Iaoeiiiaeueia aeaaeaiiy iaoe Oe?a?ie
Iinoeooo aenia?eiaioaeueii? iaoieiai?, iieieiai? i ?aaeiiaiieiai?
ii. ?.?.Eaaaoeueeiai
I?i?noa?noai ioi?iie caei?ia’y Oe?a?ie
Oe?a?inueeee iaoeiai-aeine?aeiee ?inoeooo iieieia?? ? ?aae?ieia??
Ia i?aaao ?oeiieno
Oeeiia THeiy Aieiaeeie?iaia
OAeE 632.118.3:632.35
?AAeIIAIIEIAI*II AEANOEAINO? ?AAEOe?? IAAe*OOEEAINOI
O ?INEEI
03.00.01. – ?aaeiiaiieiaiy
Aaoi?aoa?ao
aeena?oaoei? ia caeiaoooy iaoeiaiai nooiaiy
eaiaeeaeaoa aiieiai/ieo iaoe
Ee?a – 1999
Aeena?oaoei?th ? ?oeiien.
?iaioa aeeiiaia ia eaoaae?i ?aaeiiaiieiai? Ee?anueeiai oiiaa?neoaoo ii.
Oa?ana Oaa/aiea oa o a?aeaeiei aiioiceee oa ?aaeiiaiieiai? Iinoeoooo
eeioeiii? aiieiai? oa aaiaoe/ii? iiaeaia?i? IAI Oe?a?ie.
Iaoeiaee ea?iaiee – aeaaeai?e IAI Oe?a?ie, aeieoi? aiieia?/ieo iaoe,
i?ioani?
A?iaec?inueeee Aeieo?i Ieoaeeiae/,
caa?aeoaa/ a?aeae?eo a?io?ceee oa
?aae?ia?ieia?? Iinoeoooo
eeioeiii? aiieiai? oa aaiaoe/ii?
?iaeaia??? IAI Oe?a?ie
Ioioeieii iiiiaioe: aeieoi? aiieia?/ieo iaoe, i?ioani?, /eai-ei?. OAAI
Aoaeeia ?ai? Ieeieaeiae/,
Iaoe?iiaeueiee aa?a?iee oi?aa?neoao
OAAI,
caa?aeoaa/ eaoaae?e ?aae?ia?ieia??;
aeieoi? aiieia?/ieo iaoe, i?ioani?
Aieoe?oeueeee Aieiaeeie? Ieoaeeiae/,
Ee?anueeee oi?aa?neoao ?i. Oa?ana
Oaa/aiea
I?i?noa?noaa ina?oe Oe?a?ie,
i?ioani? eaoaae?e a?io?i??
I?iaiaeia onoaiiaa – ?inoeooo i?e?ia?ieia?? oa a??onieia??
?i. Ae.E.Caaieioiiai IAI Oe?a?ie
Caoeno aiaeaoaeaoueny “__” __________ 1999 ?ieo i 13 aiae 30 oa ia
caniaeaiii niaoeiaeiciaaii? a/aii? ?aaee Ae 26.155.01 a Iinoeoooi
aenia?eiaioaeueii? iaoieiai?, iieieiai? i ?aaeiiaiieiai? ii. ?.?.
Eaaaoeueeiai IAI Oe?a?ie (252022, Ee?a-22, aoe. Aaneeueeianueea, 45).
C aeena?oaoei?th iiaeia iciaeiieoenue o aiaeiioaoei IAII? IAI Oe?a?ie.
Aaoi?aoa?ao ?icineaiee “__” ________ 1999 ?.
A/aiee nae?aoa?
niaoeiaeiciaaii? a/aii? ?aaee
aeieoi? iaaee/ieo iaoe
?.A.Aa?aiaiei
CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE
Aeooaeueiinoue oaie. Aeine?aeaeaiiy ?aae?ia?ieia?/ieo aoaeo?a
?ii?coth/iai aei?ii?ithaaiiy a oe?ieiio ae?aiacii? aeic, aecia/aiiy
oeiiaeo ci?i oa iniaeeainoae ?aae?iaaiieo no?oeoo?ieo, a oaeiae
ooieoe?iiaeueieo ii?ooaiue a a?ieia?/ieo nenoaiao ? aeooaeueiith
i?iaeaiith no/anii? ?aae?ia?ieia??. I?eeeaaeii neeaaeii? a?ieia?/ii?
nenoaie aeeiai?/iiai oa?aeoa?o iiaeia aaaaeaoe nenoaio, yea caaacia/o?
cae?eniaiiy ?aaeoe?? iaae/ooeeaino? o ?ineei. ?aaeoeiy iaae/ooeeainoi
?ineei (?I*) iieyaa? a caaeaaei eeioei, iiioeieiaai?e ?ici?ciaaaiiyi
ieie iaoiaaio. Oea i?eaiaeeoue aei iae?ioecaoei? oeaiei o iinoei
eieaeicaoei? iaoiaaia oa iaiaaeaiiy eiai ciie eieiiicaoei? [Atkinson,
1993; Klement et al., 1990]. Oi/a ?I*, yea i?iyaey?oueny o i?enei?ai?e
caaeaae? ?io?eiaaieo ee?oei, ? iaeiei c iaeiioe?ai?oeo oei?a caoenoo
?ineei a?ae o?oiiaoiaaiieo i?e?ii?aai?ci?a, ? ?? aeineiaeaeaiith
i?enay/aia cia/ia eieueeinoue ?iaio, aea aei oeeo i?? ua ia noai?aii
ocaaaeueiaii? ea?oeie ?icaeoeo oe??? ?aaeoei?. Aeineiaeaeai? oieueee
ie?aii aoaie i?ioeano ?icaeoeo ?I*, yei caeaaieueoiai oi?eathoueny ??
iieaeoey?iiai ?iaiy [Kombrink , Somssich , 1995; Levine et al., 1994].
A?aoiaoth/e aacia?a/io cia/eiinoue ?acoeueoaoia, yei iiaea aeaoe
aeineiaeaeaiiy iniaeeainoae oi?ioaaiiy ?aaeiiaiieiai/ii? ?aaeoei?
neeaaeii? nenoaie, yeith ? ?I*, neiae cacia/eoe, ui aeine?aeaeaiiy
aeiciaeo caeaaeiinoae ?aae?ia?ieia?/ieo aoaeo?a neeaaeieo nenoai
aeathoue ciiao ?ice?eaaoe ?o no?oeoo?iiooieoe?iiaeueio i?aai?caoe?th.
*anoeiaei ae?ioaiiyi oei?? i?iaeaie ? aeyaeaiiy e?eoe/ieo aoai?a oa
iia’ycaieo c ieie no?oeoo?, ?aaeiaoeieia o?aaeaiiy yeeo ?iaeoue iniiaiee
aianie a ?aaeiiaiieiai/io ?aaeoeith nenoaie. Ana oea aeaei iiaenoaae
canoinoaaoe iiiicoth/a aei?iiiithaaiiy aeey aeineiaeaeaiiy iniiaieo
aoaiia ?icaeoeo ?aaeoei? iaae/ooeeainoi ca ?o ?aae?i/ooeea?noth.
Iaoa i caaea/i aeineiaeaeaiiy. Iaoith ?iaioe aoei aeine?aeeoe i?ioean
?icaeoeo ?I*, c’ynoaaoe e?eoe/ii ii aiaeiioaiith aei aieeao ii?iiiiaiiy
aoaie ?icaeoeo ?aaeoei? ca e?eoa???i ?aae?i/ooeeaino?. A caaea/i
aeineiaeaeaiiy aoiaeeei:
1. Aea/eoe aieea ?ii?coth/iai aei?ii?ithaaiiy ia ?ici? aoaie ?icaeoeo
?I*, iaea?aeaoe ?? aeiciai oa /aniai oa?aeoa?enoeee.
2. Anoaiiaeoe aieea ii?ii?iaiiy o?oiiaoiaaiieo aaeoa?ie ia i?ioean
?icaeoeo ?I*.
3. Ioeiieoe aoaeo eiia?iiaaii? aei? ii?iiiiaiiy ?ineei oa o?oiiaoiaaiieo
aaeoa?ie o oi?ioaaiii ?aaeiiaiieiai/ii? ?aaeoei? a nenoaii, ui
caaacia/o? ?icaeoie ?I*.
Iaoeiaa iiaecia iaea?aeaieo ?acoeueoaoia. Aia?oa i?iaaaeaia nenoaiaoe/ia
aeine?aeaeaiiy aieeao iiiicoth/iai aei?iiiithaaiiy ia i?ioean ?icaeoeo
?I*. Anoaiiaeaii /aniaee i aeicicaeaaeiee oa?aeoa? aoaeoia, ye?
ni?e/eiai? ii?ii?iaiiyi ?ineei ia ?icieo aoaiao ?icaeoeo ?aaeoei?
iaae/ooeeainoi. Iieacaii, ui ia ?icaeoie ?aaeoei? iaae/ooeeainoi aieeaa?
ii?iiiiaiiy ye ?ineei, oae i aaeoa?ie, aecia/aii aeiciai oa?aeoa?enoeee
oeueiai aieeao.
Ca aeiiiiiaith ?aa?an?eiiai aiae?co (iaoiae Aiena-O?eniia) iaea?aeaii
noaoenoe/io iiaeaeue ?aaeiaoeieiiai aieeao ia ?I*, c aeei?enoaiiyi yei?
aoei aecia/aii iacaeaaeio eiiiiiaioo ?aaeiaoeieiiai o?aaeaiiy ?ineei i
aaeoa?ie oa ioeiiaii aianie aca?iiaei? aoaeoia ii?iiiiaiiy ?ineei i
aaeoa?ie a i?ioean ?icaeoeo ?I*. Iieacaii, ui ia aoai? ?iaeoeoe??
?aaeoe?? o?oiiaoiaaii? aaeoa???, ye? ?iaeoeoaaee ?? ?icaeoie, oa
?ii?coth/a aei?ii?ithaaiiy ae?yee ye aioaaii?noe, aca?iii iineaaethth/e
aieea iaeia iaeiiai. Oea naiae/eoue i?i oa, ui iiaeoeoeiy caaeaaei
?ineeiieo eeioei i?e ?I* oa ii?iiiiaiii iiaea aiaeaoaaoeny ca iiae?aieie
iaoaiiciaie.
I?aeoe/ia cia/aiiy ?iaioe iieyaa? a oiio, ui anoaiiaeaiiy e?eoe/ieo
aoaiia o i?ioeani ?I* ii aiaeiioaiith aei aieeao ii?iiiiaiiy ? aaaeeeaei
oeyoii aea/aiiy iaoaiiciia oei?? ?aaeoei?. Aeine?aeaeaiiy iaoaiici?a
oiceei-oiii/ieo, aiioiii/ieo oa oiciieiai/ieo i?ioeania, ui
?icaeaathoueny i?e eiioaeoi oioiiaoiaaia oa ?ineeie a oiaei oei??
?aaeoei?, aeicaiey? aeoeaii ao?o/aoeny o oi?ioaaiiy noieeinoi ?ineei oa
noai?thaaoe iia?, aeueoa?iaoeai? caniae caoenoo ?ineei. Oea iaaoaa?
iniaeeai aaaeeeaiai cia/aiiy aeey oa?eoi??e ?c cia/ieie ??aiyie
?aae?iioee?aeiiai caa?oaeiaiiy. E?ii oiai, ?icoiiiiy nooiinoi iaoaiiciia
aca?iiaei? oioiiaoiaaia oa aeui? ?ineeie ni?ey? ciaoiaeaeaiith iaeaieueo
i?eaeaoieo e?eoa???a, ui oa?aeoa?ecothoue noieeinoue ?ineei, yei iiaeia
aeei?enoaoe a naeaeoeieiie i?aeoeoei.
Iniaenoee aianie caeiaoaa/a iieyaa? o aeeiiaii? anueiai ianyao
aenia?eiaioaeueii? /anoeie aeena?oaoe??, noaoenoe/i?e ia?iaoe?
?acoeueoao?a, i?aeai?? oa aiae?c? aeaieo iaoeiai? e?oa?aoo?e, a oaeiae –
ca o/anoth iaoeiaiai ea??aieea oa ni?aaaoi??a ioae?eaoe?e – aiae?c? oa
?ioa?i?aoaoe?? iaea?aeaieo ?acoeueoao?a. I?e iaeaaeoe?
aenia?eiaioaeueii? onoaiiaee aeey aei??thaaiiy aeaeo?iiii?o c?ace?a
aaoi?o iaaeaaaee aeiiiiiao ni?a?ia?oieee a?aeae?eo ?.?. ?eaaeea ?
TH.I.Iaaeaiei.
Ai?iaaoeiy ?acoeueoaoia aeena?oaoei?. Iniiaii iieiaeaiiy aeena?oaoeieii?
?iaioe aeiiiaiaeaeenue ia II C’?caei ?aaeiiaiieiaia Oe?a?ie
(Aeiii?iiao?ianuee,1995); IV Iiaeia?iaeiie eiioa?aioei? “AEeooy oa
nia?oue eeioeie” (Aaeiiao?a, Aiaeiy, 1996); VI Eiioa?aioei? iieiaeeo
a/aieo, i?enay/aiie 50-e ?i/ieoei c aeiy canioaaiiy Iinoeoooo oiciieiai?
?ineei i aaiaoeee IAI Oe?a?ie (Ee?a, 1996); III C’?caei ii ?aaeiaoeieiei
aeineiaeaeaiiyi (Iineaa, 1997).
Ioaeieaoei?. Iniiaiee ciino aeena?oaoeieii? ?iaioe aeeeaaeaii o 9
ioaeieaoeiyo o oi?ii iaoeiaeo noaoae oa oac aeiiiaiaeae.
No?oeoo?a oa ia’?i ?iaioe. Aeena?oaoeiy neeaaea?oueny c anooio, iaeyaeo
eioa?aoo?e (3 ?icaeiee), iieno iaoa?iaeia oa iaoiaeia aeineiaeaeaiiy,
aeeeaaeaiiy ?acoeueoaoia oa ?o iaaiai?aiiy (4 ?icaeiee), aeniiaeia,
caeeth/aiiy, ia?aeieo eioa?aoo?e (225 aeaea?ae, a oiio /enei – 145
ca?oaiaeieo i?aoeue). ?iaioa i?aaenoaaeaia ia 184 noi?iieao iaoeiiieno,
iinoeoue 21 ?enoiie oa 14 oaaeeoeue.
IA’?EOE OA IAOIAeE AeINEIAeAEAIUe
A yeino? ia’?eo?a i?e aeine?aeaeaii? i?ioean?a, iia’ycaieo c iaoiaaiacii
oa no?ee?noth o ?ineei ?, cie?aia, ?icaeoeii ?I*, oe?iei
aeei?enoiaothoue ?ineeie othothio oa o?oiiaoiaaii? aaeoa??? Pseudomonas
syringae [Klement, 1963; Klement et al., 1990]. A aeine?aeao
aeei?enoiaoaaee ?ineeie othothio Nicotiana sanderae oa ianiaoeeo?/iee
ooai oioiiaoiaaiieo aaeoa?ie Pseudomonas syringae pv. phaseolicola 291
(c eieaeoei? aiaeaeieo oioiiaoiaaiieo aaeoa?ie Iinoeoooo iie?iaiieiai?
oa ai?onieiai? ii. Ae.E.Caaieioiiai IAI Oe?a?ie). Ca etha’ycia iaaeaiiy
eoeueoo? oa oeiiii eiinoeueoaoei? aeneiaeth?ii aeeaieo iiaeyeo
caaiaeoaa/o aiaeaeieii i?ioani?o ?.I.Aaicaeyeo oa n.i.n., eaiae. aiie.
iaoe E.I.sseiae?aie.
?ineeie othothio ae?iuoaaee a oiiaao oaieeoei aei oace 5-7 ?icaeiooeo
eenoeia. A aenia?eiaioao aeei?enoiaoaaee ?icaeioo? eenoee 2-3 y?on?a.
Aaeoa?i? ae?iuoaaee ia oaa?aeiio iiaeeaiiio na?aaeiaeui – ea?oiieyiiio
aaa?i [Aaeueotheiaa e ae?., 1968]. Iiney 48 aiae eoeueoeaoaaiiy
aaeoa?iaeueii eeioeie cieaaee noa?eeueiith aiaeith c iiaa?oii aaa?o i
nociaicoaaee. Noniaici? aeiaiaeeee aei eiioeaio?aoei? 108 eeioei/ie.
Aeey iiaeoeoaaiiy ?icaeoeo ?I* aeei?enoiaoaaee iaoiae
ii’?eoeie-iioieueo?aoeie ca?aeii [Klement, 1963; Element et al., 1990] a
iaai?e iiaeeo?eaoe??. Eenoee othothio iioieueo?oaaee noniaici?th aeeaeo
aaeoa?iaeueieo eeioei oa noa?eeueiith aiaeii?iaiaeiith aiaeith
(eiio?ieue), aaiaey/e ?o ca aeiiiiiaith oi?eoea a iiaeeeioeiiee i?inoi?
eenoeia. I?e oeueiio aaeoa?i? aaiaeeee a iaeio iieiaeieo eenoea, a aiaeo
– a iioo.
Iaoiae, yeee aeei?enoiaoaaee aeey e?euee?nii? oa?aeoa?enoeee ?icaeoeo
?I*, aaco?oueny ia caeaaeiino? e?eueeino? aeaeo?ie?o?a, ui aeoiaeyoue c
?ineeiieo ee?oei aiane?aeie ci?ie i?iieeiino? ee?oeiieo iaia?ai, a?ae
nooiaith ?icaeoeo ?aaeoe?? iaae/ooeeaino? ? iieyaa? o aecia/aii?
aeaeo?ii?ia?aeiino? aethao?a, iaea?aeaieo c eenoe?a i?ioyaii iaaiiai
/ano i?ney ?o ?io?eueo?aoe?? aaeoa??aeueiith noniaic??th [Cook, 1973;
Stall et al., 1982; Klement et al., 1990]. Aeey niinoa?aaeaiiy ca
aeeiai?eith i?ioeano ?icaeoeo ?I* /a?ac 1-24-aiaeeii? iioa?aaee ?iaeee
aeni/ee c iiieoeueiaaieo eenoeia (o aeaeyaei aeeneia aeiaiao?ii 1 ni),
aeo?eioaaee ?o i?ioyaii 17 aiae o aeenoeeueiaaiie aiaei i?e e?iiaoi?e
oaiia?aoo?? (15 aeeneia ia 30 ie aiaee), iiney /iai aeii?thaaee
aeaeo?e/iee iii? iaea?aeaieo ?ic/eiia aeaeo?ieioia (aethaoia). Ii
cia/aiiyi aeaeo?e/iiai iii?o oeeo aethaoia ae?aoiaoaaee ?o ieoiio
aeaeo?ii?iaiaeiinoue (Ni.i).
Aeey aea/aiiy aieeao iiiicoth/iai aei?iiiithaaiiy ia ?icaeoie ?I*
?ineeie othothio oa noniaici? aaeoa?iaeueieo eeioei iiaeaeaaaee aei?
aino?iai aaiia-aei?iiiithaaiiy ia onoaiiaoei “Enneaaeiaaoaeue” a
aeiaiaciii aeic aiae 1 aei 600 A? (60Co, iiooaeiinoue aeice 6,3 –
7,5.10-2 A?/n). A ia?o?e na??? aeine?ae?a ?ineeie othothio ii?ii?ithaaee
a aeicao 150, 300 oa 600 A? ca 0,5 aiae aei ca?aaeaiiy ?o aaeoa??yie. A
ae?oaie na?i? aeineiaeia ?ineeie othothio nia?oo iiieoethaaee
noniaici?th aaeoa?iaeueieo eeioei i iioii, /a?ac iaaii /aniai i?iiiaeee,
ii?iiiithaaee a aeicao aiae 1 aei 600 A?.
Aeey ioeiiee aieeao aaiia-aei?iiiithaaiiy ia aeaeeaaiiy aaeoa?ie P.
syringae pv. phaseolicola 291 noniaicith 24-aiaeeiii? eoeueoo?e aaeoa?ie
(106 eeioei/ie) ii?iiiithaaee a aeicao aiae 1 aei 100 A?. Ii?iiiiaii
noniaici? aeiaiaeeee aei eiioeaio?aoei? 5.103 eeioei/ie i aeniaaee ia
oaa?aea iiaeeaia na?aaeiaeua. Eoeueoeaoaaee 48 aiae i?e oaiia?aoo?i 28
0N, iiney /iai iiae?aoiaoaaee eieueeinoue iae?ieieiiie, ui ooai?eeeny.
Aeaeeaaiiy aaeoa?ie ioeiithaaee ii aiaeiioaiith aei iaii?iiiiaiiai
eiio?ieth.
Noaoenoe/io ia?iaeo ?acoeueoao?a i?iaiaeeee caaaeueiii?eeiyoeie
iaoiaeaie aa??aoe?eii? noaoenoeee [Ieioeineee, 1980].
Aeey aeineiaeaeaiiy aoaeoia eiia?iiaaii? ae?? ii?iiiiaiiy ?ineei oa
aaeoa?ie o oi?ioaaiii ?I* aoa canoiniaaiee iaeei c iaoiae?a ?aa?an?eiiai
aiae?co – iaoiae Aiena-O?eniia [Aaeea? e ae?., 1976; Aiiaea?ue,
Noaothoa, 1976]. Aeineiaee i?iaiaeeee ii noaiaea?oi?e noai? iiaiiai
oaeoi?iiai aenia?eiaioo oeio 2n (n – /enei oaeoi?ia aieeao ia nenoaio)
aeey aeaio oaeoi?ia, eiaeai c yeeo i?eeiaa ii aeaa cia/aiiy. Ieai
aeineiaeia ia?aaeaa/aa iaea?aeaiiy aenia?eiaioaeueieo cia/aiue
aeineiaeaeoaaieo oa?aeoa?enoee i?e anio iiaeeeaeo eiiaiiaoeiyo ?iaiia
aeaieo oaeoi?ia.
I?iaaaeaiiy aeineiaeia ii oaeie noai? aeicaiey? ioeiieoe ia?aiao?e
?aa?anieiiai ?iaiyiiy oeio
y = b0 + b1x1 + b2x2 + b1b2x1x2..
Eiaoioei?ioe b1 i b2 aeacothoue ia iacaeaaeiee aianie eiaeiiai c
oaeoi?ia o cia/aiiy aeineiaeaeoaaii? ooieoei? aiaeaoeo, a eiaoioei?io
b1b2 – ia nooiiiue aca?iiaei? oeeo oaeoi?ia.
?ACOEUeOAOE OA ?O IAAIAI?AIIss
1. Aieea iiia?aaeiueiai ii?iiiiaiiy ?ineei ia ?icaeoie ?aaeoei?
iaae/ooeeainoi
Aiaeicoth/e aeai?, ui aiaeia?aaeathoue ciiie a aeaeo?ii?iaiaeiinoi
aethaoia, iaea?aeaieo c eenoeia ?ineei, ii?iiiiaieo ca 0,5 aiae aei
ca?aaeaiiy aaeoa?iyie, aiaecia/aee aeaa iaeneioie c?inoaiiy
aeaeo?ii?iaiaeiinoi, yei iaee iiaeiaiee oa?aeoa? aeey anio canoiniaaieo
aeic ii?iiiiaiiy (150, 300 ? 600 A?). ?en. 1 aiaeia?aaea? oeiiao
aeeiaiieo ciiie aeaeo?ii?iaiaeiinoi aethaoia (aeica ii?iiiiaiiy 300 A?).
?en. 1 Aeeiai?ea aeaeo?ii?ia?aeiino? aethao?a i?e ii?ii?iaii? ?ineei (
aeica 300 A?)
aei ?o ?iieoeyoe?? aaeoa??yie oa aiaeith
Ia?oee c iaeneioiia niinoa?iaaee /a?ac iaeio aiaeeio iiney iiieoeyoei?
aaeoa?iyie ii?ii?iaieo ?ineei. Aii iaa ianiaoeeoi/iee oa?aeoa?,
ineieueee aiaecia/aany ye o aeineiaeiiio (i?e ?iieoeyoe?? ?ineei
aaeoa?iyie), oae i a eiio?ieueiiio (iiieoeyoeiy ?ineei aiaeith)
aa?iaioao. O iaii?ii?iaiiio aa??aio? oeae iaeneioi a?aenooi?e.
C o?aouei? aiaeeie niinoa??aaee cieaeaiiy aeaeo?ii?iaiaeiinoi, yea
a?aeaoaaeiny, i/aaeaeii, ca ?aooiie aiaeiiaeaiiy ooieoei? iaia?ai
?ineeiieo eeioei, ? iaei aeicicaeaaeiee oa?aeoa?. Iaeaieueoa cieaeaiiy
aeaeo?ii?ia?aeiino? niinoa?iaaee i?e aeici 150 A?, oiaei ye i?e aeicao
300 i 600 A? aiii aoei cia/ii iaioei. Iiney o?aouei? (aeice 150, 600 A?)
oa i’yoi? aiaeei (aeica 300 A?) o aeiaaeeo iiieoeyoei? aaeoa?iyie
niinoa?iaaee ae?oaa c?inoaiiy aeaeo?ii?iaiaeiinoi aethaoia, yea oeueiai
?aco iaei niaoeeoi/iee oa?aeoa? i aoei iia’ycaia aaea aaciina?aaeiuei c
?icaeoeii ?aaeoei? iaae/ooeeainoi.
I?e aeicao 300 oa 600 A? aaea ia 9 aiae aeaeo?ii?iaiaeiinoue aethao?a
aoea aeecueeith aei iaeneiaeueii?, yea aecia/aeany i?e caaeaae? ee?oei
ia eiioeaaiio aoai? ?icaeoeo ?I*. O iaii?ii?iaiiio aa??aio? oaeee noai
niinoa??aaee ia 24-oo aiaeeio.
A oaae. 1 iaaaaeai? a?aeiini? aaee/eie aeaeo?ii?ia?aeiino? (ca 100 %
i?eeiaee aaee/eio aeaeo?ii?ia?aeiino? aethao?a, iaea?aeaieo c
iaii?ii?iaieo ?ineei). I?e aenieeo aeicao ii?ii?iaiiy (300, 600
A?) cia/aiiy aeaeo?ii?ia?aeiino? ia anueiio /aniaiio ?ioa?aae? cia/ii
ia?aaeuothoue 100 %-iee ??aaiue, oiaoi oaeoe/ii aanue i?ioean ?icaeoeo
?I* i?ioiaeea ia oii? i?aeaeuaii? i?iieeiino? iaia?ai ? oea ia iiaei ia
aieeiooe ia eiai aeeiai?eo.
Oaaeeoey 1. Aeeiai?ea a?aeiinieo aaee/ei aeaeo?ii?ia?aeiino?
i?e ii?ii?iaii? ?ineei aei ?iieoeyoe?? aaeoa??yie. Ca 100 %
i?eeiaee aeaeo?ii?ia?aei?noue aethao?a c eenoe?a iaii?ii?iaieo ?ineei,
?iieoeueiaaieo ?I*-?iaeoeoth/eie aaeoa??yie.
*an i?ney ?iieoeyoe?? aaeoa??e, aiae
Aeica ii?ii?iaiiy
150 A?
Aeica ii?ii?iaiiy
300 A?
Aeica ii?ii?iaiiy
600 A?
1 289(4 536(2 407(2
3 158(1 423(2 358(3
5 143(4 207(1 310(2
7 102(4 183(4 195(3
9 114(5 212(2 272(5
11 103(8 134(3 128(3
24 88(5 100(4 100(3
Oaeei /eiii, iiaeia eiinoaooaaoe i?enei?aiiy ?icaeoeo ?I* o aeiaaeeo
iiia?aaeiueiai ii?iiiiaiiy ?ineei a aeicao 300 oa 600 A?. Iineeaiith
?aaeoei? iiaee ni?eyoe ?aaeiaoeieii-iiaeoeiaaii ciiie, iia’ycaii
ianaiia?aae ci ciiiith i?iieeiinoi iaia?ai oa ia?a?iciiaeieii iiiia, yei
aiaeia?athoue ?aaoeyoi?io ?ieue, ui i?eaiaeeoue aei aeenaaeaino
aaaaoueio eeioeiieo i?ioean?a. Aiaeiii, ui ciiie a iiiiiio (ianaiia?aae,
E+ oa Na2+) aaeaini caaaeaee ia?aaeothoue i ? iaiaoiaeiith oiiaith
caaeaaei ?ineeiieo eeioei aianeiaeie ?I* [Kombrink, Somssich, 1995;
Atkinson, 1993], a ii?iiiiaiiy iiaei aeeeeeaoe oae? ciiie ?aiioa i
aoaeoeaiioa, iiae oea aiaeaoaa?oueny i?e aeoeaaoei? iiiiieo eaiaeia
aianeiaeie iiaeoeoei? ?I*. Oaeee i?aeo?ae aeyaea neia?a?ci o ae??
?ii?coth/iai aei?ii?ithaaiiy oa o?oiiaoiaaiieo aaeoa??e, i?ioa ia
aeicaieea no?oeoo?oaaoe i?ioean ?icaeoeo ?aaeoe??.
2. Aieea ?ii?coth/iai aei?ii?ithaaiiy ia ?ici? aoaie ?icaeoeo ?aaeoei?
iaae/ooeeainoi
Aeoiaey/e ci neeaaeiinoi oa aaaaoiaoaiiinoi i?ioeano ?icaeoeo ?I*,
aiaeiiiiinoae no?oeoo? oa i?ioeania, ui aeiiiiothoue ia ?icieo aoaiao
?aaeoei?, iiaeia i?eionoeoe ??ciee ??aaiue ?o e?eoe/iinoi ii
aiaeiioaiith aei aei? ?ii?coth/iai aei?iiiithaaiiy ? aeae?eeoe ie?ai?
aoaie ?aaeoe?? ca e?eoa???i ?o ?aae?/ooeeaino?. C oe??th iaoith aoei
i?iaaaeaii aeineiaeaeaiiy aieeao aaiia-aei?ii?ithaaiiy ia ?I* a ?icii
iiiaioe ?? ?icaeoeo.
sseui ?ineeie ii?iiiithaaee /a?ac 5-20 oa iiney iiieoeyoei? aaeoa?iyie,
oi i?e aeicao a?eueoa 10 A?, niinoa?iaaee i?eaii/aiiy ?icaeoeo ?aaeoei?
(?en. 2,1).
?en. 2 Caeaaei?noue a?ae aeice ii?ii?iaiiy aeaeo?ii?ia?aeiino?
aethao?a i?e ii?ii?iaii? ?ineei /a?ac 5-20 oa (1), 30-60 oa (2), 2-5
aiae (3) i?ney ?iieoeyoe?? ?ineei ?I*-?iaeoeoth/eie aaeoa??yie
Ca 100 % i?eeiaee aeaeo?ii?ia?aei?noue aethao?a c eenoe?a iaii?ii?iaieo
?ineei, ?iieoeueiaaieo ?I*-?iaeoeoth/eie aaeoa??yie
Ia 24-oo aiaeeio (/an iiaiiai ?icaeoeo ?I* a eiio?iei) i?e aeicao
ii?iiiiaiiy 50-65 A? cia/aiiy aeaeo?ii?iaiaeiinoi aethaoia noaiiaeei
aeecueei 78 % aiae eiio?ieueii? aaee/eie (iaii?iiiiaii ?ineeie,
iiieoeueiaaii aaeoa?iyie), a i?e aeicao, ui ia?aaeuoaaee 85 A? –
i?eaeecii 28 %, ui aei?iaithaaei cia/aiith aeaeo?ii?ia?aeiino? o
aeiaaeeo, eiee ?ineeie iiieoethaaee aiaeith, oiaoi i?e iiaiie
aiaenooiinoi ?icaeoeo ?I* .
I?e caieueoaiii /aniaiai iioa?aaeo iiae iiieoeyoei?th aaeoa?iyie oa
ianooiiei ii?iiiiaiiyi ?ineei aei 30-60 oa iiaiaoth/ee aieea ii?iiiiaiiy
i?iyaeyany i?e aeicao, yei ia?aaeuoaaee 65 A? (?en. 2,2), a i?e
ii?iiiiaiii /a?ac 2-5 aiae – 85 A? i aeua (?en. 2, 3).
Aiae?c iaea?aeaieo ?acoeueoaoia na?ae/eoue, ui ia?aii aeey
aeine?aeaeaiue aeice ii?iiiiaiiy ? cia/ii ieae/eie aiae aeic,
canoiniaaieo o iiia?aaeiie na?i? aeineiaeia i?e ii?iiiiaiii ?ineei aei
iiieoeyoei? aaeoa?iyie, ? ni?e/eieee cia/iee iiaiaoth/ee aieea ia
?icaeoie ?I*.
A?aeiinii aei? ?aaeiaoei? ia ?aiiio aoaiao ?I* (1-3 aiae) iiaeia
i?eionoeoe, ui iiaiaoth/ee aoaeo ii?iiiiaiiy iaoiiaeth?oueny aieeaii ia
i?ioeane ?iciiciaaaiiy oa neaiaeueii? o?ainaeoeoei? /a?ac aeith ia naii
aaeoa?iaeueii eeioeie (?o aeineoi?io aeoeaiinoue), iaoaaieici
neaiaeueieo eiiiiiaio ?ineeiieo eeioei, noai oa ooieoei? eeioeiieo
iaia?ai, aeoeaiinoue ?yaeo oa?iaioia oiui.
Ui noino?oueny aeyaeaiiai iiaiaoaaiiy ?aaeoei? i?e aei? ?aaeiaoeieiiai
oaeoi?o ia aieueo iiciio aoaiao ?? ?icaeoeo (5 aiae, cie?aia), oi iiaeia
i?eionoeoe, ui iiaiaoth/ee aoaeo ii?iiiiaiiy ? ?acoeueoaoii aieeao ia
i?ioeane, yei eaeooue a naieo ?ineeiieo eeioeiao ia iiciioeo noaaeiyo
?aaeoei?, i aaea ia iia’ycaii aaciina?aaeiuei c aca?iiaei?th ?ineeiieo
eeioei c iaoiaaiii, ye ne?aeo? c anoaiiaeaieo oaeoia ?nioaaiiy
ia?iiaeecaoei? a ?icaeoeo ?I* [Klement, 1971; Klement et al., 1990;
Atkinson, 1993]. Iniaeeainoth aieeao ?aaeiaoeieiiai oaeoi?o ia oeueiio
aoaii ? oa, ui a?i caeaoai caaeieoaaoe ?aaeicaoeith ?yaeo iaoiiaeaieo
iiia?aaeiii oiaeii iiaeie i?ioeania, cie?aia, c?inoaiiy i?iieeiinoi
iaia?ai ?ineeiieo eeioei.
3. Aianie ii?iiiiaiiy iaoiaaio o ?icaeoie ?aaeoei? iaae/ooeeaino?
?en. 3 Aeaeo?ii?ia?aei?noue aethao?a i?e ii?ii?iaii? aaeoa??e
Ca 100 % i?eeiaee aeaeo?ii?ia?aei?noue aethao?a c eenoe?a ?ineei,
?iieoeueiaaieo iaii?ii?iaieie aaeoa??yie
Aiae?c ?acoeueoao?a, i?aaenoaaeaieo ia ?en. 3 na?ae/eoue, ui i?e
?iieoeyoe?? ?ineei aaeoa??yie, ii?iiiiaieie o aeici 1 A?, ia 24-oo
aiaeeio niinoa?aaeaiiy aaee/eia aeaeo?ii?iaiaeiinoi aethaoia c ?ineeiieo
oeaiei, ca yeith, ye i a iiia?aaeiio aeineiaeao, oa?aeoa?ecoaaee
nooiiiue ?icaeoeo ?aaeoei? , aei?iaithaaea eiio?ieueiie, a i?e
caieueoaiii aeice ii?ii?iaiiy aaeoa??e aei 10 A? noaiiaeea ia?an?/ii 39
% eiio?ieueii? aaee/eie. I?e oeueiio niinoa?iaaee oei?ic iiieoeueiaaieo
cii, ui ? iaoeiiaei aeey ?I*, a iae?ioe/ii neiioiie i?iyaeyeeny iiciioa
oa neaaoa, iiae o eiio?iei, yeee iioieoaaee iaii?iiiiaieie aaeoa?iyie.
sseui canoiniaoaaee aeui aeice ii?iiiiaiiy aaeoa?ie (50, 100, 150 A?),
aeaeo?ii?iaiaeiinoue ia 24-oo aiaeeio noaiiaeea ia?an?/ii 32 % aiae
eiio?ieth, ui aiaeiiaiaeaei iiaiie aiaenooiinoi aoaeue-yeeo neiioiiia
?icaeoeo ?aaeoei?.
Aoei i?iaaaeaia ii?iaiyiiy aaee/eie aeaeeaaiiy aaeoa?ie i?e ii?iiiiaiii
oa ?o caeaoiinoi caioneaoe i?ioean ?icaeoeo ?aaeoei? iaae/ooeeainoi.
Ia ?en. 4 iaaaaeaia caeaaeiinoue “aeica-aoaeo”, aea a yeinoi aoaeoo
aenooiaei aeaeeaaiiy aaeoa?ie. Aiae?c caeaaeiino? nooiaith aeaeeaaiiy
aaeoa??e a?ae aeice ii?ii?iaiiy aeaco? ia oa, ui ye i aieueoinoue
i?aaenoaaieeia ?iaeo Pseudomonas [Caoa?ia, E?eaeineee, 1972], aaeoa?i?
P. syringae pv. phaseolicola 291 aeyaeeeny aeineoue ?aaeii/ooeeaeie.
I?e ni?anoaaeaii? nooiaith aeaeeaaiiy aaeoa??e oa aeaeo?ii?ia?aeiino?
aethao?a, ca yeith oa?aeoa?ecoaaee nooi?iue ?icaeoeo ?I*, iiaeia aa/eoe,
ui i?e aeici ii?iiiiaiiy 1 A?, eiee aeaeeaaiiy aaeoa?ie noaiiaeei a?ey
76 % ,
?en. 4 Aeaeeaaiiy ee?oei aaeoa??e P. syringae pv. phaseolicola i?e
?o aaiia-ii?ii?iaii?
aeaeo?ii?iaiaeiinoue, ui oa?aeoa?ecoaaea ?icaeoie ?aaeoei?, aoea
aeecueeith aei eiio?ieueieo cia/aiue. Aeica ii?iiiiaiiy aaeoa?ie 10 A?
(aeaeeaaiiy aei 39 %) iaoiiaethaaea ?icaeoie ?aaeoei? c iaaiith
cao?eieith (aaee/eia aeaeo?ii?iaiaeiinoi ia 24-oo aiaeeio noaiiaeea
aeecueei 46 % eiio?ieueii?). I?e aeici 50 A? (aeaeeaaiiy aei 15 %),
aaeoa?i? ao?a/aee caeaoiinoue iiaeoeoaaoe ?I*.
Ii?iaithth/e nooiiiue ?icaeoeo ?I* i?e ii?iiiiaiii ?ineei,
?iieoeueiaaieo aaeoa?iyie (?en. 2), ? i?e ii?ii?iaii? naieo aaeoa??e
aa/eii, ui i?ioean ?icaeoeo ?aaeoei? oa?aeoa?ecoaaany aieueoith
noieeinoth aei aieeao ii?iiiiaiiy, iiae o aeiaaeeo, eiee noniaicith
aaeoa?ie ii?iiiithaaee ie?aii aei iiieoeyoei? a iiaeeeioeiiee i?inoi?
?ineeiieo eenoeia (?en. 3). Iaaioue i?e iiiiiaeueiiio /aniaiio iioa?aaei
iiae iiieoeyoei?th aaeoa?ie i ii?iiiiaiiyi (5-20 oa) nooiiiue ?icaeoeo
?aaeoei? ia aiae?iciyany aiae eiio?ieth (100 %) i?e aeici 10 A?, i?e
aeici 50 A? iaaeeaeaany aei 78 % i i?e aeicao, yei ia?aaeuoaaee 85 A?,
?aaeoeiy ia ?icaeaaeanue (?en. 2 ). I?e iiieoeyoei? ae ii?iiiiaieo
aaeoa?ie ?aaeoeiy ia ?icaeaaeany, yeui aiie aoee ii?iiiiaii a aeici 50
A? (?en. 3). Iiaeeeai, ooo ia? cia/aiiy noeioeyoeiy ca ?aooiie eiioaeoo
?ineeiieo i aaeoa?iaeueieo eeioei, eiee no?oeoo?e inoaiiio ua ia caciaee
aieeao ii?iiiiaiiy, ineieueee aaeoa?i? iiaeooue noai?thaaoe neaiae aeey
caioneo ?I* c ia?oeo oaeeei ?o eiioaeoo c ?ineeiieie eeioeiaie [Huynk et
al., 1989; Alfano, Collmer , 1996], a oaeiae oa, ui aaeoa?iaeueii
eeioeie, iiaeeeai, caciaee aeayei? ?aaeiicaoenii? aei? c aieo ?a/iaei
?ineeiieo eeioei.
Aeoiaey/e c iaaaaeaieo oaeoia, iiaeia i?eionoeoe, ui i?e aei?
?ii?coth/i? ?aaeiaoei? iiaiaoaaiiy ?aaeoei? ?ineei iiaei aiaeaoaaoeny ye
ca ?aooiie ?aaeiieico /anoeie iieaeoe, ui aeeiiothoue ooieoeith
aeineoi?ia (?aaeiinoieea eiiiiiaioa a aeiciaie caeaaeiinoi ia ?en. 3),
oae i ca ?aooiie ii?ooaiiy ooieoeie aaeoa?iaeueieo oa?iaioia oa
aeni?ani? aaiia aai?oeaioiinoi, yei i?eeiathoue o/anoue o i?ioeanao
aca?iiaei? aaeoa?ie c eeioeiaie ?ineei-ainiiaea?ia (?aaeii/ooeeaa
eiiiiiaioa). I/aaeaeii, oei iiaeoeiaaii ?aaeiaoei?th ii?ooaiiy i
aiaeia?aaaee ae?ioaeueio ?ieue o aeieoaaiii ?icaeoeo ?I* i?e aieeai
ii?iiiiaiiy ia ?aiiio aoaiao ?? ?icaeoeo.
Oaeei /eiii, iaeaieueoith ?aaeii/ooeeainoth oa?aeoa?ecoaaany aoai
aca?iiaei? iaoiaaia c eeioeiaie ?ineeie, eiee aiaeaoaa?oueny iiaeoeoeiy
iaae/ooeeai? aiaeiiaiaei ?ineeie-ainiiaea?y no?oeoo?aie eeioei iaoiaaia.
4. Aoaeoe eiia?iiaaii? aei? oaeoi?ia ii?iiiiaiiy ?ineei i aaeoa?ie i?e
?icaeoeo ?I*
Iaiao?aeiith oiiaith ?icaeoeo ?I* ? aca?iiaeiy no?oeoo? ?ineeiieo oa
aaeoa?iaeueieo eeioei. Oe? no?oeoo?e iiaeooue caciaaaoe nooo?aeo ciii
iiae aieeaii ii?iiiiaiiy ?ineei aai aaeoa?ie. Ca aeiiiiiaith iaoiaeo
Aiena-O?eniia aoei aecia/aii iacaeaaeio eiiiiiaioo ?aaeiaoeieiiai
o?aaeaiiy ?ineei i aaeoa?ie oa ioeiiaii aianie aca?iiaei? aoaeoia
ii?iiiiaiiy ?ineei i aaeoa?ie a i?ioean ?icaeoeo ?I*.
Ieaioaaiiy aeineiaeia i?iaiaeeeinue o aiaeiiaiaeiinoi c noaiith iiaiiai
oaeoi?iiai aenia?eiaioo aeey aeaio oaeoi?ia, eiaeai c yeeo i?eeiaa ii
aeaa cia/aiiy. Iao?eoey ieaioaaiiy aenia?eiaioo oa ?acoeueoaoe aeii?ia
iaaaaeaii a oaae. 2.
Oaaeeoey 2. Iao?eoey ieaioaaiiy aenia?eiaioo ii aea/aiith aoaeoia
ii?iiiiaiiy ?ineei i aaeoa?ie i?e ?icaeoeo ?I*:
x1 – oaeoi? ii?iiiiaiiy ?ineei, x2- oaeoi? ii?iiiiaiiy aaeoa?ie, x1x2 –
aca?iiaeiy oaeoi?ia ii?iiiiaiiy ?ineei i aaeoa?ie, y – ooieoeiy aiaeaoeo
(aaee/eia aeaeo?ii?iaiaeiinoi aethaoia i?e ii?iiiiaiii o aeacaieo
aeicao)
Iiia?
aeineiaeo x1 x2 x1x2 y
1 A? y
10 A? y
50 A? y
100 A? y
150 A? y
300 A?
1
2
3
4 –
+
–
+ –
–
+
+ +
–
–
+ 100
100
100
100 100
100
45
100 100
100
40
100 100
100
40
31 100
100
40
30 100
100
40
30
Ia ?o iniiai aoee ?ic?aoiaai? eiao?oe??ioe ?iaiyiue ?aa?an??, ui
oa?aeoa?ecothoue aianee eiaeiiai c oaeoi?ia o i?ioean ?icaeoeo ?aaeoe??
a caeaaeiinoi aiae aeice ii?iiiiaiiy oa aianie aca?iiaei? oaeoi?ia (?en.
5).
Aiae?c caeaaeiino? aaee/eie aianeo oaeoi?a ii?ii?iaiiy ?ineei
(eiao?oe??io b1) a?ae aeice ii?ii?iaiiy iieacaa, ui ciae aieeao oeueiai
oaeoi?a ciiith?oueny ic c?inoaiiyi aeice ii?iiiiaiiy. Ioaea, aeey
oaeoi?a ii?ii?iaiiy ?ineei aeyaeeany iaiinoieiinoue eiai aianeo a
i?ioean ?icaeoeo ?I*: iaioi aeice ii?iiiiaiiy (aei 50 A?) ni?eyee
?icaeoeo ?aaeoei?, a aaeeei (aiae 100 A?) – iiaiaoaaiith. Noeioethaaiith
?icaeoeo ?I* o aeiaaeeo iaioeo aeic iiaee ni?eyoe
?aaeiaoeieiiiiaeoeiaaii ciiie, iia’ycaii ianaiia?aae ci ciiiith
i?iieeiinoi iaia?ai oa ia?a?iciiaeieii iiiia, yei aiaeia?athoue
?aaoeyoi?io ?ieue, ui i?eaiaeeoue aei aeenaaeaino aaaaoueio eeioeiieo
i?ioean?a. Iiaiaoaaiiy ia?aa?ao ?I* i?e c?inoaiii aeice ii?iiiiaiiy
iiaei aooe iaoiiaeaia ?aaeiaoeieiei aieeaii ia i?ioeane ?iciiciaaaiiy oa
o?ainaeoeoei? neaiaeia (ooieoeiiioaaiiy iiaa?oiaaeo ?aoeaioi?ia oa
eiiiiiaioia iieioinoiiiiceoeaeii? neaiaeueii? nenoaie i nenoaie oeAIO,
Na2+-, oinoieiiiaecaeaaeieo i?ioa?ieiiac oiui), a oaeiae i?ioeania
o?aine?eioei? i o?aineyoei? a ii?iiiiaieo ?ineeiieo eeioeiao. I/aaeaeii,
oea i?ecaaei aei aioaaiiicio o aei? ii?iiiiaiiy ii aiaeiioaiith aei
i?ioeano ?icaeoeo ?I*.
Aiae?c caeaaeiino? aianeo ii?iiiiaiiy aaeoa?ie (eiao?oe??io b2) a?ae
aeice ii?ii?iaiiy iieacaa, ui o anueiio canoiniaaiiio aeiciaiio
aeiaiaciii ii?ii?iaiiy aaeoa??e caeienithaaei iiaiaoth/ee aieea ia
?icaeoie ?I*. Oeae aieea c?inoaa ic caieueoaiiyi aeice ii?iiiiaiiy,
ni?e/eiyth/e iiaia aeieoaaiiy ?I* i?e aeicao, ui ia?aaeuoaaee 100 A?.
?en. 5 Caeaaei?noue a?ae aeice ii?ii?iaiiy aiane?a oaeoi??a
ii?ii?iaiiy ?ineei (b1), aaeoa??e-?iaeoeoi??a ?I* (b2) oa ?o aca?iiae??
(b1b2) o ?icaeoie ?I*
Ii cia/aiith eiaoioei?ioa b1b2 (?en. 5) ioeiithaaee nooiiiue
iaaaeeoeaiinoi eiia?iiaaii? aei? ii?iiiiaiiy ?ineei i aaeoa?ie ia
?aaeoeith iaae/ooeeainoi. I?e iaioeo aeicao ii?iiiiaiiy, eiee oeae
eiaoioei?io iaa iiceoeaia cia/aiiy, aca?iiaeiy oaeoi?ia iaea
aioaaiiinoe/iee oa?aeoa?: ii?iiiiaiiy ?ineei ni?eyei ?icaeoeo ?I*, a
ii?iiiiaiiy aaeoa?ie, iaaiaee, iaoiiaethaaei ?? iiaiaoaaiiy.
Ciaoiaeaeaiiy eiaoioei?ioa b1b2 i?e caieueoaiii aeice (iiiaae 100 A?) a
iaeanoi iaaaoeaieo cia/aiue icia/a?, ui oaeoi?e ii?iiiiaiiy ?ineei i
aaeoa?ie, aca?iiaeith/e, caeienithaaee iaaaoeaiee aieea ia ?icaeoie
?aaeoei?, oiaoi ooo iaa iinoea neia?ai/iee aoaeo.
Aeoiaey/e c iaea?aeaieo o iaoie ?iaioi ?acoeueoaoia, iaeaieueo e?eoe/iei
ii aiaeiioaiith aei aeii ii?iiiiaiiy neiae aaaaeaoe aoai iiaeoeoei? ?I*,
c eio?ei iia’ycaii i?ioeane ?iciiciaaaiiy iaoiaaio oa o?ainaeoeoei?
neaiaeia o eeioeiao ?ineeie-ainiiaea?y, ui ? aecia/aeueiei aeey anueiai
iiaeaeueoiai ?icaeoeo ?aaeoei?, yeee caeii/o?oueny caaeaaeeth
iioieiaaieo eeioei.
Iaea?aeai? a ?iaio? ?acoeueoaoe, ui noinothoueny ae?? aino?iai
ii?ii?iaiiy ia aca?iiae?th ?ineeie c o?oiiaoiaaiieie aaeoa??yie
iiaeooue ni?eyoe oi?ioaaiith iiaeo iai?yie?a aeine?aeaeaiue. Cie?aia, a
i?iaaaeaieo aeine?aeao canoiniaai? aeice ii?ii?iaiiy i?eaiaeeee aei
cia/iiai cieaeaiiy aeeoo?caeaoiino? aaeoa??e, c /ei iiaeia ianaiia?aae
iia’ycaoe ii?ooaiiy i?ioean?a ?ici?ciaaaiiy ?ineeiith iaoiaaio. I?ioa
caea?oueny oe?eeii i?eionoeiei, ui i?e o?ii?/iiio ii?ii?iaii? c
iecueeeie iiooaeiinoyie iiaeeiooeo aeic, ?nio? iaaiee ae?aiacii aeic,
eiee aiane?aeie ?iaeoeiaaieo iooaoe?eieo i?ioean?a oa ?aaoeyoi?ieo
ii?ooaiue, iaoiiaeaieo ae??th ?ii?coth/iai aei?ii?ithaaiiy, iiaeeea?
ci?ie aeni?an?? a?aeiia?aeieo aai?a (cie?aia, aai?a aa??oeaioiino?) o
ee?oeiao o?oiiaoiaai?a aac nooo?aiai cieaeaiiy ?o aeaeeaaiiy. Oea
iiaea i?eaanoe aei ci?i o neioac? oa aeni?an?? iaoiaaiii iieaeoe
ae?neoi??a, ui ?iaeoeothoue caoenio ?aaeoe?th ?ineeie. Oae? iaoiaaie ia
aoaeooue ?ici?ciaaaoeny ?ineeiieie ee?oeiaie ? iaoeiooue oiiae aeey
naiai ?icaeoeo a oeaieiao ?ineei, aeeeeeath/e iaoieia?/iee i?ioean.
Ca aeiiiiiaith ?aa?an?eiiai aiae?co aoei iieacaii, ui ia aoai?
?iaeoeoe?? ?aaeoe?? o?oiiaoiaaii? aaeoa???, ye? ?iaeoeoaaee ?? ?icaeoie,
oa ?ii?coth/a aei?ii?ithaaiiy o aenieeo aeicao ae?yee ye aioaaii?noe,
aca?iii iineaaethth/e aieea iaeia iaeiiai. Ia aeeeth/aii, ui i?e
iiaeoeoei? caaeaaei ?ineeiieo eeioei i?e ?I* oa ii?iiiiaiii ? caae?yieie
iaei? ? o? ae iaoai?cie. Oe? iaoai?cie iiaeooue aooe iiae?aieie aei
aiiioico, yeee niinoa??aaee a ?yae? aeiaaee?a i?e ?icaeoeo ?I* [Bachmair
et al., 1990; Greenberg, Ausubel, 1993; Dietrich et al., 1994; Greenberg
et al., 1994; Lamb, 1994; Vaux et al., 1994; Mittler et al., 1995].
AENIIAEE
1. Anoaiiaeaii, ui ?icaeoie ?aaeoe?? iaae/ooeeaino? (?I*)
i?enei?th?oueny i?e ii?iiiiaii? ?ineei o aenieeo aeicao (50-600 A?) aei
?iieoeyoe?? ?o o?oiiaoiaaiieie aaeoa??yie, ? cao?eio?oueny i?e
ii?ii?iaii? ?ineei i?ney ?o ?iieoeyoe??.
2. ?I* ? iaeaieueo ?aaeii/ooeeaith ia ii/aoeiaiio aoai? aca?iiaei?
?ineeiieo ee?oei c aaeoa??yie, eiee aiaeaoaa?oueny iiaeoeoeiy ?I* (30-60
oa i?ney ?iieoeyoe?? ?ineei aaeoa??yie). Ia oeueiio aoai? ?aaeoe?y
iiai?noth ?ia?aoaaeany i?e ii?ii?iaii? o aeicao, ui ia?aaeuoaaee 85 A?.
3. Ii?ii?iaiiy aaeoa??e i?eaiaeeoue aei ?ia?aoaaiiy ?icaeoeo ?I*, ui
aeaco? ia caeaaei?noue ia?aa?ao oe??? ?aaeoe?? a?ae ooieoe?iiaeueii?
aeoeaiino? aaeoa??aeueieo ee?oei. ?aaeoe?y ia ?icaeaaeany i?e
?iieoeyoe?? ?ineei aaeoa??aeueiith noniaic??th, ii?ii?iaiith o aeic? 50
A?, eiee aeaeeaaiiy aaeoa??e noaiiaeei i?eaeecii 15 %.
4. ?aa?an?eiee aiae?c ?acoeueoao?a aeine?ae?a ii aea/aiith eiia?iiaaii?
ae?? o?oiiaoiaaiieo aaeoa??e oa ?ii?coth/iai aei?ii?ithaaiiy iieacaa, ui
o aieea? aei?ii?ithaaiiy oa iaoiaaio ia aoai? ?iaeoeoe?? ?I* (1-3 aiae
i?ney ca?aaeaiiy ?ineei aaeoa??yie) ia? i?noea aioaaii?ci, a ia
iiaeaeueoeo aoaiao i?ioeano ?icaeoeo ?aaeoe?? – neia?a?ci.
5. Aecia/aeueiith aeey ?icaeoeo ?I* ? ooieoe?iiaeueia aeoeai?noue
aaeoa??aeueieo ee?oei, ui aeyaeythoue cia/io /ooeea?noue aei ae??
?aae?aoe?eiiai oaeoi?o.
IA?AEIE ?IAIO, IIOAEIEIAAIEO CA OAIITH AeENA?OAOeI?
1. Oeeiia TH.A., Iio??a I.I., Aoua I.I., A?iaeciinueeee Ae.I.
Canoinoaaiiy ?aaeiiaiieiai/ieo iaoiaeia aeey aeineiaeaeaiiy ?aaeoei?
iaae/ooeeainoi o ?ineei // Aeiiiaiaei IAI Oe?a?ie.- 1997. – N 11. – C.
175-177.
2. Oeeiia TH.A., Iio??a I.I., Aoua I.I., A?iaeciinueeee Ae.I.
Canoinoaaiiy ?aaeiiaiieiai/iiai iaoiaeo aeey aeineiaeaeaiiy iaoaiiciia
?aaeoei? iaae/ooeeainoi aei o?oiiaoiaai?a o ?ineei // Oiciieiaiy i
aiioiiiy eoeueoo?ieo ?ineei. -1998.- O. 30, N 3.- N. 187-193.
3. Oeeiia TH.A., Iio??a I.I., Aoua I.I., A?iaeciinueeee Ae.I.
Canoinoaaiiy iiiicoth/iai aei?iiiithaaiiy aeey aeineiaeaeaiiy aeaiaioia
aca?iiaei? ainiiaea?-iaoiaai ia i?eeeaaei ?aaeoei? iaae/ooeeainoi o
?ineei // Iie?iaiie. aeo?i. -1998.- O. 60, N 4. – N. 94-100.
4. Oeeiia TH.A., Iio??a I.I., Aoua I.I., A?iaeciinueeee Ae.I.
Aeineiaeaeaiiy aieeao iiiicoth/iai aei?iiiithaaiiy ia ?icaeoie no?aniai?
?aaeoei? ia i?eeeaaei ?aaeoei? iaae/ooeeainoi o ?ineei // Aeooaeueii
i?iaeaie oiciieiai? aiaeiiai ?aaeeio oa iinooinoieeinoi ?ineei. -E.: OIA
“Iiaeia?. oii. aaaioeiy”, 1997. – N. 35-37.
5. Oeeiia TH.A. Aeanoeainoi ?aaeoei? aiia?/ooeeainoi o ?ineei // Cai?iee
i?aoeue nooaeaioia oa anii?aioia Ee?anueeiai oi?aa?neoaoo ii. Oa?ana
Oaa/aiea. I?e?iaeie/i iaoee. /Aeione 2. – Ee?a. Oi-o.- Ee?a, 1995. – N.
19-27.
6. Oeeiia TH.A., A?iaec?inueeee Ae.I. Aieea aino?iai
aaiia-aei?iii-ithaaiiy ia ?aaeoeith aiia?/ooeeainoi o ?ineei // II
C`?cae ?aaeiiaiieiaia Oe?a?ie (Aeiii?iiao?ianuee, 22-24 aa?aniy 1995):
Oac. aeii. – Ee?a. 1995. – N. 199.
7. Shilina J.V., Mikheev A.N., Guscha M.I., Grodzinsky D.M. The
influence of gamma-radiation on hypersensitive reaction of Nicotiana
sanderae plant // 4th IUBMB Conference “The life and death of the cell”
(Edinburg International Conference Centre, 14-17 July 1996): Abstr.
//Biochemical Society Transactions.London. -1996.- Vol. 24, Part 4. – P.
C83.
8. Oeeiia TH.A. Aeineiaeaeaiiy ?aaeoei? iaae/ooeeainoi ?ineei ic
canoinoaaiiyi ?aaeiiaiieiai/iiai iaoiaeo // Aeooaeueii i?iaeaie
oiciieiai? ?ineei i aaiaoeee.- Ee?a, 1996. – N. 61-62.
9. Oeeeia TH.A., Ieoaaa A.I., Aoua I.E., A?iaeceineee Ae.I. Eco/aiea
?aaeoeee naa?o/oanoaeoaeueiinoe ?anoaiee n i?eiaiaieai
?aaeeiaeieiae/aneiai iaoiaea // 3 Nuecae ii ?aaeeaoeeiiiui
enneaaeiaaieyi (Iineaa, 14-17 ieoya?y 1997). – Oacenu aeieeaaeia.-
Ioueii, 1997. – Oii 1. – N. 169.
Oee?ia TH.A. ?aae?ia?ieia?/i? aeanoeaino? ?aaeoe?? iaae/ooeeaino? o
?ineei. – ?oeiien.
Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy eaiaeeaeaoa a?ieia?/ieo iaoe
ca niaoe?aeuei?noth 03.00.01 – ?aae?ia?ieia?y. -?inoeooo
aenia?eiaioaeueii? iaoieia??, iieieia?? ? ?aae?ia?ieia?? ?i. ?.?.
Eaaaoeueeiai IAI Oe?a?ie, Oe?a?inueeee iaoeiai-aeine?aeiee ?inoeooo
iieieia?? oa ?aae?ieia?? IIC Oe?a?ie, Ee?a, 1999.
Aeena?oaoe?y i?enay/aia ieoaiith aeine?aeaeaiiy ?aae?ia?ieia?/ieo
oa?aeoa?enoee ?aaeoe?? iaae/ooeeaino? (?I*) o ?ineei othothio Nicotiana
sanderae ca?aaeaieo aaeoa??yie Pseudomonas syringae pv. phaseolicola.
I?e aieea? aaiia-aei?ii?ithaaiiy a aeicao 50-600 A? iaea?eueoith
?aae?i/ooeea?noth oa?aeoa?ecoaaany aoai aca?iiae?? ?ineei ? aaeoa??e,
eiee a?aeaoaa?oueny ?iaeoeoe?y ?aaeoe?? ?ineeie-ainiiaea?y. Anoaiiaeaii,
ui aecia/aeueiith eaieith aeey ?icaeoeo ?I* ? ooieoe?iiaeueia
aeoeai?noue aaeoa??aeueieo ee?oei, ye? aeyaeythoue cia/io
?aae?i/ooeea?noue.
Eeth/ia? neiaa: ?aaeoe?y iaae/ooeeaino? o ?ineei, aaiia-aei?ii?ithaaiiy,
othothi, inaaaeiiiian, ?aae?i/ooeea?noue.
Oeeeia TH.A. ?aaeeiaeieiae/aneea naienoaa ?aaeoeee
naa?o/oanoaeoaeueiinoe o ?anoaiee. – ?oeiienue.
Aeena?oaoeey ia nieneaiea o/aiie noaiaie eaiaeeaeaoa aeieiae/aneeo iaoe
ii niaoeeaeueiinoe 03.00.01 – ?aaeeiaeieiaey. – Einoeooo
yenia?eiaioaeueiie iaoieiaee, iieieiaee e ?aaeeiaeieiaee ei.
?.A.Eaaaoeeiai IAI Oe?aeiu, Oe?aeineee iao/ii-enneaaeiaaoaeueneee
einoeooo iieieiaee e ?aaeeieiaeee IIC Oe?aeiu, Eeaa, 1999.
Aeena?oaoeey iinayuaia aii?ino enneaaeiaaiey ?aaeeiaeieiae/aneeo
oa?aeoa?enoee ?aaeoeee naa?o/oanoaeoaeueiinoe (NA*) o ?anoaiee oaaaea
Nicotiana sanderae ca?aaeaiiuo aaeoa?eyie Pseudomonas syringae pv.
phaseolicola. I?e aicaeaenoaee aaiia-eceo/aiey a aeicao 50-600 A?
iaeaieueoae ?aaeei/oanoaeoaeueiinoueth oa?aeoa?eciaaeny yoai
acaeiiaeaenoaey ?anoaiey e aaeoa?ee, eiaaea i?ienoiaeeo eiaeoeoeey
?aaeoeee ?anoaiey-oicyeia. Onoaiiaeaii, /oi ii?aaeaeythuei caaiii aeey
?acaeoey NA* yaeyaony ooieoeeiiaeueiay aeoeaiinoue aaeoa?eaeueiuo
eeaoie, eioi?ua i?iyaeytho cia/eoaeueioth ?aaeei/oanoaeoaeueiinoue.
Eeth/aaua neiaa: ?aaeoeey naa?o/oanoaeoaeueiinoe o ?anoaiee,
aaiia-eceo/aiea, oaaae, inaaaeiiiian, ?aaeei/oanoaeoaeueiinoue.
J.V.Shilina. The radiobiology property of hypersensitive reaction of
plants.- Manuscript.
Thesis for obtaining scientific degree of Candidate of biological
sciences on speciality 03.00.01 – radiobiology. – Institute of
experimental pathology, onkology and radiobiology, National Academy of
Sciences of Ukraine, Ukrainian Research Institute of Oncology and
Radiology Ministry of Health Care of Ukraine, Kiyv, 1999.
The dissertation is devoted to investigation on radiobiological
characteristics of the hypersensitive reaction (HR) of tobacco plants
Nicotiana sanderae against Pseudomonas syringae pv. phaseolicola 291.
The time-dependent and dose-dependent nature of gamma-irradiation
effects on different stages of the HR development was established. The
first stage of bacteria-plant interaction (30-60 min after bacteria
inoculation) that induction of the HR was occured, was the most
radiosensitive. It is shown that the ingibition effect of radiation on
this stage could be conditioned by fall functional activity of banteria
cells which were to be very radiosensitive.
Key words : hypersensitive reaction of plants, pseudomonads, tobacco,
gamma-irradiation, radiosensitivity.
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