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