytohormones on the cell cycle&

+u=in7re8ulated 8enes controllin8 the cell elon8ation. Au*ins are

'hytohormones stimulating the cell elongation +"/& %he 'lant cell gro(th is

kno(n t3 )e initiated )y (ater u'take resulting from a stress rela*ation of cell

(alls +$& %he 'hytohormones, stimulating gro(th, cause cell (all elongation

)ut it is not yet clear ho( it is carried out& %he acidicD gro(th theory 'ostulatesthat in the 'lasmalemma au*in induces the 2T'um' o'eration and secretion of

hydrogen into the cell (all and conse


17/44

'ro)lems such, as measuring the cell (all '2, ham'er the 'rocess of making a

correct decision in this dis'ute&

A clue for understanding the mechanism of the au*in5regulated cell elongation

is, undou)tedly, a detail inestigation of the )iogenesis of cell (all 'rotein and

non5'rotein com'onents as (ell as hydrolytic en.ymes characteri.ed )y a s'ecific

actiity to(ard these com'onents& In com'liance (ith the data o)tained oer thelast fe( years, 'lant cell (alls hae )een found t3 hae fie )asic 'rotein classes

'laying a central 'art in the cell elongation gro(th +10-& %hey include

hydro*y'roline5rich glyco'roteins 62?RPs7 5 e*tensins and e*'ansins, glycine5

rich 'roteins 6R?Ps7, 'roline5rich 'roteins 6P?Ps7, Solanaceous lectins and

ara)inogalactan 'roteins& %he a)oe5enumerated 'roteins[ functions are

numerous )ut the main one is the inolement in the organi.ation of the 'rimary

cell (all car)ohydrate frame(ork that 'oints t3 their essential role in the cell

elongation gro(th regulation& Qolutionary and functionally, all these 'rotein

classes are related as regards hydro*y'roline5rich residues and similar as regardsnucleotide seor 'olymer 'roduced )y 'lasmatic mem)rane of higher 'lant&

%his en.yme maintains the synthesis of callose 'olysaccharide 61,-5H5glucan7&

Mor'hologically essential elements, 'roiding s'atial information for ne(ly

synthesi.ed cellulose microfi)rils and controlling their orientation, are cortical

1


18/44

M% re'resenting dynamic 'olymers com'osed of heterodimers of tu)ulin E4 and

H5su)units )eing found in correlation 1=1 +1", 1-, (hile a minor com'onent of

M% is tu)ulin \5isoform 'laying some 'art in the M% nucleus organi.ation&

Various tu)ulin isoforms may differentially modulate the M% function&

In the 'eriod of cell elongation the M% sta)ility is controlled )y arious

mechanisms +1$, 1/= 17 as a result of


19/44

1,"5H595*ylosyl 'olymer chains7 of the elongating 'rimary cell (all in

dicotyledonous 'lants forming a t(o5layer secondary (all, hae )een identified

in many 'lants +10#, 181& Studies results +181 5 18$ sho( that some increase in

the01gene e*'ression is o)sered under the influence of au*ins, gi))erellins

and )rassinosteroids in


20/44

measuring the de*transe actiity in deca'itated and non5deca'itated oats

coleo'tylesusing the thin5layer chromatogra'hy techni


21/44

treatment the m&m& alue of hemicellulose 'olysaccharides shifts t3 the range of

alues ty'ical of lo(5molecular com'ounds& %his fact indicates that the

e*'ression of endo51,-=1,$5H595 and e*o5H595glucanase genes is regulated

differentially )y IAA&

It should )e noted that during the last fe( years 1,$5H5endoglucanases,

characteri.ed )y s'ecific effects on 'lant cell (alls, hae also )een identified andisolated in many )acteria and fungi& For e*am'le, e*'eriments on studying the

en.ymic actiity of fungal 1,$5 H5endoglucanase 6encoded )y the gene 'el#=A7

from richoderma reesei (ith m&m& of "- k9a, 'ertaining t3 the glycoside

hydrolase family 1", hae esta)lished that in inactiated under the influence of

high tem'eratures and au*in5treated cell (alls of cucum)er 6'ucumis sativusc

Bur'ee Pickler7 hy'ocotyls this en.yme hydrolytic actiity enhances to(ard cell

(all *yloglucan 1,-=1,$5H5glucan 'olysaccharides& 2o(eer, this en.yme does

not sho( a similar actiity to(ard *ylan, ara)ino*ylan, galactomannan and

galactan 'olysaccharides +1#-& %he results o)tained in this study and in otherinestigations +1#$, conducted t3 reeal a role of the H5glucanase hydrolytic

en.yme in the molecular mechanisms of 'lant 'rotection res'onses t3 'athogens

action in to)acco 'lants, transformed )y )inar ector 'lasmids, constructed )y

genetic engineering technior

'olysaccharides 6i&e& causing, unlike hydrolytic en.ymes, a smooth shift of

'olysaccharides in the cell (all matri* (ithout su)stantial changes in itsstructure7 +1#0&

"1


22/44

A functional interaction (as esta)lished t3 e*ist )et(een e*'ansins 6(hich

hae the 'ro'erties of a 'rimary (all5loosening agents, i&e& e*'ansins can directly

induce (all e*tension7 and some hydrolases 6they may )e considered indirect, or

secondary, (all5loosening agents7 making cells more sensitie t3 an e*'ansin

action +1#-& Moreoer, the e*'ansin action (as found t3 )e s'ecific for cellulose

*yloglucan com'onents and not for cellulose glucomannan andhomogalacturonan 'olymers of the cell (all +1#& 9ata 'roing the correlation

)et(een the au*in hormone leel and e*'ansin actiity (ere also o)tained& For

e*am'le, some enhancement in the e*'ression of the e*'ansin5e12p=gene (as

o)sered in ethyolated hy'ocotyls of gro(ing tomato fruits 65ycopersicon

esculentumc Moneymaker7 treated (ith the synthetic au*in ",$59 for 1" hours

+1#!&

%hus, the 'lant cell elongation is controlled )y the t(o co5ordinated

mechanisms of the cell (all restructuring= the first one is the 'olymer (all

dissociation and the second one is a smooth changing of its structure (ithouthydrolysis of matri* 'olysaccharides& In the architecture creation of a t(o5layer

secondarily differetiated cell (all 6containing u' t3 0! O of cellulose relatie t3

the total cell (all mass7 )oth in monocotyledonous and dicotyledonous 'lants a

key role is 'layed )y cellulose synthase and callose synthase en.ymes 6'ertaining

t3 the class of integral mem)rane endo51,$5H5endogluconases, )elonging t3 the

cellulose synthase com'le*, res'onsi)le for )iosynthesis of 'lasmatic mem)rane

cellulose com'onents in secondary cell (alls and controlling the arrangement of

ne(ly synthesi.ed amor'hous cellulose chains as (ell as their length through

regulating the termination of their )iosynthesis7 and )y a num)er of en.ymes

inoled in glycosylation and fucosylation of *yloglucan, ara)inogalactan and

RA 'olysaccharides tightly interlacing cellulose microfi)rils t3 make a (all

stronger&

Se


23/44

C9!3, encoding QRs, locali.ed in mem)ranes and classified as cellulose

synthases&

%he 6orriganmutation of one of QRs, associated (ith the 'lasma mem)rane

and encoded )y the C9!gene, has )een esta)lished t3 cause formation of an

a))erant cell 'lates 6that testifies that K@? 'lays a key role during cytokinesis7,

an incom'lete cell (alls 6o(ing t3 a lo( cellulose leel7, and multinucleated cellsthat results in arious defects in Arabidopsis seedling mor'hology +"!!& %he

C9!gene has )een found t3 )e e*'ressed in all 'arts of mem)ranes of arious

'lant tissues (hile the C9!= and C9!3 genes are differentially e*'ressed in

deelo'ing leaf trichomes& Furthermore, C9!=gene e*'ression is o)sered in

deelo'ing root hairs (ithin the root differentiation .one, in the )asal region of

leaes, and floral organs, (hereas the C9!3 gene is e*'ressed in the )undle

sheath cells that surround the ascular )undle (ithin the leaf meso'hyll tissue as

(ell +"!1&

It has )een esta)lished that the au*in does not considera)ly affect the increasein the e*'ression leel of theC9!gene, encoding 'rotein (ith m&m& of " k9a,

and its homological genes 5 'el3 gene of tomato 657 1sculentum7, encoding

'rotein (ith m&m& of 08,/ k9a, and 'el#> gene of +rassica napus, encoding

'rotein (ith m&m& of 0# k9a that is eidence of a functional difference of QRs,

locali.ed in mem)ranes, from endo'lasmatic reticulum5secreted au*in5regulated

hydrolitic QRs +"!"&

Studies on rsw= mutants of Arabidopsis, allelic for 6orriganmutants, hae

sho(n that rsw= mutants are tem'erature5sensitie, 'henoty'ically similar t3

rsw#mutant'lants in their cellulose )iosynthesis, and in com'arison (ith (ild

'lants at a certain tem'erature 'roduce less than /! O of cellulose in roots +"!-&

Moreoer, rsw= mutants (ere found t3 hae some deiations in 'olysaccharide

'roduction of the cell (all hemicellulose matri*& In 6orriganmutants there (ere

o)sered some increase in the cell diameter, formation of holes in cell (alls and

final cell destruction )ecause of cellulose lo( leels& Similar 'henomena hae

)een detected also in rsw=mutants that is eidence of a key role of 'el#> and

C9!genes in the cellulose )iosynthesis in cell elongation 'eriod&

9uring a fe( last years genes of en.ymes, inoled in the )iosynthesis of

*yloglucan, ara)inogalactan and RA 'olysaccharides of the cell (all

hemicellulose matri*, tightly interlacing cellulose microfi)rils, hae )eenidentified and descri)ed )y the functional genome analysis& %hey include some

mem)ers of the en.yme family of genes, inoled in the )iosynthesis of

*yloglucans 4 *yloglucan *ylosyltransferases 6for e*am'le, At0# gene and

grou' of its homological genes, called At4=- in Arabidopsis +"!$, the '?5

gene family 6cellulose synthase5like7, encoding *yloglucan glucan synthase,

'artici'ating in the )iosynthesis of hemicellulose matri* com'onents, connecting

(ith each other *yloglucan 'olysaccharides 6in 'articular, in seeds of guarthere

(as identified a homologue of the Arabidopsis '?5Agene, locali.ed in Rolgi

com'le* mem)ranes and encoding H51,$5mannan synthase7 +"!/, genes of *ylansynthaseen.yme 6in 'articular,0?#gene of rice seedlings7 +"!0, res'onsi)le for

"-


24/44

the )iosynthesis of RA 'olysaccharides, com'osed of H51,$5*ylans, )ound (ith

H51,-5ara)inose and H51,"5linked glucuronic acid, as (ell as genes of glucan

synthase en.yme, cataly.ing the )iosynthesis of H51,$5glucans that hae a high

m&m& 6for e*am'le, the 4?#gene identified in corn mesocotyles7 +"!0&

%he PC?5analysis techni

It has )een esta)lished that all a)oe mentioned en.ymes of the 'olysaccharide

)iosynthesis of the cell (all hemicellulose matri* are confined t3 the Rolgi

a''aratus and are mem)rane 'roteins, containing a short amino5therminal

domain, facing the cyto'lasm, single transmem)rane domain, se'arated from a

glo)ular catalytic 'art of 'roteins )y se


25/44

and 'athogens 6it is suggested that e*tensin )inds negatiely charged 'athogens

and 'reents their su''ly into cells7& Q*tensin inceases the cell (all mechanical

strength through its intercalation as (ell as ionic and coalent interaction (ith

other 'olymers, in 'articular, (ith 'ectins& It (as esta)lished that catalysts of the

cell 'rotection res'onse t3 the action of elisitors or o*idatie stress are 'ero*idase

en.ymes, stimulating a


26/44

*onclusion. A considera)le 'rogress in the inestigation of the mechanism of

au*in regulatory effects on the cell differentiation key stages 4 cell diision and

elongation, has made the )asis for further fundamental studies of its role in the

gene e*'ression regulation in all stages of the 'lant ontogenesis that (ill

contri)ute t3 the adancement in the deelo'ment of ne( genetic engineering

)iotechnologies& In the future such deelo'ments may )e a''lied t3 correct thekey stages of the 'lant mor'hogenesis, to create of more 'roductie and resistant

t3 unfaoura)le enironmental conditions sorts of 'lants&

&)/)&)N*)3

1& 5eopold A7'7$ %ooden 5787 2ormonal regulatory system in 'lants 2orm&

?egul& 9eelo'& 4 1#8$& 4 "& 5 P& $ 4 ""&

"& ?now !& %he correlatie inhi)ition of the gro(th of a*illary )uds Ann& Bot& 41#"/& 5 -#& 4 P& 8$1 4 8/#&

-& Eareing "7;7Qndogenous inhi)itors in seed germination and dormancy

Qncyclo'edia of 'lant 'hysiology Qd& :& ?uhland& 4 Berlin; R`ttingen;

2eidel)erg= S'ringer, 1#0/& 4 Vol&1/, 't "& 4 P& #!# 4 #"$&

$& 8arwin '7$ 8arwin ;7 %he 'o(er of moement in 'lants Qd& & Murray& 4

Gondon, 188!& 4 88! '&

/& +rown 77$ @orris 477 ?esearches of the germination of some of the

4ramineae & Chem& Soc& 4 18#!& 4 Vol& /& 4 P& $/8 4 /"8&

0&5oeb F7 Chemical )asis of correlation Bot& Ra.& 4 1#18& 4 Vol& 0/& 4 P& 1/! 4

1$&

& @olisch 79ie Ge)ensdauer der Pflan.e& 4 ena= Fischer, 1#"8 6%ransl& )y

Fulling Q&2& 5 Gancaster= Science 'ress, 1#-87&

8& ?ytni6 C7@7 %i6olay 4rygorevitch Cholodny 6%o the century from the

)irthday7 Ukr& Bot& & 5 1#8"& 4 Vol& -#, -& 5 P& 1 5 -&

#&@er6is A7I7 %ro'isms of 'lants in the light of Kholodny 5 :ent theory Ukr&

Bot& & 5 1#8"& 4 Vol& -#, -& 5 P& 10 5 -1&

1!& Eent ;7$ himann C7D7 Phytohormones& 5 Ne( Work= MacMillan, 1#-& 4

"#$ b&

11& himann C7D7Plant gro(th Fundamental as'ects of normal and malignantgro(th Qd& :&:& No(inski 4 Amsterdam= Qlseier, 1#0!& 4 P&$8 4 8""&

1"&"ilet "717Action des gi))erellins sur lactiite au*ines5o*ydasiit& 5 Ne( Work= 2a(orth Press, "!!!& 4 "//

P&

1$& Arteca !7 Plant Rro(th Su)stances= Princi'les and A''lications& 5 Ne(

Work= Cha'man and 2all, 1##0& 5 "// '&

"0


27/44

1/& 'reelman !7 A7$ @ullet F7 17 @ligosaccharins, )rassinolides, and >asmonates=

nontraditional regulators of 'lant gro(th, deelo'ment, and gene e*'ression

Plant Cell& 4 1##& 4 Vol& #& 4 P& 1"11 5 1""-&

10& 4ross 87$ "arthier +7 Noel natural su)stances acting in 'lant gro(th

regulation & Plant Rro(th ?egul& 5 1##$& 4 Vol& 1-& 5 P& #- 5 11$&

1&5utsen6o 17C7$ @aruch6o 17A7$ 5eonova 747 %he fusicoccine action on theearly stages of germination of sorghum at salting %hes& ?e'& IV Internat& Conf&

Plant Rro(th and 9eelo'ment ?egulatorsD& 4 Mosco(= %he Mosco( State

Agronomic Uni& Pu)l&, 1##& 5 P& 1!/&

18& 'haila6hian@7'h7$ +uten6o !747$ Culaeva 97%7$ Cefeli D7I7$ A6senova %7"7

%erminology of gro(th and deelo'ment of higher 'lants& 5 Mosco(= Science,

1#8"& 5 #0 '&

1#& @c'ourt "7 Renetic analysis of hormone signalling Annu& ?e& Plant

Physiol& Plant Mol& Biol& 5 1###& 5 Vol& /!& 5 P& "1# 5 "$-&

"!& +ecraft "7E7$ ?u6-oon Cang$ ?ang-4on ?uh& %he mai.e C?INKGW$rece'tor kinase controls a cell5autonomous differentiation res'onse Plant

Physiol& 4 "!!1& 4 Vol& 1", "& 4 P& $80 4 $#0&

"1& Divanco F7 @7$ ;lores 7 17 Control of root formation )y 'lant gro(th

regulation %he Plant Rro(th ?egulators in Agriculture and 2orticulture& ?ole

and Commercial Uses Qd& S& B&Amar>it 5 Ne( Work= 2a(orth Press, "!!!& 5 P& 1

5 "/&

""& ?hneider 17A7$ Eightman ;7Au*ins Phytohormones and related com'ounds

5 a com'rehensie treatise&5 Amsterdam= Qlseier, North 2olland Biomedical

Press, 1#8& 5 P& "# 5 1!/&

"-&Cefeli D7Natural gro(th inhi)itors and 'hytohormones& 5 2aague= unh Pu)l&,

1#8& 5 "#$ P&

"$&8orffling C79as 2ormonsystem der 'flan.en& 4 Stuttgard; Ne( Work= Reorg

%hieme Verlag, 1#8"& 5 -!$ P&

"/&Cende 7$ eevaart F7 A7 87%he fie classical 'lant hormones Plant Cell&

4 1##& 4 Vol& #& 4 P& 11# 5 1"1!&

"0&Fa6ubows6a A7$ Cowalczy6 ?7%he au*in con>ugate 15@5indole5-5acetyl5595

glucose is synthesi.ed in immature legume seeds )y IARlc synthase and may )e

used for modification of some high molecular (eight com'ounds & Q*'& Bot& 4

"!!$& 4 Vol&//, -#8& 4 P& #1 4 8!1&"& edden ;7$ "hillips A7 Mani'ulation of hormone )iosynthetic genes in

transgenic 'lants Curr& @'in& Biotechnol& 5 "!!!& 4 Vol&11& 5 P& 1-!51-&

"8&;iccadenti %7$ ?estili ?7$ "andolfini 7$ 'irillo '7$ !otino 47 57$ ?pena A7

Renetic engineering of 'arthenocar'ic fruit deelo'ment in tomato Mol&

Breeding& 4 1###& 4 Vol& /& 4 P& $0- 5 $!&

"#&!omano '7$ ein @7$ Clee 7Inactiation of au*in in to)acco transformed

(ith the indoleacetic acid5lysine synthetase gene of"seudomonas savastonoi

Renes and 9eelo'& 4 1##1& 4 Vol&/& 4 P& $-8 4 $$0&

-!&+artel +7Au*in )iosynthesis Annu& ?e& Plant& Physiol& Plant Mol& Biol& 41##& 4 Vol& $8& 4 P& /1500&

"


28/44

-1&%ormanly F7$ ?lovin F7 "7$ 'ohen F7 87?ethinking au*in )iosynthesis and

meta)olism Plant Physiol& 4 1##/& 4 Vol&1!& 4 P& -"- 5 -"#&

-"& @Gller A7$ Eeiler 17E7 Indolic costituents and indole5-5acetic acid

)iosinthesis in the (ild5ty'e and a try'to'han au*otro'h mutant of Arabidopsis

thaliana Planta& 4 "!!!& 4 Vol& "11& 4 P& 8// 4 80-&

--&!e6oslavs6aya %7I7$ +andurs6i !7?7 Indole as a 'recursor of indole5-5aceticacid inea mays Phytochemistry& 4 1##$& 4 Vol& -/& 4 P& #!/ 4 #!#&

-$&8olan 57Pointing roots in the right direction= the role of au*in trans'ort in

res'onse to graity Ren& and 9eelo'& 4 1##8& 4 Vol&1", 1$& 4 P& "!#1 5

"!#/&

-/&+artel +7$ ;in6 47 !79ifferential regulation of au*in5'roducing nitrilase gene

family inArabidopsis thaliana Proc& Nat& Acad& Sci& USA& 4 1##$& 4 Vol& #1& 4

P& 00$# 5 00/-&

-0& ?chmidt !7 '7$ @Gller A7$ ain !7$ +artling 87$ Eeiler 17 E7 %ransgenic

to)acco 'lants e*'ressing theArabidopsis thaliananitrilase II en.yme Plant &4 1##0& 4 Vol& #& 4 P& 08- 5 0#1&

-& 4rsis ?7$ ?auerteig ?7$ %euhaus C7$ Albrecht @7$ !ossiter F7$ @Gller 57F7

Physiological analysis of transgenicArabidopsis thaliana'lants e*'ressing one

nitrilase isoform in sense or antisense direction Plant Physiol& 4 1##8& 4

Vol&1/-& 4 P& $$0 5 $/0&

-8&+a6 ?7$ ;eyereisen !7%he inolement of t(o P$/! en.ymes, CWP8-B1 and

CWP8-A1, in au*in homeostasis and glucosinolate )iosynthesis Plant Physiol& 4

"!!1& 4 Vol& 1"& 4 P& 1!8 5 118&

-#&+a6 ?7$ a2 ;717$ ;eldmann C7A7$ 4albraith 87A7$ ;eyereisen !7 CWP8-B1, a

cytochrome P$/! at the meta)olic )ranch'oint in au*in and indole glucosinolate

)iosynthesis inArabidopsis thaliana Plant Cell& 4 "!!1& 4 Vol& 1-& 4 P& 1!1 4

111&

$!& 8elarue @7$ "rinsen 17$ 9nc6elen 7D7$ 'aboche @7$ +ellini '7 ?ur=

mutations ofArabidopsis thalianadefine a ne( locus inoled in the control of

au*in homeostasis Plant & 4 1##8& 4 Vol& 1$& 4 P& 0!- 4 011&

$1& 'ollett '717$ arberd %7"7$ 5eyser 97 2ormonal interactions in the control of

Arabidopsishy'ocotyl elongation Plant Physiol& 4 "!!!& 4 Vol&1"$& 4 P& //- 4

/01&

$"& @izutani @7$ Eard 17$ 9hta 87 Cytochrome P$/! gene su'erfamily inArabidopsis thaliana= isolation of c9NAs, differential e*'ression, and ?FGP

ma''ing of multi'le cytochromes P$/! Plant Mol& Biol& 4 1##8& 4 Vol& -& 4 P&

-# 4 /"&

$-& !eymond "7$ Eeber 7$ 8amond @7$ ;armer 1717 9ifferential gene

e*'ression in res'onds to mechanical (ounding and insect feeding in

Arabidopsis Plant Cell& 4 "!!!& 4 Vol& 1"& 4 P& ! 4 1#&

$$&


29/44

$/&u F7 ?egulation of 'ero*isome )iogenesis and function MSU59@Q Plant

?esearch Ga)oratory& %hirty5Qighth Annual ?e'ort& USA& 4 Ne( Work, "!!-& 5

P& 1# 4 "0&

$0&olman +7C7$ ?ilva I787$ +artel +7 %he Ara)ido'sisp2a#mutant is defectie

in an A%P5)inding cassette trans'orter5like 'rotein re


30/44

/#& ayashi @7$ oriyama C7$ Condo @7$ %ishimura @7 ",$5

9ichloro'heno*y)utyric acid5resistant mutants of Arabidopsis hae defects in

glyo*ysomal fatty acid 5o*idation Plant Cell& 5 1##8& 4 Vol& 1!& 4 P& 18- 4

1#/&

0!&ayashi @7$ %ito C7$ oriyama-Cato C7$ Condo @7$ amaya 7$ %ishimura @7

AtPe*1$' maintains 'ero*isomal functions )y determining 'rotein targeting tothree kinds of 'lant 'ero*isomes QMB@ & 4 "!!!& 4 Vol& 1#& 4 P& /!1 5 /1!&

01& ?ytni6 C7@7$ @usaten6o 57I7$ Dasu6 %7"7$ Dedenicheva %7"7$ 4eneralova

D7@7$ @artin 4747$ %esterova A7%7 2ormonal com'le* of 'lants and mushrooms&

4 Kie, "!!-& 4 180 '&

0"& ?lovin F7"7$ +andurs6i !7?7$ 'ohen F787 Au*in Biochemistry and molecular

)iology of 'lant hormones Qds& P&&& 2ooykaas, M&A& 2all, K&?& Gi))enga& 4

Amsterdam= Qlseier Science BV, 1###& 4 P& 11/ 4 1$!&

0-& all "7F7 Indole5-5acetyl5myo5inositol in kernels of 9ryza sativa

Phytochemistry& 4 1#8!& 4 Vol& 1#& 4 P& "1"1 4 "1"-&0$&8omagals6i E7$ ?chulze A7$ +andurs6i !7?7 Isolation and characteri.ation of

ester of indole5-5acetic acid from the li


31/44

-&Clee F7 7%he effects of oer'roduction of t(oAgrobacterium tumefaciens

%59NA au*in )iosynthetic gene 'roducts in transgenic 'etunia 'lants Renes

and 9eelo'& 4 1#8& 4 Vol&1& 4 P& 80 5 #0&

$& 'osgrove 87F7 ?ela*ation in a high5stress enironment= %he molecular )ases

of e*tensi)le cell (alls and cell enlargement Plant Cell& 4 1##& 4 Vol& #& 4 P&

1!-1 4 1!$1&/&Ito @7Factors controlling cyclin B e*'ression Plant& Mol& Biol& 4 "!!!& 4

Vol& $-, /0& 4 P& 0 5 0#!&

0& ?etiady 7 7$ ?c6ine @7$ ariguchi %7$ amamoto 7$ Couchi 7$ ?hinmyo A &

%o)acco mitotic cyclins= cloning, characteri.ation, gene e*'ression and

functional assay Plant & 4 1##/& 4 Vol& 8& 4 P& #$# 5 #/&

&ata ?7$ Couchi 7$ ?uzu6a F7$ Ishii 7Isolation and characteri.ation of c9NA

clones for 'lant cyclins QMB@ & 4 1##1& 4 Vol& 1!& 4 P& "081 5 "088&

8& 4olstyen !7$ ?tandart %7$ @ac6ie ?7$ 'olman A7$ +low F7$ !uderman F7$ Eu

@7$ unt & %he role of cyclin synthesis, modification and destruction in thecontrol of cell diision & Cell Sci& 4 1#8#& 4 Vol& 1"& 4 P& 5 #&

#& Couchi 7$ ?e6ine 7$ ata ?7 9istinct classes of mitotic cyclins are

differentially e*'ressed in the soy)ean shoot a'e* during the cell cycle Plant

Cell& 4 1##/& 4 Vol& & 4 P& 11$- 5 11//&

8!&%igg 17 A7 Cyclin5de'endent 'rotein kinases= key regulators of the eukaryotic

cell cycle Bioessays& 4 1##/& 4 Vol& 1& 4 P& $1 5 $8!&

81&!enaudin F7 "7$ 8oonan F7 7$ ;reeman 87$ ashimoto F7$ irt 7$ InzH 87$

Facobs 7 Plant cyclins= a unified nomenclature for 'lant A5, B5 and 95ty'e

cyclins )ased on se


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8& @ironov D7$ 8e Deylder 57$ Dan @ontagu @7$ InzH 87 Cyclin5de'endent

kinases and cell diision in 'lants= the ne*us Plant Cell& 4 1###& 4 Vol& 11& 4 P&

/!# 5 /""&

88& %igg 17 A7 Cyclin5de'endent kinase = at the cross5roads of transcri'tion,

9NA re'air and cell cycle control Curr& @'in& Cell& Biol& 4 1##0& 4 Vol& 8& 4

P& -1" 5 -1&8#& @HszrKros 7$ @is6olczi "7$ Ayaydin ;7$ "ett6L-?zandther A7$ "eres A7$

@agyar 7$ orvKth 47 D7$ +a6o 57$ ;ehHr A7$ 8udits 87 Multi'le cyclin5

de'endent kinase com'le*es and 'hos'hatases control R"M 'rogression in

alfalfa cells Plant Mol& Biol& 4 "!!!& 4 Vol& $-, /0& 4 P& /#/ 5 0!/&

#!& ?egers 47$ !ouzH "7$ Dan @ontagu @7$ InzH 87Cyclin5de'endent kinases in

'lants Plant cell 'roliferation and its regulation in gro(th and deelo'ment

Qds & Bryant, & :iley& 5 Chichester, UK, 1##& 5 P& 1 5 1#&

#1& Foubes F7$ 'hevalier '7$ 8udits 87$ eberle-+ors 17$ Inze 87$


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1!!7 Asada 7$ Curiyama !7$ ?hibao6a 7%K?P1"/, a kinesin5related 'rotein

inoled in the centrosome5inde'endent organi.ation of the cytokinetic a''aratus

in to)acco BW5" cells & Cell Sci& 4 1##& 4 Vol& 11!& 4 P& 1# 5 18#&

1!1&@easday D7$ @oore 57$ !etna6aran !7$ 5ee F7$ 8onoviel @7$ %eiman A7 @7$

Andrews +7A family of cyclin5like 'roteins that interacts (ith Pho8/ cyclin5

de'endent kinase Moll& Cell Biol& 4 1##& 4 Vol& 1& 4 P& 1"1" 5 1""-&1!"& 8urfel 7$ ;eiler 7 ?7$ 4ruissem E7 ?etino)lastoma5related 'roteins in

'lants= homologues or orthologues of their meta.oan counter'arts Plant Mol&

Biol& 4 "!!!& 4 Vol& $-& 4 P& 0-/ 5 0$"&

1!-& +elmont 57$ @itchison 7 Identification of a 'rotein that interacts (ith

tu)ulin dimers and increases the catastro'he rates of microtu)ules Cell& 5 1##0&

4 Vol& 8$& 5 P& 0"- 5 0-1&

1!$&5iu +7$ 'yr !7$ "alevitz +7 A7A kinesin5like 'rotein, KatA', in the cell of

Arabidopsisand other 'lants Plant Cell& 5 1##0& 4 Vol& 8& 5 P& 11# 5 1-"&

1!/& Dos F7 E7$ ?afadi ;7$ !eddy A7 ?7$ epler "7 C7 Kinesin5like calmodulin5)inding 'rotein is differentially inoled in cell diision Plant Cell& 5 "!!!& 4

Vol& 1"& 5 P& ## 5 ##!&

1!0&Asada 7$ 'ollings 87Molecular motors in higher 'lants %rends Plant Sci&

5 1##& 4 Vol& "& 5 P& "# 5 -&

1!& immerman E7$ ?par6s '7 A7$ 8o2sey ?7 F7Amor'hous no longer= the

centrosome comes into focus Curr& @'in& Cell Biol& 5 1###& 4 Vol& 11& 5 P& 1""

5 1"8&

1!8&8ictenberg F7 +7$ immerman E7$ ?par6s '7 A7$ oung A7$ Didair '7$ heng

7$ 'arrington E7$ ;ay ;7 ?7$ 8o2sey ?7 F7 Pericentrin and 5tu)ulin form a

'rotein com'le* and are organised into a noel lattice at the centrosome & Cell

Biol& 5 1##8& 4 Vol& 1$1& 5 P& 10- 5 1$&

1!#& 8urso %7 A7$ 'yr !7 F7 A calmodulin5sensitie interaction )et(een

microtu)ules and a higher 'lant homologue of 'rotein translation elongation

factor QF5 Plant Cell& 5 1##$& 4 Vol& 0& 5 P& 8#- 5 #!/&

11!& Dantard @7$ 'owling !7$ 8elichere '7 Cell cycle regulation of the

microtu)ular cytoskeleton Plant Mol& Biol& 5 "!!!& 4 Vol& $-& 5 P& 0#1 5 !-&

111& 'ai 47$ !omagnoli ?7$ @oscatelli A7$ 'resti @7Qidence for microtu)ule5

)ased organelle trans'ort in the 'ollen tu)e Cell Biology of Plant and Fungal

%i' Rro(th Qds& A& Reitmann, M& Cresti&5 Amsterdam= I@S Press, "!!1& 5 P& 1

5 1"&

11"& @oscatelli A7$ 'ai 47$ 'resti @79ynein related 'oly'e'tides during 'ollen

tu)e gro(th Cell Biology of Plant and Fungal %i' Rro(th Qds A& Reitmann,

M& Cresti& 5 Amsterdam= I@S Press, "!!1& 5 P& 1- 5 "0&

11-& Didali 57$ oldaway-'lar6e @7$ epler "7C7 %he calciumcytoskeleton

connection in 'ollen tu)e gro(th Cell Biology of Plant and Fungal %i'

Rro(th Qds A& Reitmann, M& Cresti& 5 Amsterdam= I@S Press, "!!1& 5 P& " 5

-/&

--


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11$& eath +7$ ?6alamera 87 ?egulation of ti' mor'hogenesis )y the

cytoskeleton and calcium ions Cell Biology of Plant and Fungal %i' Rro(th

Qds& A& Reitmann, M& Cresti& 5 Amsterdam= I@S Press, "!!1& 5 P& - 5 /-&

11/& o6ota 17$ @uto ?7$ ?himmen 7Inhi)itory regulation of higher5'lant myosin

)y Ca"Tions Plant Physiol& 5 1###& 4 Vol& 11#& 5 P& "-1 5 "$!&

110& o6ota 17$ @uto ?7$ ?himmen 7 Ca"T

5calmodulin su'resses the F5actin)inding actiity of a 1-/ k9a actin5)undling 'rotein isolated from lily 'ollen

tu)es Plant Physiol& 5 "!!!& 4 Vol& 1"-& 5 P& 0$/ 5 0/$&

11& @endentrall @7 87$ odge A7 17 ?egulation of cdc "8 cyclin5de'endent

'rotein kinase actiity during the cell cycle of the yeast ?accharomyces

'erevisiae Micro)iol& Mol& Biol& ?e& 4 1##8& 4 Vol& 0"& 4 P& 11#1 5 1"$-&

118& 4overse A7$ de Almeida 1ngler F7$ Derhees F7$ Dan der Crol ?7$ elder F7$

'heysen 47 Cell cycle actiation )y 'lant 'arasitic nematodes Plant& Mol& Biol&

4 "!!!& 4 Vol& $-, /0& 4 P& $ 501&

11#& assan F7 "7$ FaMuenoud @7$ 5eopold "7$ ?chultz ?7 F7$ %igg 17 A7Identification of human cyclin5de'endent kinase 8, a 'utatie 'roteine kinase

'artner for cyclin C Proc& Nat& Acad& Sci& USA& 4 1##/& 4 Vol& #"& 4 P& 881 5

88/&

1"!&+urssens ?7$ Dan @ontagu @7$ InzH 87%he cell cycle inArabidopsis Plant

Physiol& Biochem& 4 1##8& 4 Vol& -0& 4 P& # 5 1#&

1"1&Fohn "7 '7$ hang C7$ 8ong '7$ 8iederich 57$ Eightman ;7 '-$cdc"related

'roteins in control of cell cycle 'rogression, the s(itch )et(een diision and

differentiation in tissue deelo'ment and stimulation of diision )y au*in and

cytokinin Aust& & Plant Physiol& 4 1##-& 4 Vol& "!& 4 P& /!- 5 /"0&

1""& Eang 7$ Ji J7$ ?chorr "7$ 'utler A7 F7$ 'rosby E7 57$ ;ow6e 57 '7 ICK1, a

cyclin5de'endent 'rotein kinase inhi)itor from Arabidopsis thaliana interacts

(ith )oth Cdc"a and Cyc 9- and its e*'ression is induced )y a)scisic acid

Plant & 4 1##8& 4 Vol& 1/& 4 P& /!1 5 /1!&

1"-&!eding "7$ ?houl 97$ InzH 87$ Dan @ontagu @7$ Dan 9rc6elen 7Geels of

endogenous cytokinins, indole5-5acetic acid and a)scisic acid during the cell

cycle of synchroni.ed to)acco BW5" cells FQBS Gett& 4 1##0& 4 Vol& -#1& 4 P&

1/ 5 18!&

1"$&8oerner "7$ Forgensen F7 17$ ou !7$ ?teppuhn F7$ 5amb '7Control of root

gro(th and deelo'ment )y cyclin e*'ression Nature& 4 1##0& 4 Vol& -8!& 4 P&/"! 5 /"-&

1"/& 'hung ?7 C7$ "arish !7 E& Studies on the 'romoter ofArabidopsis thaliana

cdc=agene FQBS Gett& 4 1##/& 4 Vol& -0"& 4 P&"1/ 5 "1#&

1"0&+Ngre 57$ @es6iene I7$ eberle-+ors 17$ irf 7Stressing the role of MAP

kinases in mitogenic stimulation Plant& Mol& Biol& 4 "!!!& 4 Vol& $-, /0& 4

P& !/ 5 18&

1"&CGltz 8& Phylogenetic and functional classification of mitogen5 and stress5

actiated 'rotein kinases & Mol& Qol& 4 1##8& 4 Vol& $0& 4 P& /1 5 /88&

1"8& irt 7MAP kinases in 'lant signal transduction ?esults Pro)l& Cell&9iffer& 4 "!!!& 4 Vol& "& 4 P& 1 5 #&

-$


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1"#&CGltz 87Phylogenetic and functional classification of mitogen5 and stress5

actiated 'rotein kinases & Mol& Qol& 4 1##8& 4 Vol& $0& 4 P& /1 5 /88&

1-!&@izoguchi 7$ 4otoh 7$ %ishida 17$ amaguchi-?hinoza6i C7$ ayashida %7$

Iwasa6i 7$ Camada 7$ ?hinoza6i C7 Characteri.ation of t(o c9NAs that encode

MAP kinase homologues inArabidopsis thalianaand analysis of the 'ossi)le role

of au*in in actiating such kinase actiities in cultured cells Plant & 4 1##$& 4Vol& /& 4 P& 111 5 1""&

1-1& @orris "7 '7$ 4uerrier 87$ 5eung F7$ 4iraudat F7 Cloning and

characteri.ation of@1C#, anArabidopsisgene encoding a homologue of MAP

kinase kinase Plant Mol& Biol& 4 1##& 4 Vol& -/& 4 P& 1!/ 5 1!0$&

1-"&Fona6 '7$ Ciegerl ?7$ 5igterin6 E7$ +ar6er "7 F7$ us6isson %7 ?7$ irt 7

Stress signalling in 'lants= a MAP kinase 'ath(ay is actiated )y cold and

drought Proc& Nat& Acad& Sci& USA& 41##0& 4 Vol& #-& 4 P& 11"$ 5 11"#&

1--&unter 7Signalling= "!!! and )eyond Cell& 4 "!!!& 4 Vol& 1!!& 5 P& 11- 5

1"&1-$&8ecroocM-;errant D7$ 8ecroocM ?7$ Dan Eent F7$ ?chmidt 17$ Creis @7 A

homolog of the @A"B1!C family of 'rotein kinase genes is e*'ressed in

egetatie and in female re'roductie organs of"etunia hybrida Plant& Mol&

Biol& 4 1##/& 4 Vol& "& 4 P& --# 5 -/!&

1-/&!omeis 7$ "iedras "7$ hang ?7 J7$ Clessig 87 ;7$ irt 7$ Fones F7?a'id

Ar #5 and cf5#5de'endent actiation of MAP kinases in to)acco cell cultures and

leaes= conergence of resistance gene, elicitor, (ound, and salicylate

res'onses Plant Cell& 4 1###& 4 Vol& 11& 4 P& "-5"8&

1-0&5igterin6 E7$ Cro 7$ zur %ieden


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1$-& 4arrington 7 "7$ Fohnson 47 [email protected] and regulation of mitogen5

actiated 'rotein kinase signaling 'ath(ays Curr& @'in& Biol& 5 1###& 4 Vol& 11&

5 P& "11 5 "18&

1$$& amal A7$ Fouannic A7 ?7$ 5eprince @7$ Creis @7$ enry 7 Molecular

characterisation and e*'ression of anArabidopsis thalianaG& MAP kinase kinase

c9NA, AtMAP"Ka Plant Sci& 5 1###&5 Vol& 1$!& 5 P& $# 5 0$&1$/&Ichimura C7$ @izoguchi 7$ ayashida %7$ ?e6i @7$ ?chinoza6i C7Molecular

cloning and characteri.ation of three c9NAs encoding 'utatie mitoge5actiated

'rotein kinase kinases 6MAPKKs7 inArabidopsis thaliana 9NA ?es& 5 1##8 a&

4 Vol& /& 5 P& -$1 5 -$8&

1$0&Fouannic ?7$ amal A7$ 5eprince A7 ?7$ regear F7 E7$ Creis @7$ enry 7

Characterisation of noel 'lant genes encoding MQKKS%Q11 and ?AF5related

'rotein kinases Rene& 5 1###& 4 Vol& ""#& 5 P& 11 5 181&

1$& 5eprince A7 ?7$ Fouannic ?7$ amal A7$ Creis @7$ enry 7 Molecular

characterisation of 'lant c9NAs BnMAP$K1 and BnMAP$K" )elonging tothe RCKSPS1 su)family of MAP kinase kinase kinase kinase Biochim& etBio'hys& Acta& 5 1###& 4 Vol& 1$$$& 5 P& 1 5 1-&

1$8&!ommel '7$ afen 17 ?as, a ersatile cellular s(itch Curr& Biol& 5 1##8& 4

Vol& 8& 5 P& $1" 5 $18&

1$#&


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1/& !ogers ?7$ Eells !7$ !echsteiner @7 Amino acid se


38/44


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18&Arrowsmith 87A7$ de ?ilva F7Characteri.ation of t(o tomato fruit5e*'ressed

c9NAs encoding *yloglucan endotransglycosylases Plant Mol& Biol& 4 1##/& 4

Vol& "8& 4 P& -#1 4 $!-&

188&Co6a '7D7$ 'erny !717$ 4ardner !747$ %oguchi 7$ ;uio6a ?7$ a6atsuto ?7$

oshida ?7$ 'louse ?787A 'utatie role for the tomato genes8

18#& 'ampbell "7$ +raam F7 yloglucan endotransglycosylases= diersity of

genes, en.ymes and 'otential (all5modifying functions %rends Plant Sci& 4

1###)& 4 Vol& $& 4 P& -01 5 -00&

1#!&Cota6e 7$ %a6agawa %7$ a6eda C7$ ?a6urai %7Au*in5induced elongation

gro(th and e*'ressions of cell (all5)ound e*o5 and endo55glucanases in Barley

coleo'tiles Plant and Cell Physiol& 4 "!!!& 4 Vol& $1, 11& 4 P& 1"" 5 1"8&

1#1&@PlhP @7$


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/%(4&)3

/i8. 1. %he scheme for the indole5-5acetic acid 6IAA7 )iosynthesis from

try'to'han +-8= the en.yme 6mem)er of cytochrome P$/! 6monoo*igenase7

family7 encoded )y '"*+gene is cataly.er of synthesis indole5-5acetaldo*ime

4 the general 'redecessor of IAA and their con>ugates 4 indole glucosinolates, in

the synthesis of (hich is inoled en.yme 6mem)er of cytochrome P $/! family7

encoded )y '",3+#gene&

$"


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/i8. 2.A suggested model for PA1 'rotein function in Ara)ido'sis +$0& PA1

'rotein, homological for human and yeast 'roteins, is locali.ed in the

'ero*isomal mem)rane and trans'orts acyl5CoA esters of FA 6FA5CoA7, indole5

-5)utyric acid 6IBA537 and also ",$5dichloro'heno*y)utyric acid 6",$59B5

37 into the 'ero*isomes (here they are cata)olised to succinate, IAA, and ",$5

9 res'ectiely& Mutant 'lants resistant t3 the action of IBA and ",$59B are

defectie in acyl5CoA o*idase 6ac237, multifunctional 'rotein 6aim#7 and thiolase

6ped#7 that is a direct eidence of these en.ymes inolement in IBA and ",$59B

H5o*idation&

$-


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/i8. #.Schematic re'resentation of the structure of a *yloglucan ty'ical of that

found in the cell (alls of many dicots, and the en.ymes needed to synthesi.e it

+"!0= glucan synthases inoling in the glucan 6Rlc7 )iosynthesis, 5

fucosyltransferases, 5galactosyltransferases and 5*ylosyltransferases, >oining

olygosaccharide molecules 4 fucose 6Fuc7, galactose 6Ral7 and *ylose 6yl7

accordingly to glucan )ack)one synthesi.ed de novo&

Genes of Auxin Biosynthesis and Auxin-regulated Genes

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