Механізми природної імунності рослини

Природна імунність рослини забезпечується конститутивним та індукованими механізмами. Конститутивний механізм обумовлений будовою клітинної стінки рослини, а різновиди інду­кованих — виникають теля взаємодії рослини з патогенними некротичними мікрорганізмами, непатогенними бактеріями, а також після...

Full description

Saved in:
Bibliographic Details
Published in:Біополімери і клітина
Date:2006
Main Author: Козировська, Н.О.
Format: Article
Language:Ukrainian
Published: Інститут молекулярної біології і генетики НАН України 2006
Subjects:
Online Access:https://nasplib.isofts.kiev.ua/handle/123456789/156471
Tags: Add Tag
No Tags, Be the first to tag this record!
Journal Title:Digital Library of Periodicals of National Academy of Sciences of Ukraine
Cite this:Механізми природної імунності рослини / Н.О. Козировська // Біополімери і клітина. — 2006. — Т. 22, № 2. — С. 91-101. — Бібліогр.: 112 назв. — укр., англ.

Institution

Digital Library of Periodicals of National Academy of Sciences of Ukraine
_version_ 1859915944899379200
author Козировська, Н.О.
author_facet Козировська, Н.О.
citation_txt Механізми природної імунності рослини / Н.О. Козировська // Біополімери і клітина. — 2006. — Т. 22, № 2. — С. 91-101. — Бібліогр.: 112 назв. — укр., англ.
collection DSpace DC
container_title Біополімери і клітина
description Природна імунність рослини забезпечується конститутивним та індукованими механізмами. Конститутивний механізм обумовлений будовою клітинної стінки рослини, а різновиди інду­кованих — виникають теля взаємодії рослини з патогенними некротичними мікрорганізмами, непатогенними бактеріями, а також після обробки деякими природними або синтетичними речовинами. Метою представленого огляду є висвітлення обох типів механізмів формування стійкості до патогенів та інших стресів. Plant innate immunity is assured by both constitutive and induced mechanisms. The constitutive barrier for pathogens relies on the plant cell wall structure, and the varieties of inducible systemic resistance result from interaction of the plant with pathogenic necrotrophic microorganisms, nonpathogenic bacteria, and also after the contact with some natural or synthetic substances. Description of both mechanisms of plant systemic resistance to pathogens and other stressors is the purpose of the review. Природная иммунность растений обеспечивается конститу­тивным и индуцироваными механизмами. Конститутивный механизм обусловлен строением клеточной стенки растения, а разновидности индуцированных — возникают после взаимо­действия растения с патогенными некротрофными микроор­ ганизмами, непатогенными бактериями, а также после кон­такта с некоторыми натуральными или синтетическими веществами. Целью представленного обзора является освеще­ние обоих типов механизмов формирования устойчивости растений к патогенам и прочим стрессорам
first_indexed 2025-12-07T16:05:23Z
format Article
fulltext Mech a nisms of plant in nate im mu nity N.O. Kozyrovska The In sti tute of Mo lec u lar Bi ol ogy and Ge net ics, NAS of Ukraine 150, Ac a de mi cian Zabolotny Str, Kyiv, 03143, Ukraine Plant in nate im mu nity is as sured by both con sti tu tive and in duced mech a nisms. Con sti tu tive bar rier for patho gens re lies on the plant cell wall struc ture, and the va ri et ies of in duc ible sys temic re sis tance re sult from in ter ac tion of the plant with patho - genic necrotrophic mi cro or gan isms, non patho genic bac te ria, and also af ter the con tact with some nat u ral or syn thetic mat - ters. De scrip tion of both mech a nisms of plant sys temic re sis tance to patho gens and other stress ors is the pur pose of the re view. Keywords: plant in nate im mu nity, sys temic ac quired re sis tance, in duced sys temic re sis tance Plant in nate im mu nity is strik ingly sim i lar to de fense sys tem of ver te brates and in sects ac cord ing to or ga ni za tion prin ci ples and mo lec u lar mech a nisms, which lie in the ba - sis of the re sponse to ex ter nal fac tor (patho gen) and, trust - wor thily, is the evolutionarily old sys tem of host de fense from patho gen [1]. As well as other higher or gan isms, plants are ca pa ble of rec og niz ing surficial struc tures of mi - cro or gan isms or the elicitors of plant de fense sys tem. The plants have the re cep tors sim i lar to Toll an i mal pro teins, which rec og nize patho gen [2]. Be sides, both an i mals and plants have sim i lar sig nal cas cades that ac ti vate im mu nity re spon si ble genes tran scrip tion. There fore, ni tric ox ide and pro tein kinase cas cades ac ti va tion in flu ence the de - fense re sponse in all Metazoa rep re sen ta tives, and, as a con se quence, the syn the sis of antimicrobial sub stances takes place [2, 3]. There are also dif fer ences in im mune sys tem or ga ni za - tions of plants and an i mals. First of all, plants do not have im mune cells like B-lym pho cytes, which rec og nize the patho gen, as in plant or gan ism ev ery cell has to do it it self. Sec ondly, plants have spe cial pro tec tion pro grams, which oc cur due to the abil ity of some sorts of plants to rec og nize spe cific vir u lence fac tors of some mi cro or gan isms [1]. Thirdly, plants have chan nels for “mes sages” transduction from the place of patho gen at tack. Such in for ma tion spread ing most likely is am pli fied by bac te ria and vi ruses, which are con stantly pres ent in side plant tis sues. Prior to de scrib ing plant im mu nity for ma tion mech a nisms proper, 91 ISSN 0233-7657. Biopolymers and cell. 2006. Vol. 22. ISS 2. Translated from Ukrainia. ãN.O. Kozyrovska, 2006 Reviews let us re vise ba sic terms and def i ni tions that are used in the ma te rial set forth be low. Patho genic mi cro or gan isms are de ter mined to be vir u - lent, if they cause symp toms of the dis ease in sen si tive plants, or avirulent, if they en able de fense re ac tion of plants and block the patho logic pro cess, as a con se quence. Plant patho genic Gram-neg a tive bac te ria Pseu do mo nas syringae, Erwinia carotovora, E. amylovora, Pantoea stewartii and many oth ers ac ti vate plant im mune sys tem through the in ser tion of the effector pro teins (vir u lence fac tors) into the plant cell, us ing con ser va tive type III se - cre tory sys tem (TTSS) [4]. Pili-trans port cor ri dors to ex - port vir u lence fac tors into the host cell are nec es sary for TTSS for ma tion [5]. TTSS and pili con trol the gene patho - ge nic ity clus ter (hrp), the prod ucts of which (harpins) are nec es sary for hy per sen si tiv ity re ac tion (HR) and for fast pro cess of pro grammed cell death (PCD) in the in fec tion lo cus to limit the patho gen spread ing into the host plant [6]. The genes, en cod ing TTSS effectors, are called the avirulence genes (avr). At the pres ence of cor re spond ing plant cell wall de fense R pro teins in re sis tant plant the prod ucts of the avr genes cause HR that en ables de fense re - ac tion cas cades, which re sults in patho gen re pro duc tion block ing in plant or gan ism [7]. The Avr pro tein causes dis ease only in ab sence of cor - re spond ing R pro tein pair, but the ex cep tions are also pos - si ble. Avirulent pro teins are suppressors of de fense pro teins of the first wave of de fence. The de ci pher ing ge nome of P. syringae pv. to mato (Pst) DC3000, dam ag ing to mato and Arabidopsis, showed that among 5763 open read ing frames there are not less than 298 of vir u lence genes, en cod ing about 50 effector TTSS pro teins and lo cal iz ing on mo bile ge netic el e ments [8]. Be side the TTSS effector genes, con - ser va tive effector locuses (CEL) were found, en cod ing the fam ily of con ser va tive type III effectors, like HopPtoM, AvrE etc in some spe cies of plant patho genic bac te ria [9-10]. These vir u lence fac tors sup press basal plant cell im mune sys tem in an other way, than those that were en - coded by the hrp genes [11]. There fore, the plant suc cess fully de fends it self only when it has the cor re spond ing R pro tein, which rec og nizes the sig nal, gen er ated by patho gen, more spe cif i cally, by a prod uct of its Avr gene. This con cep tion on plant de fense mech a nism was in tro duced by H.H. Flor in the 40s-50s of the last cen tury [12], and it be came the ba sis of “gene-for-gene” or “R-for-Avr” con cept. Study ing ge net - ics of a host–patho gen sys tem on the flax–rust model, the au thor de fined that ev ery rust tol er ance gene in host had the cor re spond ing fun gus patho ge nic ity gene. Later the R-Avr con nec tion was ex panded to other com bi na tions of plant-host–patho gen, where the lat ter could be rep re - sented by bac te ria, vi ruses, nem a todes, in sects etc [13]. Then it was as cer tained that men tioned gene prod ucts in - ter act di rectly as an ex cep tion, and in re al ity Avr pro tein in ter acts not with R pro tein but with so called guard of R pro tein, and only af ter this the for eign pro tein is rec og nized [14]. R genes that en code de fense R pro teins are im mensely struc tur ally di verse and clus tered in plants. This pro vides the in creased re com bi na tion prob a bil ity and, con se - quently, the ad ap ta tion to fast-chang ing genes of avirulence in patho genic bac te ria. The first R gene, which codes for tol er ance to Pst was cloned in to ma toes in 1998 [15]. At the pres ent mo ment it is known al ready that the ma jor ity of R pro teins which rec og nize mi cro bial patho ge - nic ity fac tors have got a com mon mo tive con sist ing of leucine-rich re peats (LRR). This el e ment plays a cer tain role in the pro tein-pro tein in ter ac tions, as well as dur ing sig nal transduction. R pro teins are di vided into sev eral groups; the big gest one is rep re sented by pro teins which have the nu cle o tide bind ing site (NBS) [16]. These pro - teins are spe cific for both prokaryotes and eukaryotes and they have com mon fea ture of bind ing with ATP or GTP for ac com plish ing their bi o log i cal func tions. NBS-LRR R pro teins, in their turn, are di vided into two sub groups de - pend ing on sec ond ary struc ture of the N-ter mi nal re gion. The first one com bines the pro teins that have super-coiled aminoterminal (CC-NBS-LRR sub group), the sec ond one – the pro teins which dif fer by the pres ence of do main, ho mol o gous do main of Drosophila Toll pro tein and interleukine re cep tor (Il-1R) of mam mals (TIR-NBS-LRR sub group). Fol low ing the anal ogy with men tioned pro teins, they play the sig nif i cant role in sig nal transduction in the plant, ac ti vat ing the de fense sys tem [17]. In Arabidopsis they are di vided into sev eral phylogroups, and num ber not less than 220 pro teins [18]. TIR do main is an evolutionarily old struc ture, and some plants, i.e. mono cot y le dons, have al ready lost it. The group of “clas si cal” NBS-LRR R- pro teins func tions in cytosol, at the same time, pro teins as so ci ated with the plant cell mem brane (LRR-RLK, LRR-RLP), play the role of re - cep tors [19]. Typ i cal re cep tor-like proteinkinases have the ex ter nal aminotherminal do main, for sig nal per cep tion, and cy to plas mic kinase do main (Ser/Thr) on the carboxyterminal, for sig nal transduction [20]. The sig nals from TIR- and CC-pro teins are trans - duced in dif fer ent ways, namely, the TIR-do main pro teins in duce the re sis tance via the gene-reg u la tor EDSI (en - hanced dis ease sus cep ti bil ity), whereas the group of CC-NBS-LRR pro teins re quires ex pres sion of NDR1 (non-race spe cific dis ease re sis tance) [17]. Some R pro - teins of NBS-LRR group re quire SGT1 pro tein 92 N.O. Kozyrovska (orthologue yeast pro tein ubiquinone-ligase) for their func tion ing [21]. Ob vi ously there are some other in de - pend ent ways of sig nal transduction in plants. Basal plant im mu nity re stricts de vel op ment of avirulent patho gen in sen si tive plants in the case of R gene ab sence. It pro vides cell sur face in teg rity which many patho gens can not over come. First of all, cu ti cle cov ers the leaf sur - face to pre vent patho gen pen e tra tion into aloplast, if they do not have cutinase. Sec ondly, a cell wall for ma tion out of strong poly mers (cel lu lose, pec tin) is a re li able bar rier for patho gens. In cor po rated antimicrobial pro teins, re leased at plant cell wall hy dro ly sis by patho gens, in crease strength of de fence [15]. The sen si tive plant has the de fense mech a - nisms, sim i lar to re sis tance, con di tioned by R-pro teins, though the pro cesses of this de fence pass slower and less ef - fec tively [22]. In plants that do not have the R gene to a cer tain patho gen, HR and PCD take place and the mech a - nisms sim i lar to sys temic ac quired re sis tance oc cur [23, 24]. Be sides that, the sen si tive plants in crease the phytohormone pro duc tion level [25, 26] and re pro duc tive de vel op ment as a re sponse to in fec tion to ob tain de scen - dants fast and to save the ge nus [27]. These events in plants are sim i lar to the ones which oc cur in stress con di tions, caused by abiotic fac tors [29]. Thus, due to basal sys tem of cell de fense the plant dis ease oc curs not al ways in case of the R-Avr pair ab sence. Non spe cific or nonhost re sis tance (NHR) of the plant is par tially con trolled by basal plant de fense sys tem. The mech a nisms of non spe cific re sis tance of the plants to the patho gens al low them to ex ist in the en vi ron ment and to evolve in the plant world re gard less of a wide patho gen spec trum. It has the sig nif i cant mean ing for both prac ti cal farm ing and ag ri cul ture in gen eral. From the fun da men tal re search point of view, un der stand ing NHR is nec es sary for host spec i fic ity phe nom ena and pathogenesis de ter - min ing in plants as such. The iden ti fi ca tion and de tailed char ac ter is tic of genes re spon si ble for NHR will pro vide the an swer to fol low ing ques tions: Is this re sis tance in - duced? Why is it not ef fec tive against vir u lent patho gens? What is the re la tion be tween NHR and host plant re sis - tance? It was dem on strated on ge netic pathosystem Arabidopsis–P. syringae pv. phaseolicola that the wild-type plant pro hib its the pseu do mo nas spread and there fore pre - vents the dis ease, whereas the mu tant, de fec tive by the gene NHO1 (nonhost re sis tance), sup ports the patho gen de vel op ment [29, 30]. This gene codes for glyc erol kinase, nec es sary for re sis tance to non patho genic and avirulent bac te ria. Thus, its ac tiv ity against vir u lent patho gens is not enough and it is not in duced by them dur ing the in fec tious pro cess, i.e. NHO1 is in duced by the bac te rium, for which the plant is not the host. It is sup posed that the gene in duc - tion is me di ated by mo lec u lar struc tures, lo cated on the sur face of mi cro or gan isms, so called PAMP (Patho - gen-As so ci ated Mo lec u lar Pat tern), rep re sented by peptidoglycans, lipoteichoic acid of Gram-pos i tive bac te - ria and liposaccharides and flagella of Gram-neg a tive bac - te ria [2, 31]. Typ i cal PAMP-struc ture rep re sen ta tive is flagellin, the ba sic pro tein of flagella used by bac te ria for move ment. The N-ter mi nal of pro tein com prises 22 aminoacid re sid u - als (flg 22), and its struc ture is one of the most con ser va tive among plant patho genic bac te ria and acts as a de fense elicitor [32]. The plant rec og nizes flagellin due to transmembrane re cep tor-kinase (MAPKKK, mitogen ac - ti vated kinase kinase kinase), and also by other sig nal sys - tem com po nents, e.g. tran scrip tion fac tors [33]. In Arabidopsis cas cade of MAPKinases (MEKK1, MPK4/MPK5 and MPK3/MPK6), which act af ter pro - cess ing leaf cell sus pen sion with flagellin and rec og niz ing by the re cep tor, was as cer tained com pletely [34]. In gen - eral, 23 pu ta tive MAPK, 10 MAPKK and more than 20 MAPKKK were dis cov ered in Arabidopsis ge nome [35]. Be sides patho gens, MAPKinases are ac ti vated by hor - mones, as well as by abiotic fac tors, which stress the plants. Mind ing the sig nif i cant num ber of dis cov ered kinase genes, the im por tant role of MAPK in cell pro cess reg u la - tion is sup posed. For ex am ple, knock out of one of such to - bacco gene, NPK1, ob structed R gene ac tiv ity dur ing the vi ral in fec tion, and also led to dwarf phe no type. Spe cific MAPKs (e.g. MPK6) are the pos i tive reg u la tors of ex pres - sion of some genes, which con di tion plant re sis tance to pri - mary in fec tion by cer tain patho gens, i.e. take part in basal de fense sys tem, whereas ac quired sys temic re sis tance in plants is me di ated by other MAPKinases [36]. Nonhost and avirulent patho gens (more pre cise, their PAMP, vol a tile bac te rial ex cre tion etc) are rec og nized by plant cell wall re cep tors. The lat ter trans mit risk sig nals through pro tein kinase cas cades into the cell, where af ter all HR and lo cal PCD would take place. The sig nals, which are trasduced by kinase cas cade, lead to bio chem i cal trans - for ma tions in the cell, which is at tacked by patho genic mi - cro or gan ism. The first sign of these trans for ma tions is the ox i da tive burst [37, 38]. As a con se quence, re ac tive ox y gen spe cies (ROS) like superoxide an ion (O-2), which trans - forms fast into hy dro gen per ox ide, are pro duced. Ni tric ox ide is also gen er ated [40]. Both pro cesses cause the pro - duc tion of toxic sub stances by the plant for self-pro tec tion from patho gen. On the other hand, they are sig nal mol e - cules in plant basal de fense sys tem, and, in their turn, pro - voke the for ma tion of new sig nal com po nents. ROS and ni tric ox ide re pro gram transcriptional events in the cell, 93 Mech a nisms of plant in nate im mu nity which re sult in the syn the sis of sig nal in ter me di ates, namely sal i cylic acid (SA), eth yl ene (ET) or jasmonic acid (JA), in HR and PCD, antimicrobial sub stances syn the sis, cell wall re in force ment, ac ti va tion of de fense genes, en - cod ing PR (patho ge nic ity re lated) and other pro teins [41]. It is proved in many re searches that SA plays the main role in the ac ti va tion of lo cal re sponse to patho genic bac te ria and oomycetes [42-46]. SA takes part in co-reg u la tion of plant de fense ways de pend ent on ET and JA [25, 47]. The lat ter hor mones me di ate de fense mech a nism against necrotrophic fungi through defensin and thionine in duc - tion [48], ac ti vate the en zymes in volved in phytoalexin syn the sis [51, 52], in duce sys temic re sis tance in plants by some non patho genic rhizobacteria [53-55]. The link, which joints the sig nals that go from reductive-ox i da tive pro cesses with the sig nals, trans duced by kinase cas cades, is OXI1-kinase. It is in duced in a re - sponse to H2O2 for ma tion and in its turn drives MPK3 and MPK6, i.e. is a sig nif i cant fac tor of sig nal transduction on the way to HR [56]. The sig nals from H2O2 to wards PCD are trans duced through the in ter ac tion be tween pro teins also on a rea son of the Ca+2 level changes. At the same time, other events take place in the cell. Cell wall thick ens be - cause of syn the sis of pro teins, rich in hydroxyproline, calose de po si tion (b-gluconic poly mer) and papillae for - ma tion, and these can not be over come by non-spe cific patho gens. The cell syn the sizes antimicrobial sub stances – de fense pro teins (defensins), phytoalexins, and PRproteins. Defensins are struc tur ally sim i lar to de fense pro teins of in sects, e.g. drosomycin, and are sim i lar to antimicrobial sub stances of ver te brates, and their ex pres - sion is con trolled by plant hor mones. Phytoalexins are low-mo lec u lar sub stances, ex creted by un af fected cells on the bor der with tis sues, af fected by patho gens or pests (by in sects in par tic u lar, but not by biotrophs). PR pro teins are dif fer ent in their struc ture and have gen er ally antifungal ac tiv ity (gluconases, chitinases), and some other fea tures among which is the ca pa bil ity to cryoprotection. The PR pro teins may ini ti ate so called sec ond wave of im mune re - sponse, rec og niz ing sig nal mol e cules of its own. For ex am - ple, glucanes (glucanase ac tiv ity prod ucts of sep a rate PR pro teins) are the de fense re sponse elicitors and thus, they in di rectly in duce antimicrobial ac tiv ity of plants [15]. Basal de fense sys tem of plants is the first stage in the plant-patho gen con test, and it is ac ti vated or in ac ti vated pe ri od i cally by patho gens on ei ther SA-de pend ent, or SA-in de pend ent ways. It also hap pens that the plants, treated by the elicitor only, may have dif fer ent re sponses, de pend ing on its na ture. Thus, Arabidopsis and to bacco plants, treated by vir u lence de ter mi nants of E. carotovora subsp. carotovora, gen er ate the im mune re sponse in dif fer - ent ways – on polygalacturonase - through the me di a tion of JA and ET, to harpin (HrpN) – SA, JA and ET [57]. There is a way of sig nal transduction from patho gen to plant, de pend ent on abscisic acid [58], and it is quite pos si - ble that the search of new ways of sig nal transduction is not fin ished yet. Fig.1 il lus trates the func tion ing scheme of Arabidopsis basal de fense sys tem af ter in fec tion by P. syringae pv. to mato DC3000. In the be gin ning of the 20th cen tury two sci en tists, Buverie and Ray, sep a rately from each other, came to the con clu sion that the in fec tion of plants by patho gens led to for ma tion of plant re sis tance in the case of re peated in fec - tion. In 30 years, Ches ter fi nal ized the phe nom e non of ac - quired sys tem re sis tance of plants, and later (in the be gin - ning of the 90s of the last cen tury) sys tem atic study on dif - fer ent types of in duced re sis tance of plants started [59]. Sys temic ac quired re sis tance (SAR) ap pears as a re sult of lo cal re ac tion to necrotrophic patho gen. At the same time the plant ac quires re sis tance to the sec ond at tack of the same patho gen or other classes of patho gens, and not only lo cally but ev ery where and [60-62]. In the pro cess of cell de fence PR genes are ac ti vated on cer tain dis tance from the place of pri mary plant-host in fec tion. SA ac cu - mu la tion is oblig a tory for SAR in duc tion [46, 63]. Trans - gen ic plants, which ex press the bac te rial gene nahG (codes for salicylatehydrolase, which trans forms SA into catechol) can not be re sis tant to patho gen in fec tions [42, 64]. Be sides necrotrophs, SAR in duc tion is caused by cer - tain con cen tra tions of exogenically added SA or its func - tional an a logues [65-69]. In the case of ac quir ing sys temic re sis tance, PR gene and some other gene ex pres sion takes place. The fea ture of SAR is the ap pear ance of HR state and ev ery thing con nected to it (see above), but it is more pro nounced. This im por tant cell de fense mech a nism re - quires the NPR1 (non-expressor of patho ge nic ity re lated genes) to reg u late sig nal transduction, pass ing from SA [70]. The mu tant npr1 ac cu mu lates nor mal SA level af ter in fec tion by patho gen, but it is un able to ex press PR genes and to form SAR [71, 72]. Overexpression of NPR1 causes the re sis tance of plants to both plant patho genic bac te ria and fungi [73], how ever, it is not needed in de fence against vi ruses. The NPR1 gene en codes the pro tein of two do - mains, which pro vide pro tein-pro tein in ter ac tions. Dur ing SAR, NPR1 prod uct is lo cal ized in the nu cleus, where it ac ti vates PR gene pro mot ers in a phys i cal con tact with some of TGA tran scrip tion fac tor. The NPR1 gene ex pres - sion is reg u lated by pro teins such as WRKY transcriptional fac tor which binds with DNA, rec og niz ing W-box of the pro mot ers [75-79]. NPR1 par tic i pa tion in de fense re - sponse in the plant cell de pends on both the type of patho - gen and the type of avirulence pro teins, which get into the 94 N.O. Kozyrovska cell through TTSS. For ex am ple, when AvrB of patho gen bac te rium pen e trates to the cell, the in ter ac tion with RPM1 pro tein (ac cord ing to “gene-for-gene” type) and fast HR re gard less of NPR1 take place. If the bac te rium at - tacks the plant by the effector, like AvrRpt2, then it is rec - og nized by RPS2 pro tein and af ter rec og ni tion ei ther fast SA ac cu mu la tion, , or slow ac cu mu la tion of HR- as so ci - ated sig nals takes place, and this de pends on what genes, NPR1 or NDR1 will be ac ti vated [11, 14]. In cytosol NPR1 plays an im por tant role, namely it is the role of the me di a tor be tween SA- and JA-de pend ent ways of plant de fense [80]. What hap pens in cytosol in par - tic u lar re mains un known, though it is sup posed that NPR1 ei ther sup presses the JA-de pend ent gene reg u la tor or de - liv ers the neg a tive reg u la tor to the nu cleus [47]. The Arabidopsis mu tants, which con sti tu tively ac cu mu late SA (cpr1 or cpr5) do not re quire “ex ter nal” prim ing, they are primed con stantly by the cell, but the NPR1 prod uct is not re quired for this [81]. Plant spe cific transcriptional fac tor WRKY70 is the con nect ing link be tween SA- and YA- or ET- me di ated ways of plant de fense. It ac ti vates the genes, which are in duced in a re sponse to SA, and re presses the genes de pend ent on YA. Overexpression of WRKY70 in - creases plant re sis tance level through NPR1-SA-mech a - nism, and the NPR1 gene antisense-sup pres sion leads to ac ti va tion of an other mech a nism, con di tioned by JA and eth yl ene [82]. Neg a tive reg u la tor of SAR is the EDR1 (en - hanced dis ease re sis tance) gene, mu ta tion in which does not lead to con sti tu tive plant de fense re sponse, and this re - sponse is primed [83]. The mech a nisms of sig nal transduction through SA are not fi nally cleared out, though there is the in for ma tion that re cep tor for SA is SABP2 hydrolase with lipase ac tiv ity [84]. The rep re sen ta tives of б/в-hy dro las es are known as the ones that take part in sig nal transduction, me di ated by hor mones, and there fore SABP2 func tion con sists in methyl group re moval from methyl salicylate, re leas ing SA, and in sig nal in duc tion in the form of lipid and its de - riv a tives [85, 86]. Hence forth, the sig nals are trans duced by pro teins sim i lar to VADI (vas cu lar as so ci ated death) [87]. Cer tain con cen tra tions of ex og e nously in tro duced SA, which in duce SAR in Arabidopsis, lead to ex pres sion in - crease of at least 12 genes in the plant. One of the func - tional groups of these genes is in volved in cell de fence (they en code glycosile transferase, glutathione-S-transferase etc). These are fast re ac tion genes, and their role con sists in pro vid ing an ti ox i dant and detoxication func tions in the plant. The other group of genes par tic i pates in sig nal transduction they code for pro tein kin ases and transcriptional fac tors) and re quire NPR1 for their in duc - tion [88]. This gen er ally com pli cated net work of SA-, JA- and ET-de pend ent ways of plant de fense from patho gens and abiotic stress ors has rather clear re ac tion to ex ter nal fac tors and de fines the range and power of de fense re ac tion. For ex am ple, both SA and JA are nec es sary for NPR1-in de - pend ent Arabidopsis de fense from P. syringae, Peronospora parasitica [89, 90]. In con trast, JA-de pend ent sig nal path com petes with SA-NPR1-de pend ent mech a nism, which re stricts the de vel op ment of P. syringae [91]. SA ad di tion leads to sup pres sion of JA- and ET- me di ated plant sig nal sys tem, which tes ti fies to the pri or ity of SA- NPR1-de - pend ent way of plant de fense. [80]. On the other hand, at NPR1 ab sence con sti tu tive ex pres sion of PDF1.2 in ssi1 and cpr6-mu tants of Arabidopsis is in creased [89, 92]. Thus, there are sev eral vari ants, which the plant chooses for de fense in or der to stay healthy. Fig 2. shows pos si ble im mune re sponse ways de pend ing on patho gen va ri ety. It should be men tioned, that abiotic stress ors cause SAR in ac cor dance with SA-sig nal path anal o gously to how it is done by necrotrophic patho gens. Thus, in creased 95 Mech a nisms of plant in nate im mu nity Fig 1. Hy po thet i cal model of ac ti va tion-in ac ti - va tion of basal de fense sys tem dur ing in fect ing a sen si tive plant of Arabidopsis by bac te rium Pst DC3000 [11]: 1- de fec tive bac te rium, which has a mu ta tion in the hrp genes clus ter, ac ti vates the basal sys tem in SA-in de pend ent way, namely – through PAMP, flagellin, is rec og nized by kinase do main of R pro tein-re cep tor FLS2 and trans - mits sig nals through MPK3/MPK6 cas cade; 2 - CEL mu ta tions in ac ti vate SA-in de pend ent sys - tem with the help of AvrPto effectors but the plant cell evades par tially such a pathogene ma - neu ver and ac ti vates SA-de pend ent de fense sys - tem; 3 - HopPtoM, AvrE-effectors in ac ti vate SA-de pend ent de fense scheme and cause cell heavy metal con cen tra tions lead to the in crease of both SA level and cor re spond ing me tab o lites in volved in this sig nal path in hyperaccumulating plants, and, as a con se quence, ul ti mately the re sis tance to some patho gens is formed [93]. In duced sys temic re sis tance (ISR), caused by non-patho genic rhizobacteria, is phenotypically sim i lar to ac quired sys temic plant im mu nity [94, 95]. Among rhizobacteria the most ac tive ones are the Pseu do mo nas rep re sen ta tives, that sup press patho gens in de pres sive soils, first of all, through the ex cre tions of an ti bi ot ics, lytic en zymes, siderophores, as well as by means of com pet ing for sources of nu tri tion [96]. Be sides, Pseu do mo nas re duce the de vel op ment of dis eases in above ground plant parts through the mech a nism, which is me di ated by the plant it - self. For ex am ple, P. fluorescens CHA0 ac ti vates the de - fense mech a nism in to bacco, sim i lar to SAR due to own SA ex cre tion, which “starts” this mech a nism in the plant [97]. Other Pseu do mo nas rep re sen ta tives stim u late ISR in many plant va ri et ies, suc cess fully col o niz ing root sys tem, how ever, the pos i tive re sult com pletely de pends on com bi - na tions bac te ria – plant-host [98]. It is worth men tion ing also that the bac te ria dif fer by the ca pa bil ity to cause ISR even in side the ge nus [99]. It is pos si ble that the dif fer ence be tween them is con di tioned by the ca pa bil ity of the plant to rec og nize the bac te rium, and more pre cisely, some of its com po nents or ex cre tions. It is known that for for ma tion of ISR a met a bol i cally ac tive bac te rium is not re quired, i.e. cell mem brane com po nents, lipopolysaccharides, siderophores, an ti bi ot ics, flagella in duce ISR [31, 100]. Some other bac te ria, which im prove plant de vel op ment (Ba cil lus subtilis, B. amyloliquefaciens), ac ti vate the plant de fense sys tem through the vol a tile or ganic com pounds (VOC) sim i lar to 2,3-butanediol [101]. It is op por tunely to add that be sides the bac te rial VOC, cell de fense mech a - nism is ac ti vated by some other low-mo lec u lar or ganic com pounds, e.q. volicitin, which is pro duced by in sects [102]. Some mu tants that lose cer tain ISR multicomponent inductors, are still ca pa ble of caus ing plant sys temic re sis tance anyway. The sim i lar ity of ISR and SAR con sists in the fact that the plant be comes re sis tant to a wide range of patho gens af - ter the con tact with the bac te ria. For ex am ple, as well as the typ i cal patho gens, P. fluorescens WCS417r causes plant re sis tance to dif fer ent kinds of patho gens, such as fun gal root patho gen Fusarium oxysporum, oomycete leaf patho - gen P. parasitica, bac te rial plant patho gens Xanthomonas campestris pv. armoracie and Pst DC300 [99, 103]. ISR and SAR ef fec tive ness spec tra some times over lap, but any way the mech a nisms, which con di tion them, are dif fer ent. It be came ev i dent af ter the ex per i ments with trans gen ic plants (NahG), that re vealed ISR to F. oxysporum and Pst DC3000 with out the PR-1 and PR-2 gene ex pres sion (sig - nif i cant for SA-de pend ent sig nal transduction path) af ter their in oc u la tion with WCS417r [54, 104]. There fore, it was de ter mined that ISR is in duced in de pend ently from SA. Fur ther ex per i ments with plants, in sen si tive to JA and ET, al lowed sup pos ing that for ISR oc cur rence the sig nal transduction ways me di ated by JA and ET are nec es sary [105]. For ex am ple, eth yl ene plays an im por tant role in the sig nal transduction from the VOC, and the sig nal way, me - di ated by JA, is in duced in the re sponse to in fect ing by micorrhizal fungi or trichoderma [105, 106]. The clas sic ex am ple of syn er gism be tween the rhizobacteria in duced sig nal ways, de pend ent on eth yl ene and JA, is ex pres sion of the PDF1.2 gene, en cod ing defensine in Arabidopsis (Fig. 2). To un der stand whether ISR is con nected to the in - creased genes ac tiv ity which re act to JA and eth yl ene, there was the study on the ex pres sion of a se ries of Arabidopsis genes in re sponse to the plant col o ni za tion by WCS417r. It was found that none of the plant genes was upregulated af - ter the con tact with the bac te rium ei ther lo cally, or in the dis tance [107]. Thus, the con clu sion was made, that ISR is based on the sen si tiv ity to these hor mones [108]. De spite dif fer ent mech a nisms, both ISR and SAR de - pend on NPR1 [53, 109]. Al though in case of SAR it in - duces the ex pres sion of known PR genes, it is still un der ques tion, what oc curs dur ing ISR. To re veal hy po thet i cal genes, ac ti vated af ter the con tact of the plant with the rhizobacterium, tran scrip tion anal y sis of Arabidopsis root was con ducted af ter its col o ni za tion with WCS417r, and it was found that 98 genes had in creased ex pres sion, in par - tic u lar, the AtMYB72 gene, which codes for the tran scrip - 96 N.O. Kozyrovska Fig 2. Sig nal ways at the re sis tant Arabidopsis plant for ma tion in case of in fect ing by vi rus, patho gen or non-patho gen bac te ria, fungi [53, 80, 108]. tion fac tor [110], and by this gene knock-out-mu ta tion of Arabidopsis lost the ca pac ity of cre at ing ISR in re sponse to WCS417r. Be sides, it was shown that its reg u la tion oc curs in the way, me di ated by eth yl ene, and in de pend ently from SA and JA, but eth yl ene proper is not needed for its ac tiv - ity. It is in ter est ing that in the leaf all 8000 genes did not change the ex pres sion level, and it did not ex clude ISR reg - u la tion in post-trans la tion way. The mo bi li za tion of plant de fense forces for fast re ac - tion to the patho gen at tack was called prim ing [68]. Plant prim ing with non-patho gen bac te ria (as well as nat u ral or syn thetic chem i cal sub stances) quick ens the re ac tion of the cell and plant in gen eral to bac te rial, fun gal, and vi ral in fec tion as well as to other stresses. The works on prim ing the cell cul tures of pars ley and Arabidopsis by bac te ria al - lowed the con clu sion that prim ing is the main mech a nism of in duced sys temic re sis tance of plants [67, 103, 111, 112]. The prim ing plant cell de fence is more rea son able en - ergy-wise, than con sti tu tive de fense mech a nism, be cause the de fense re ac tion is needed for the plant only at the mo - ment of patho gen at tack or the in flu ence of abiotic fac tors. Be sides, per ma nent syn the sis of ac tive pro tein sub stances, in volved di rectly to the de fense pro gram may pre vent nor - mal me tab o lism in the cell. The au thor ex presses sin cere grat i tude to Pro fes sor R.I. Gvozdyak for crit i cal re marks. Н. А. Козыровская Механизмы природной иммунности растений Ðåçþìå Природная иммунность растений обеспечивается конститутивным и индуцироваными механизмами. Конститутивный механизм обусловлен строением клеточной стенки растения, а разновидности индуцированных – возникают после взаимодействия растения с патогенными некротрофными микроорганизмами, непатогенными бактериями, а также после контакта с некоторыми натуральными или синтетическими веществами. Целью представленного обзора является освещение обоих типов механизмов формирования устойчивости растений к патогенам и прочим стрессорам. Ключевые слова: природная иммунность растений, приобретенная системная резистентность. REF ER ENCES 1. Nurenberger T., Brun ner F., Kemmerling, Piater L. In nate im mu - nity in plants and an i mals: strik ing sim i lar i ties and ob vi ous dif fer - ences// Immunol. Rev.- 2004.- 198, N1.- P. 249-253. 2. Aderem A., Ulevitch R. Toll-like re cep tors in the in duc tion of the in nate im mune re sponse // Na ture.- 2000.- 406.- P. 782787. 3. Imler J.-L., Hoffmann J. A. Toll re cep tors in in nate im mu nity // Trends Cell Biol.- 2001.-11.- P. 304311. 4. Hueck C. J. Type II pro tein se cre tion sys tems in bac te rial of an i - mals and plants // Microbiol. Mol. Biol. Rev.- 1998.- 62.- P. 379-433. 5. Hauck P., Thilmony R., He S. Y. A Pseu do mo nas syringae type III effector sup presses cell wall-based extracellular de fense in sus cep - ti ble Arabidopsis plants // PNAS.- 2003.- 100.- P. 8577-8582. 6. Buell C. R., Joardar V., Lindeberg M., Selengut J., Paulsen I. T., Gwinn M. L., Dodson R. J., Deboy R. T., Durkin A. S., Kolonay J. F., Madupu R., Daugherty S., Brinkac L., Beanan M. J., Haft D. H., Nel son W. C., Davidsen T., Zafar N., Zhou L., Liu J., Yuan Q., Khouri H., Fedorova N., Tran B., Rus sell D., Berry K., Utterback T., Van Aken S. E., Feldblyum T. V., D’Ascenzo M., Deng W. L., Ramos A. R., Alfano J. R., Cartinhour S., Chatterjee A. K., Delaney T. P., Lazarowitz S. G., Mar tin G. B., Schnei der D. J., Tang X., Bender C. L., White O., Fra ser C. M., Collmer A. The com plete ge nome se quence of the Arabidopsis and to mato patho gen Pseu do mo nas syringae pv. to mato DC3000 // Proc. Natl. Acad..Sci. USA.- 2003.- 100.- P. 10181-10186. 7. Bogdanove A. J., Kim J. F., Wei Z., Kolchinsky P., Charkowski A. O., Conlin A. K., Collmer A., Beer S. V. Homology and func tional sim i lar ity of an hrp-linked patho ge nic ity lo cus, dspEF, of Erwinia amylovora and the avirulence lo cus avrE of Pseu do mo nas syringae pathovar to mato // Proc. Natl. Acad. Sci.USA. – 1998.- 95.- P. 1325-1330. 8. Gaudriault S., Malandrin L., Paulin J.P., Barny M. A. DspA, an es sen tial patho ge nic ity fac tor of Erwinia amylovora show ing homology with AvrE of Pseu do mo nas syringae, is se creted via the Hrp se cre tion path way in a DspB-de pend ent way // Mol. Microbiol.- 1997. – 26, N 5.- P. 1057-69. 9. He S.Y., Jin Q. The Hrp pilus: learn ing from flagella // Curr. Opin. Microbiol.- 2003.- 6.- P. 15-29. 10. Lee J., Klusener B., Tsiamis G., Stevens C., Neyt C., Tampakaki A. P., Panopoulos N. J., Noller J., Weiler E. W., Cornelis G. R., Mansfield J. W., Nurnberger T. HrpZ(Psph) from the plant patho - gen Pseu do mo nas syringae pv. phaseolicola binds to lipid bilayers and forms an ion-con duct ing pore in vi tro // Proc. Natl. Acad. Sci. USA.- 2001.- 98.- P. 289-294. 11. Debroy S., Thilmony R., Kwack Y.-B., Nomura K., He S. A fam - ily of con served bac te rial effectors in hib its sal i cylic acid-me di ated basal im mu nity and pro motes dis ease ne cro sis in plants // Proc. Natl. Acad. Sci. USA.- 2004.- 10.- P. 9927-9932. 12. Flor H. H. The com ple men tary geneic sys tems in flax rust // Adv. Genet.- 1956.- 8.- P. 29-54. 13. Flor H. H. Cur rent sta tus of the gene-for-geneconcept // Annu. Rev. Phytopathol.- 1971.- 9.- P. 275-296. 14. Mackey D., Belkhadir Y., Alonso J. M., Ecker J. R., Dangl J. L. Arabidopsis RIN4 is a tar get of the type III vir u lence effector AvrRpt2 and mod u lates RPS2-me di ated re sis tance // Cell.- 2003.- 112.- P. 379-389. 15. Veronese P., Ruiz M. T., Coca M. A., Hernandez-Lopez A., Lee H., Ibeas J. I., Damsz B., Pardo J. M., Hasegawa P. M., Bressan R. A., Narasimhan M. L.. In de fense against patho gens. Both plant 97 Mech a nisms of plant in nate im mu nity sen ti nels and foot sol diers need to know the en emy // Plant Physiol.- 2003.- 131. P. 1580-1590. 16. Dong J., Chen C., Chen Z. Ex pres sion pro files of the Arabidopsis WRKY gene superfamily dur ing plant de fense re sponse // Plant Mo lec u lar Bi ol ogy. –2003.-51, N1.- P. 21-37. 17. Shirano Y., Kachroo P., Shah J., Klessig D.F. A gain-of-func tion mu ta tion in an Arabidopsis Toll interleukin1 re cep tor–nu cle o tide bind ing site–leucine-rich re peat type R gene trig gers de fense re - sponses and re sults in en hanced dis ease re sis tance // Plant Cell.- 2002.- 14.- P. 3149-3162. 18. Fluhr R. Sen ti nels of dis ease. Plant re sis tance genes // Plant Physiol.- 2001.- 127.- P.1367-1374. 19.Verica J., Chae L., Tong H., Ingmire P., He Z. Tis sue-spe cific and de vel op men tally reg u lated ex pres sion of a clus ter of tandemly ar - rayed cell wall-as so ci ated kinase-like kinase genes in Arabidopsis // Plant Physiol.- 2003.- 133.- P. 1732-1746. 20. Ellis J., Dodds P., Pryor T. Struc ture, func tion and evo lu tion of plant desease re sis tance genes // Curr. Opin. Plant Biol.- 2000.- 3.- P. 279-284. 21. Peart J. R.., Lu R.., Sadanandom A., Malcuit I., Moffett P., Brice D. C., Schauser L., Jaggard D. A., Xiao S., Coleman M..J., Dow M., Jones J. D., Shirasu K., Baulcombe D. C. Ubiquitin ligase-as - so ci ated pro tein SGT1 is re quired for host and nonhost dis ease re - sis tance in plants // Proc. Natl. Acad. Sci. USA.- 2002.- 99.- P. 10865-10869. 22. Maleck K., Le vine A., Eulgem T., Mor gan A., Schmid J., Lawton K. A., Dangl J. L., Dietrich R. A. The transcriptome of Arabidopsis thaliana dur ing sys temic ac quired re sis tance // Nat. Genet.- 2000.- 26.- P. 403-410. 23. Yang Y. O., Shah J., Klessig D. F. Sig nal per cep tion and transduction in de fense re sponses // Genes Dev.- 1997.- 11.- P. 1621-1639. 24. Glazebrook J. Genes con trol ling ex pres sion of de fense re sponses in Arabidopsis: 2001 sta tus // Curr. Opin. Plant Biol.- 2001.- 4.- P. 301-308. 25. Dong X. N. SA, JA, eth yl ene, and dis ease re sis tance in plants // Curr. Opin. Plant Biol.- 1998.- 1.- P. 316-323. 26. Lund S. T., Stall R. E., Klee H. J. Eth yl ene reg u lates the sus cep ti - ble re sponse to patho gen in fec tion in to mato // Plant Cell.- 1998.- 10.- P. 371-382. 27. Korves T., Bergelson J. A de vel op men tal re sponse to patho gen in - fec tion in Arabidopsis // Plant Physiol.- 2003.- 133.- P. 339-347. 28. Chen W., Provart N., Glazebrook J., Katagiri F., Chang H., Eulgem T., Mauch F., Luan S., Zou G., Whitham S., Budworth P., Tao Y., Xie Z., Chen X., Lam S., Kreps J., Harper J., Si-Ammour A., Mauch-Mani B., Heinlein M., Kobayashi K., Hohn T., Dangl J., Wang X., Zhu T. Ex pres sion pro file ma trix of Arabidopsis tran scrip tion fac tor genes sug gests their pu ta tive func - tions in re sponse to en vi ron men tal stresses // Plant Cell. – 2002.- 14.- P. 559-574. 29. Ming L., Tang X., Zhou J. -M. Arabidopsis NHO1 is re quired for gen eral re sis tance against Pseu do mo nas bac te ria // Plant Cell.- 2001.- 13.- P. 437-447. 30. Kang L., Li J., Zhao T., Xiao F., Tang X., Thilmony R., He S., Zhou J.- M. In ter play of the Arabidopsis nonhost re sis tance gene NHO1 with bac te rial vir u lence // Proc. Natl. Acad. Sci. USA.- 2003.- 100.- P. 3519-3524. 31. Lee man M., Van Pelt J. A., Den Ouden F. M., Heinsbroek M., Bakker P. A. H. M., Ship pers B. In duc tion of sys temic re sis tance against fusarium wilt of rad ish by lipopolysaccharides of Pseudommonas fluorescens // Phytopathology.- 1995.- 85.- P. 1021-1027. 32. Fe lix G., Duran J. D., Volko S., Boller T. Plants have a sen si tive per cep tion sys tem for the most con served do main of bac te rial flagellin // Plant J.- 1999.- 18.- P. 265-276. 33. Navarro L., Zipfel C., Rowland O., Keller I., Robatzek S., Boller T., Jones J. The transcriptional in nate im mune re sponse to flg22. In ter play and over lap with Avr gene-de pend ent de fense re sponses and bac te rial pathogenesis // Plant Physiol.- 2004.- 135.- P. 1113-1128. 34. Asai T., Tena G., Plotnikova J., Willmann M. R., Chiu W. L., Gomez-Gomez L., Boller T., Ausubel F. M., Sheen J. MAP kinase sig nal ling cas cade in Arabidopsis in nate im mu nity // Na ture.- 2002.- 415.- P. 977-983. 35. Jin H., Axtel M., Dahlbeck D., Ekwenna O., Zhang S., Staskawicz B., Baker B. NPK1, an MEKK1-like mitogen-ac ti - vated pro tein kinase kinase, reg u lates in nate im mu nity and de vel - op ment in plants // De vel op men tal Cell.- 2002.- 3.- P. 291-297. 36. Menke F., van Pelt J. A., Pieterse C. M. J., Klessig D. F. Si lenc ing of the mitogen-ac ti vated pro tein kinase MPK6 com pro mises dis - ease re sis tance in Arabidopsis // Plant Cell.- 16.- 2004.- P. 897-907. 37. Gechev T. S., Hille J. Hy dro gen per ox ide as a sig nal con trol ling plant pro grammed cell death // J. Cell Biol.- 2005.- 168.- P. 17-20. 38. Doke N., Miura Y., Sanchez L. M., Park H. J., Noritake T., Yoshioka H., Kawakita K. The ox i da tive burst pro tects plants against patho gen at tack: mech a nism and role as an emer gency sig - nal for plant bio-de fence - a re view // Gene.- 1996.- 179.- P. 45-51. 39. Newman M. A., von Roepenack-Lahaye E., Parr A., Daniels M. J., Dow J. M. Prior ex po sure to lipopolysaccharide po ten ti ates ex - pres sion of plant de fenses in re sponse to bac te ria // Plant J.- 2002.- 29.- P. 487-495. 40. Zeidler D., Zдhringer U, Gerber I., Dubery I., Hartung T., Bors W., Hutzler P. , Durner J. In nate im mu nity in Arabidopsis thaliana: Lipopolysaccharides ac ti vate ni tric ox ide synthase (NOS) and in duce de fense genes // Proc. Natl. Acad. Sci. USA.- 2004.- 101.- P. 15811-15816. 41. de Torres M., Sanchez P., Fernandez-Delmond I., Grant M. Ex - pres sion pro fil ing of the host re sponse to bac te rial in fec tion: the tran si tion from basal to in duced de fence re sponses in RPM1-me - di ated re sis tance // The Plant Jour nal.- 2003.- 33.- P. 665-676. 42. Ryals J. A., Neuenschwander U. H., Willits M. G., Molina A., Steiner H. Y., Hunt H. Y. Sys temic ac quired re sis tance // Plant Cell.- 1996.- 8.- P. 1809-1819. 98 N.O. Kozyrovska 43. Durner J., Shah J., Klessig, D. F. Sal i cylic acid and dis ease re sis - tance in plants // Trends Plant Sci.- 1997.- 2.- P. 266-274. 44. Demp sey D., Shah J., Klessig D. F. Sal i cylic acid and dis ease re - sis tance in plants // Crit. Rev. Plant Sci.- 1999.- 18.- P. 547-575. 45. O’Donnell P. J., Jones J. B., Antoine F. R., Ciardi J., Klee H. J. Eth yl ene-de pend ent sal i cylic acid reg u lates an ex panded cell death re sponse to a plant patho gen // Plant J.- 2001.- 25.- P. 315-323. 46. Gaffney T., Friedrich L., Vernooij B., Negrotto D., Nye G., Uknes S., Ward E., Kessmann H., Ryals J. Re quire ment of sal i - cylic acid for the in duc tion of sys temic ac quired re sis tance // Sci - ence.- 1993.- 261.- P. 754-756. 47. Pozo M. J., Van Loon L. C., Pieters C. M. J. Jasmonates – sig nals in plant-mi crobe in ter ac tions // J. Plant Growth Regul.- 2005.- 23.- P. 211-222. 48. Knoester M., Van Loon L.C., Heuvel J.V.D., Hennig J., Bol J.F., Linthorst H.J.M. Eth yl ene-in sen si tive to bacco lacks nonhost re - sis tance against soil-borne fungi // Proc. Natl. Acad. Sci. USA.- 1998.- 95.- P.1933-1937. 49. Penninckx I. A. M. A., Eggermont K., Terras F. R. G., Thomma B. P. H. J., De Samblanz G. W., Buchala A., Mйtraux J.-P., Man - ners J. M., Broekaert W. F. Patho gen-in duced sys temic ac ti va tion of a plant defensin gene in Arabidopsis fol lows a sal i cylic acid–in - de pend ent path way // Plant Cell.- 1996.- 8.- P. 2309-2323. 50. Epple P., Apel K., Bohlmann H. An Arabidopsis thaliana thionin gene is in duc ible via a sig nal transduction path way dif fer ent from that for pathogenesis-re lated pro teins // Plant Physiol.- 1995.- 109.- P. 813-820. 51. Boller T., Gehri A., Mauch F., Vцgeli U. Chitinase in bean leaves: in duc tion by eth yl ene, pu ri fi ca tion, prop er ties, and pos si ble func - tion // Planta.- 1983.- 157.- P. 22-31. 52. Mauch F., Staehelin L. A. Func tional im pli ca tions of the subcellular lo cal iza tion of eth yl ene-in duced chitinase and Я-1,3-glucanase in bean leaves // Plant Cell.- 1989.- 1.- 447-457. 53. Pieterse C. M. J., Van Wees S. C. M., Van Pelt J. A., Knoester M., Laan R., Gerrits H., Weisbeek P. J., Van Loon L. C. A novel sig - nal ing path way con trol ling in duced sys temic re sis tance in Arabidopsis // Plant Cell.- 1998.- 10.- P. 1571-1580. 54. Pieterse, C. M. J., Van Loon, L. C. Sal i cylic acid–in de pend ent plant de fense path ways // Trends Plant Sci.- 1999.- 4.- P. 52-57. 55. Ivacoli A., Boutet E., Metraux J. P. In duced sys temic re sis tance in Arabidopsis thaliana in re sponse to root in oc u la tion with Pseu - do mo nas flu o res cence CHA0 // Mol. Plant Mi crobe In ter act.- 2003.- 16.- P. 851-858. 56. Rentel M. C., Lecourieux D., Ouaked F., Usher S. L., Petersen L., Okamoto H., Knight H., Peck S. C., Grierson C. S., Hirt H., Knight M. R. OXI1 kinase is nec es sary for ox i da tive burst- me di - ated sig nal ling in Arabidopsis // Na ture.- 2004.- 427, N6977- P. 858-861. 57. Kariola T., Palomaki T. A., Brader G., Palva E. T. Erwinia carotovora subsp. carotovora and Erwinia-de rived elicitors HrpN and PehA trig ger dis tinct but in ter act ing de fense re sponses and cell death in Arabidopsis // Mol. Plant Mi crobe In ter act.- 2003.- P. 179-187. 58. Ton J., Mauch-Mani B. Beta-amino-bu tyric acid-in duced re sis - tance against necrotrophic patho gens is based on ABA-de pend ent prim ing for callose // Plant J.- 2004.- 38.- P. 119-130. 59. Conrath U., Thulke O., Katz V., Schwindling S., Kohler A. prim - ing as a mech a nism in in duced sys temic re sis tance of plants // Eu - ro pean J. Plant Pathol.- 2001.- 107.- P. 113-119. 60. Metraux J.-P., Nawrath C, Genoud T. Sys temic ac quired re sis - tance // Euphytica.- 2002.- 124.- P. 237–243. 61. Sticher L., Mauch-Mani B., Metraux J.-P. Sys temic ac quired re - sis tance. Annu. Rev. Phytopathol.- 1997.- 35.- P. 235-270. 62. Uknes S., Mauch-Mani B., Moyer M., Pot ter S, Wil liams S., Dincher S., Chandle D., Slusarenko S., Ward S., Ryals J. Ac - quired re sis tance in Arabidopsis // Plant Cell.- 1992.- 4.- P. 645-656. 63. van Loon L. C., van Strien E. A. The fam i lies of pathogenesis-re - lated pro teins, their ac tiv i ties, and com par a tive anal y sis of PR-1 type pro teins // Physiol. Molec. Plant Pathol.- 1999.- 55.- P. 85–97. 64. Nawrath C., Heck S., Parinthawong N., Metraux J. -P. EDS5, an es sen tial com po nent of sal i cylic acid-de pend ent sig nal ing for dis - ease re sis tance in Arabidopsis, is a mem ber of the MATE trans - porter fam ily // Plant Cell.- 2002.- 14.- P. 275-286. 65. Zimmerli L., Jakab G., Metraux J.-P., Mauch-Mani B. Potentiation of patho gen-spe cific de fense mech a nisms in Arabidopsis by beta -aminobutyric acid // Proc. Natl. Acad. Sci. USA.- 2000.- 97.- P. 12920-12925. 66. Ton J., Jakab G., Toquin V., Flors V., Iavicoli A., Maeder M. N., Mйtraux J.-P., Mauch-Mani B. Dis sect ing the Я-aminobutyric acid–in duced prim ing phe nom e non in Arabidopsis // Plant Cell.- 2005.- 17.- P. 987-999. 67. Katz V. A., Thulke O. U., Conrath U. A benzothiadiazole primes pars ley cells for aug mented elic i ta tion of de fense re sponses // Plant Physiol.- 1998.- 117.- P. 1333-1339. 68. Conrath U., Pieterse C. M., Mauch-Mani B. Prim ing in plant-patho gen in ter ac tions // Trends Plant Sci.- 2002.- 7.- P. 210-216. 69. Conrath U., Chen Z., Ricigliano J. R., Klessig D. F. Two in duc - ers of plant de fence re sponses, 2,6-dichloroisonicotinec acid and sal i cylic acid, in hibit catalase ac tiv ity in to bacco // Proc. Natl. Acad. Sci. USA.- 1995.- 92.- P. 7143-7147. 70. Kohler A., Schwindling S., Conrath U. Benzothiadiazole-in - duced prim ing for po ten ti ated re sponses to patho gen in fec tion, wound ing, and in fil tra tion of wa ter into leaves re quires the NPR1/NIM1 gene in Arabidopsis // Plant Physiol.- 2002.- 128.- P. 1046-1056. 71. Cao H., Bowl ing S. A., Gordon A. S., Dong X. Char ac ter iza tion of an Arabidopsis mu tant that is nonresponsive to in duc ers of sys - temic ac quired re sis tance // Plant Cell.- 1994.- 6.- P. 1583-1592. 72. Delaney T. P., Friedrich L., Ryals J. A. Arabidopsis sig nal transduction mu tant de fec tive in chem i cally and bi o log i cally in - duced dis ease re sis tance // Proc. Natl. Acad. Sci. USA.- 1995.- 92.- P. 6602-6606. 99 Mech a nisms of plant in nate im mu nity 73. Cao H., Li X., Dong X. Gen er a tion of broad-spec trum dis ease re - sis tance by overexpression of an es sen tial reg u la tory gene in sys - temic ac quired re sis tance // Proc. Natl. Acad. Sci. USA.- 1998.- 95.- P. 6531-6536. 74. Kachroo P., Yoshioka K., Shah J., Dooner H. K., Klessig D. F. Re sis tance to tur nip crin kle vi rus in Arabidopsis is reg u lated by two host genes and is sal i cylic acid de pend ent but NPR1, eth yl ene, and jasmonate in de pend ent // Plant Cell.- 2000.- 12.- P. 677-690. 75. Zhang Y., Fan W., Kinkema M., Li X., Dong X. In ter ac tion of NPR1 with ba sic leucine zip per pro tein tran scrip tion fac tors that bind se quences re quired for sal i cylic acid in duc tion of the PR-1 gene // Proc. Natl. Acad. Sci. USA.- 1999.- 96.- P. 6523-6528. 76. Kinkema M., Fan W., Dong X. Nu clear lo cal iza tion of NPR1 is re quired for ac ti va tion of PR gene ex pres sion // Plant Cell.- 2000.- 12.- P. 2339-2350. 77. Yu D., Chen C., Chen Z.. Ev i dence for an im por tant role of WRKY DNA bind ing pro teins in the reg u la tion of NPR1 gene ex - pres sion // Plant Cell.- 2001.- 13.- P. 1527-1540. 78. Fan W., Dong X. In vivo in ter ac tion be tween NPR1 and tran - scrip tion fac tor TGA2 leads to sal i cylic acid–me di ated gene ac ti - va tion in Arabidopsis // Plant Cell.- 2002.- 14.- P. 1377-1389. 79. Yu D., Chen C., Chen Z.. Ev i dence for an im por tant role of WRKY DNA bind ing pro teins in the reg u la tion of NPR1 gene ex - pres sion // Plant Cell.- 2001.- 13.- P. 1527-1540. 80. Spoel S. H., Koornneef A., Claessens S. M. C., Korzelius J. P., Van Pelt J. A., Mueller M. J., Buchala A. J., Mйtraux J.-P., Brown R., Kazan K., Van Loon L. C., Dong X., Pieterse C. M.. NPR1 Mod u lates cross-talk be tween salicylate- and jasmonate-de pend ent de fense path ways through a novel func tion in the cytosol // Plant Cell.- 2003.- 15.- P. 760-770. 81. Bowl ing S. A., Guo A., Cao H., Gordon A. S., Klessig D. F., Dong X.. A mu ta tion in Arabidopsis that leads to con sti tu tive ex - pres sion of sys temic ac quired re sis tance // Plant Cell.- 1994.- 6.- P. 1845-1857. 82. Li J., Brader G., Palva E. T. The WRKY70 tran scrip tion fac tor: a node of con ver gence for jasmonate-me di ated and salicylate-me di - ated sig nals in plant de fense // Plant Cell.- 2004.- 16.- P. 319-331. 83. Tang D., Innes R. W. Overexpression of a kinase-de fi cient form of the EDR1 gene en hances pow dery mil dew re sis tance and eth yl - ene-in duced se nes cence in Arabidopsis // Plant J.- 2002.- 32.- P. 975-983. 84. Kumar D., Klessig D. F. High-af fin ity sal i cylic acid-bind ing pro - tein 2 is re quired for plant in nate im mu nity and has sal i cylic acid-stim u lated lipase ac tiv ity // Proc. Natl. Acad. Sci. USA.- 2003.- 100.- P. 16101-16106. 85. Forouhar F., Yang Y., Kumar D., Chen Y., Fridman E., Park S. W., Chiang Y., Acton T. B., Montelione G., Pichersky E., Klessig D. F., Tong L. Struc tural and bio chem i cal stud ies iden tify to bacco SABP2 as a methyl salicylate esterase and im pli cate it in plant in nate im mu nity // Proc. Natl. Acad. Sci. USA..- 2005.- 102.- P. 1773-1778. 86. Maldonado A. M., Doerner P., Dixon R. A., Lamb C. J., Cameron R. K. A pu ta tive lipid trans fer pro tein in volved in sys - temic re sis tance sig nal ling in Arabidopsis // Na ture.- 2002.- 419.- P. 399-403. 87. Lorrain S., Lin B., Auriac M. C., Kroj K., Saindrenan P., Nicole M., Balaguй C., Roby D. VASCULAR ASSOCIATED DEATH1, a novel GRAM do main–con tain ing pro tein, is a reg u - la tor of cell death and de fense re sponses in vas cu lar tis sues // Plant Cell.- 2004.- 16.- P. 2217-2232. 88. Blanco F, Garreton V, Frey N, Dominguez C, Perez-Acle T, Van der Straeten D, Jordana X, Holuigue L. Iden ti fi ca tion of NPR1-de pend ent and in de pend ent genes early in duced by sal i - cylic acid treat ment in Arabidopsis // Plant Mol Biol. – 2005.- 59.- P. 927-944. 89. Jo seph D. Clarke, Sigrid M. Volko, Heidi Ledford, Fred er ick M. Ausubel, and Xinnian Dong Roles of Sal i cylic Acid, Jasmonic Acid, and Eth yl ene in cpr-In duced Re sis tance in Arabidopsis // Plant Cell 12: 2175-2190. 90. Nandi A., Kachuroo P., Fukushig H., Hildebrand D., Klessing D., Shah J. Eth yl ene and jasmonic acid sig nal ing path ways affectNPR1-in de pend ent ex pres sion of de fence genes with out im - pact ing re sis tance to Pseu do mo nas syringae and Peronospora parasitica in the Arabidopsis ssiI mu tant // Mol. Plant-Mi crobe In ter act.- 2003.- 16.- P. 588-599. 91. Kloek AP, Verbsky ML, Sharma SB, Schoelz JE, Vogel J, Klessig DF, Kunkel BN. The Pseu do mo nas syringae type III effector AvrRpt2 func tions down stream or in de pend ently of SA to pro mote vir u lence on Arabidopsis thaliana // Plant J. 2004 Feb;37(4):494-504. 92. Shah Jyoti Shah, Pradeep Kachroo, and Dan iel F. Klessig The Arabidopsis ssi1 Mu ta tion Re stores Pathogenesis-Re lated Gene Ex pres sion in npr1 Plants and Ren ders Defensin Gene Ex pres sion Sal i cylic Acid De pend ent // Plant Cell 11: 191-206 93. Free man J. L., Gar cia D., Kim D., Hopf A., Salt D. E. Con sti tu - tively el e vated sal i cylic acid sig nals glutathione-me di ated nickel tol er ance in Thlaspi nickel hyperaccumulators // Plant Physiol.-2005.- 137.- P. 1082-1091. 94. Van Peer R., Niemann G. J., Schippers B. In duced re sis tance and phytoalexin ac cu mu la tion in bi o log i cal con trol in Fusarium wiltof car na tion by Pseu do mo nas sp. strain WCS417r // Phytopathology.- 1991.- 81.- P. 728-734. 95. Wei G., Klopper J. W., Tuzun S. In duc tion of sys temic re sis tance of cu cum ber to Colleotrichum orbiculare by se lect strains of plant growth-pro mot ing rhizobacteria // Phytopathology.- 1991.- 81.- P. 1508-1512. 96. Lottmann J., Berg G. Phenotypic and genotypic char ac ter iza tion of an tag o nis tic bac te ria as so ci ated with roots of trans gen ic and non-trans gen ic po tato plants // Microbiol. Res. – 2001. – 156. – P. 75-82. 97. Maurhofer M., Reimmann C., Schmidli-Sacherer P., Heeb S., Haas D., Dйfago G. Sal i cylic acid biosynthetic genes ex pressed in Pseu do mo nas fluorescens strain P3 im prove the in duc tion of sys - temic re sis tance in to bacco against to bacco ne cro sis vi rus // Phytopathology.- 1998.- 88.- P. 678-684. 98. Bloemberg G. V., Lugterberg B. J. Mo lec u lar ba sis of plant growth pro mo tion and biocontrol by rhizobacteria // Curr. Opin. Plant Biol. – 2001. – 4. – P. 343-350. 99. van Wees S. C., Pieterse C. M., Trijssenaar A., Van’t Westende Y. A. M., Hartog F., van Loon L. C. Dif fer en tial in duc tion of sys - temic re sis tance in Arabidopsis by biocontrol bac te ria // Mol. Plant-Mi crobe In ter act.- 1997.- 10.- P. 710-716. 100 N.O. Kozyrovska 100. Schnider-Keel U., Seematter S., Maurhofer M., Blumer C., Duffy B., Gi got-Bonnefoy C., Reimmann C., Notz R., Dйfago G., Haas D., Keel C. Autoinduction of 2,4-Diacetylphloroglucinol Biosynthesis in the Biocontrol Agent Pseu do mo nas fluorescens CHA0 and Re pres sion by the Bac te rial Me tab o lites Salicylate and Pyoluteorin // J. Bacteriol.- 2000.- 182.- P. 1215-1225. 101. Choong-Min Ryu, Mohamed A. Farag, Chia-Hui Hu, Munagala S. Reddy, Jo seph W. Kloepper, and Paul W. Parй Bac - te rial Volatiles In duce Sys temic Re sis tance in Arabidopsis Plant Physiol. 134: 1017-1026; 102. Pare P. W., Farag M. A., Krishnamachari V., Zhang H., Ryu C. M., Kloepper J. W. Elicitors and prim ing agents ini ti ate plant de - fense re sponses // Photosynth Res. – 2005.- 85.- P. 149-59. 103. Van Loon L. C., Bakker P. A., Pieterse C. M. Sys temic re sis - tance in duced by rhizosphere bac te ria // Annu. Rev. Phytopathol.- 1998.- 36.- P. 453-483. 104. Pieterse C. M., van Wees S. C., Hoffland E., van Pelt J. A., van Loon L. C. Sys temic re sis tance in Arabidopsis in duced by biocontrol bac te ria is in de pend ent of sal i cylic acid ac cu mu la tion and pathogenesis-re lated gene ex pres sion // Plant Cell.- 1996.- 8.- P. 1225-1237. 105. Pozo MJ, Cordier C, Dumas-Gaudot E, Gianinazzi S, Barea JM, Azcon-Aguilar C. Lo cal ized ver sus sys temic ef fect of arbuscular mycorrhizal fungi on de fence re sponses to Phy toph - thora in fec tion in to mato plants // J Exp Bot. 2002 Mar;53(368):525-34. 106. Christelle Mar ti nez, Frйdйric Blanc, Emilie Le Claire, Olivier Besnard, Michel Nicole, and Jean-Claude Baccou Sal i cylic Acid and Eth yl ene Path ways Are Dif fer en tially Ac ti vated in Melon Cot - y le dons by Ac tive or Heat-De na tured Cellulase from Trichoderma longibrachiatum // Plant Physiol. 127: 334-344. 107. Ton J., Pieterse C. M., Van Loon L. C. Iden ti fi ca tion of a lo cus in arabidopsis con trol ling both the ex pres sion of rhizobacteria-me di ated in duced sys temic re sis tance (ISR) and basal re sis tance against Pseu do mo nas syringae pv. to mato // Mol. Plant Mi crobe In ter act.- 1999.- 12.- P. 911-918. 108. Pieters C. M. J, Van Loon L. C. NPR1: the spi der in the web of in duced re sis tance sig nal ling path ways // Curr. Opin. Plant Biol..- 2004.- 7.- P. 456-464. 109. Van Wees S. C., Luijendijk M., Smoorenburg I., van Loon L. C., Pieterse C. M. Rhizobacteria-me di ated in duced sys temic re sis - tance (ISR) in Arabidopsis is not as so ci ated with a di rect ef fect on ex pres sion of known de fense-re lated genes but stim u lates the ex - pres sion of the jasmonate-in duc ible gene Atvsp upon chal lenge // Plant Mol Biol.- 1999.-41.- P. 537-49. 110. Verhagen B. W., Glazebrook J., Zhu T., Chang H. S., van Loon L. C., Pieterse C. M. The transcriptome of rhizobacteria-in duced sys temic re sis tance in arabidopsis // Mol Plant Mi crobe In ter act.- 2004.- 17.- P. 895-908. 111. Van Loon L. C., Bakker P. A. H. M., Pieterse C. M. J. Sys temic re sis tance in duced by rhizobacteria // Annu. Rev. Phytopathol.- 1998.- 36.- P. 453-483. 112. Pieterse C. M. J, Van Wees S. C. M., Ton J., Van Pelt J. A., Van Loon L. C. Sig nal ling in rhizobacteria-in duced sys temic re sis tance in Arabidopsis thaliana // Plant Biol.- 2002.- 4.- P. 535-544. 101 Mech a nisms of plant in nate im mu nity
id nasplib_isofts_kiev_ua-123456789-156471
institution Digital Library of Periodicals of National Academy of Sciences of Ukraine
issn 0233-7657
language Ukrainian
last_indexed 2025-12-07T16:05:23Z
publishDate 2006
publisher Інститут молекулярної біології і генетики НАН України
record_format dspace
spelling Козировська, Н.О.
2019-06-18T14:30:19Z
2019-06-18T14:30:19Z
2006
Механізми природної імунності рослини / Н.О. Козировська // Біополімери і клітина. — 2006. — Т. 22, № 2. — С. 91-101. — Бібліогр.: 112 назв. — укр., англ.
0233-7657
DOI: http://dx.doi.org/10.7124/bc.000721
https://nasplib.isofts.kiev.ua/handle/123456789/156471
632.938
Природна імунність рослини забезпечується конститутивним та індукованими механізмами. Конститутивний механізм обумовлений будовою клітинної стінки рослини, а різновиди інду­кованих — виникають теля взаємодії рослини з патогенними некротичними мікрорганізмами, непатогенними бактеріями, а також після обробки деякими природними або синтетичними речовинами. Метою представленого огляду є висвітлення обох типів механізмів формування стійкості до патогенів та інших стресів.
Plant innate immunity is assured by both constitutive and induced mechanisms. The constitutive barrier for pathogens relies on the plant cell wall structure, and the varieties of inducible systemic resistance result from interaction of the plant with pathogenic necrotrophic microorganisms, nonpathogenic bacteria, and also after the contact with some natural or synthetic substances. Description of both mechanisms of plant systemic resistance to pathogens and other stressors is the purpose of the review.
Природная иммунность растений обеспечивается конститу­тивным и индуцироваными механизмами. Конститутивный механизм обусловлен строением клеточной стенки растения, а разновидности индуцированных — возникают после взаимо­действия растения с патогенными некротрофными микроор­ ганизмами, непатогенными бактериями, а также после кон­такта с некоторыми натуральными или синтетическими веществами. Целью представленного обзора является освеще­ние обоих типов механизмов формирования устойчивости растений к патогенам и прочим стрессорам
Автор огляду висловлює щиру подяку проф. P. І. Гвоздяку за критичні зауваження.
uk
Інститут молекулярної біології і генетики НАН України
Біополімери і клітина
Огляди
Механізми природної імунності рослини
Механизмы природной иммунности растений
Mechanisms of plant innate immunity
Article
published earlier
spellingShingle Механізми природної імунності рослини
Козировська, Н.О.
Огляди
title Механізми природної імунності рослини
title_alt Механизмы природной иммунности растений
Mechanisms of plant innate immunity
title_full Механізми природної імунності рослини
title_fullStr Механізми природної імунності рослини
title_full_unstemmed Механізми природної імунності рослини
title_short Механізми природної імунності рослини
title_sort механізми природної імунності рослини
topic Огляди
topic_facet Огляди
url https://nasplib.isofts.kiev.ua/handle/123456789/156471
work_keys_str_mv AT kozirovsʹkano mehanízmiprirodnoíímunnostíroslini
AT kozirovsʹkano mehanizmyprirodnoiimmunnostirastenii
AT kozirovsʹkano mechanismsofplantinnateimmunity