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...
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| Published in: | Біополімери і клітина |
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| Date: | 2006 |
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| Language: | Ukrainian |
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Інститут молекулярної біології і генетики НАН України
2006
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| Cite this: | Механізми природної імунності рослини / Н.О. Козировська // Біополімери і клітина. — 2006. — Т. 22, № 2. — С. 91-101. — Бібліогр.: 112 назв. — укр., англ. |
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Digital Library of Periodicals of National Academy of Sciences of Ukraine| _version_ | 1859915944899379200 |
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| 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. |
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| 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
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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.
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|
| 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 |