S.N. Elansky, L. Yu. Kokaeva, N.V. Statsyuk, Yu.T. Dyakov
Selelekela
Oomycete Phytophthora infestans (Mont.) de Bary, sesosa sa blight ea morao, lefu la bohlokoa ka ho fetisisa moruong la litapole le tamati, e hohetse tlhokomelo e haufi ea bafuputsi ba tsoang linaheng tse sa tšoaneng ka lilemo tse fetang lekholo le halofo. Ka tšohanyetso e hlaha Europe bohareng ba lekholo la bo19 la lilemo, e ile ea baka seoa sa litapole se ileng sa sala mohopolong oa meloko e mengata.
Ho tla fihlela kajeno, hangata e bitsoa “Irish tlala mushroom.” Hoo e ka bang lilemo tse lekholo ka mor'a lefu la seoa la pele, ho ile ha sibolloa mefuta ea litapole tse hlaha tsa Mexico tse hananang le blight ea morao-rao, mekhoa ea ho li tšela ka litapole tse lenngoeng li ile tsa ntlafatsoa (Muller, 1935), 'me mefuta ea pele ea morao-rao e hanyetsanang le blight e ile ea fumanoa (Pushkarev, 1937) . Leha ho le joalo, kapele ka mor'a ho qala temo ea bona ea khoebo, merabe ea likokoana-hloko tsa morao-rao tse neng li le kotsi ho mefuta e manganga li ile tsa bokellana. 'me liphatsa tsa lefutso tse ncha tsa ho hanyetsa tse kentsoeng mefuteng e tsoang ho litapole tse hlaha tsa Mexico li ile tsa qala ho lahleheloa ke katleho ea tsona ka potlako.
Ho hloleha ho sebelisa khanyetso ea monogenic (tse otlolohileng) ho qobelletse barui ho batla mekhoa e rarahaneng ea ho sebelisa khanyetso e sa tobang ea polygenic (ho rapama). Lilemong tsa morao tjena, merabe e mabifi e qalile ho bokellana lichabeng tse itseng tsa likokoana-hloko, e leng se bakang khoholeho ea ho hanyetsa ka mokhoa o sa tobang. Ho hlaha ha mefuta e hananang le fungicide ho bakile mathata ts'ebelisong ea lihlahisoa tsa k'hemik'hale tse sireletsang litapole.
Ka lebaka la phapang e kholo lipakeng tsa li-oomycete le li-fungus ka sebopeho sa lik'hemik'hale, sebopeho sa ultrastructure le metabolism, li-fungicides, haholo-holo tsa systemic, tse sebelisetsoang ho sireletsa limela ho tsoa ho mafu a mangata a fungal, ha li sebetse khahlano le oomycetes.
Ka hona, ts'ireletsong ea lik'hemik'hale khahlano le blight ea morao, ho ne ho sebelisoa ho fafatsa ka makhetlo a mangata (ho fihla ho makhetlo a 12 ka nako kapa ho feta) ka litokisetso tsa ho kopana ka bophara. Mohato oa phetoho e ne e le tšebeliso ea li-phenylamide, tse chefo ho oomycetes le ho ajoa ka mokhoa o hlophisitsoeng limela. Leha ho le joalo, tšebeliso ea bona e atileng kapele e ile ea lebisa ho bokelleng ha mefuta e hanyetsanang ho batho ba fungal (Davidse et al., 1981), e leng se ileng sa thatafatsa ts'ireletso ea limela haholo. P. infestans ke eona feela likokoana-hloko tsa sebaka se futhumetseng, tšenyo e ke keng ea fokotsoa maemong a temo ntle le tšebeliso ea lik'hemik'hale tsa tšireletso (Van Bruggen, 1995).
Lintlha tse ka holimo li hlalosa tlhokomelo e kholo eo bafuputsi ba tsoang linaheng tse fapaneng ba e lefang thutong ea P. infestans populations, matla a lipalo tsa bona le liphatsa tsa lefutso, hammoho le mekhoa ea liphatsa tsa lefutso tsa ho fetoha.
Phetoho ea bophelo ba P. INFESTANS
Oomycete Phytophthora infestans e hlahisa mycelium ea intercellular le haustoria ka hare ho makhasi a litapole. Ka ho fepa ka mahlaku a makhasi, e etsa hore ho thehoe matheba a lefifi, a fetohang a ntšo 'me a bola ha a le mongobo. Ka tšenyo e kholo, lekhasi lohle lea shoa. Ka mor'a nako ea ho fepa, limela li thehoa holim'a mycelium - sporangiophores - e hōlang ka ntle ka stomata. Boemong ba leholimo bo mongobo ba etsa lesela le lesoeu ho pota-pota matheba a ka tlas'a makhasi. Qetellong ea li-sporangiophores, li-zoosporangia tse bōpehileng joaloka lemone li thehoa, tse senyang 'me li tsamaisoa ke lipula tsa pula (setšoantšo sa 1). Ho kena ka har'a marotholi a metsi holim'a lekhasi la litapole, sporangia e mela ho li-zoospores tse 6-8, tseo, kamora nako ea ho sisinyeha, li pota-potileng, li koahetsoe ka khetla ebe li mela ka har'a tube ea kokoana-hloko. Lehlomela le phunyeletsa lesela la makhasi ka stomata. Tlas'a maemo a itseng, sporangium e ka hola ka tšupu ea kokoana-hloko ka ho toba letlapeng la makhasi. Tlas'a maemo a matle, nako ea ho tloha tšoaetsong ho fihlela ho thehoa ha sporulation e ncha ke matsatsi a 3-4 feela.
Hang ha e se e le fatše 'me e sefshoa mobung, sporangia e khona ho tšoaetsa li-tubers. Li-tubers tse amehang haholo li bola nakong ea polokelo; libakeng tse amehileng ka bofokoli tšoaetso e ka 'na ea tsoela pele ho fihlela sehla se latelang. Ho feta moo, sesosa sa blight sa morao-rao se ka phela mariha ka mokhoa oa li-oospores (li-spores tsa thobalano tse teteaneng) mobung oa maloanlahla a limela le peo ea tamati. Li-Oospores li thehoa lithong tsa limela tse phelang ha mefuta ea mefuta e fapaneng ea ho nyalana e kopana le mongobo o feteletseng. Nakong ea selemo, mefuta ea li-asexual sporulation ho li-tubers tse lenngoeng tse nang le tšoaetso le litšila tsa limela tse nang le li-oospores li kena mobung ebe li baka tšoaetso ea makhasi a tlase a limela. Maemong a mang, mycelium e ka hola ho tloha ho tuber e tšoaelitsoeng haufi le karolo e tala ea semela 'me hangata e hlaha karolong e ka holimo ea kutu.
Phapang e tebileng pakeng tsa oomycete le li-fungus tse ngata ke ho ata ha diplophase bophelong ba bona ka gametic meiosis le ho mela ha li-zygotes (oospores) ntle le ho fokotsa karohano ea nyutlelie. Tšobotsi ena, hammoho le dipolar heterothallism, e nkang sebaka sa bisexuality, e ka bonahala e etsa hore ho khonehe ho sebelisa mekhoa ea oomycetes e ntlafalitsoeng bakeng sa ho ithuta ka bongata ba li-eukaryote tse phahameng (tlhahlobo ea panmixia le karohano ea baahi, phallo ea liphatsa tsa lefutso tsa intra- le tse ling, joalo-joalo). Leha ho le joalo, lintlha tse tharo li thibela mekhoa ena hore e se ke ea fetisetsoa ka ho feletseng thutong ea batho ba P. infestans.
1. Hammoho le hybrid oospores, self-fertile le parthenogenetic oospores li thehoa ka bongata (Fife, Shaw, 1992; Anikina et al., 1997a; Savenkova, Cherepnikoba-Anirina, 2002; Smirnov, 2003), le ho etsoa khafetsa ha bona. e ka ba e lekaneng ho susumetsa liphetho tsa liteko.
2. Ts'ebetso ea thobalano ho P. infestans e etsa tlatsetso e sa reng letho ho matla a baahi, hobane li-fungus li ikatisa haholo-holo ka limela tsa limela, ho etsa liphello tsa tlhahlobo ea PCR ho sebelisa li-primers tse khethehileng tse fetang 90% tse tsamaellanang le liphello tsa tlhahlobo ea mofuta oa ho nyalana ka mokhoa o tloaelehileng. tsela ea phepo e nepahetseng. nako ea ho hōla meloko e 'maloa ea asexual sporulation (polycyclic tsoelo-pele ea lefu lena). Li-Oospores li phetha karolo ea bohlokoa ho bolokeng lintho tse phelang nakong eo ho se nang limela tse tala (mariha) le ts'oaetso ea mantlha ea lipeo. Joale, nakong ea lehlabula, ho hlahisa li-clonal ho etsahala le ho eketseha kapa, ka lehlakoreng le leng, ho fokotseha ha palo ea li-clone ka bomong tse hlahang ka lebaka la ho kopana hape ka thobalano, e khethiloeng haholo-holo ka khetho ea ho lekana haholoanyane. Ka hona, karo-karolelano ea li-clone ka bomong ho baahi qalong le qetellong ea epiphytoty e ka fapana ka ho feletseng.
3. Potoloho e hlalositsoeng e tloaelehile bakeng sa baahi ba matsoalloa a P. infestans naheng ea habo bona - Amerika Bohareng. Libakeng tse ling tsa lefats'e, ts'ebetso ea thobalano e ne e sa tsejoe ka lilemo tse fetang 100; Phetoho ea bophelo e ne e le monate ka ho feletseng, 'me ho ata ho ne ho tsepamisitse maikutlo tlhahong: ts'oaetso e tsoang ho li-tubers tse lenngoeng tse lenngoeng tse nang le tšoaetso e ile ea hasana makhasi, ea etsa motheo oa lefu lena, o ka kopanang le tsoelo-pele e kholo ea lefu lena.
Ka hona, libakeng tse ling ho ka 'na ha e-ba le phetoho ea potoloho ea thobalano le ea ho kopanela liphate, ha ho tse ling ho ka ba le potoloho ea asexual feela.
Tšimoloho ea P. INFESTANS
P. infestans e ile ea hlaha Europe qetellong ea halofo ea pele ea lekholo la bo1991 la lilemo. Kaha litapole li hlaha ka leboea-bochabela ho Amerika Boroa, ho ne ho nahanoa hore kokoana-hloko eo e ile ea tlisoa ho tloha moo ho ea Europe nakong ea ho phahama ha letsoai la Chile. Leha ho le joalo, liphuputso tse entsoeng seteisheneng sa litapole sa Rockefeller Center Phuleng ea Toluca (Mexico) li ile tsa qobella ho shebisisa ntlha ena ea maikutlo (Niederhauser, 1993, XNUMX).
1. Phuleng ea Toluca, mefuta ea litapole tsa li-tuberous ea sebakeng seo (Solanum demissum, S. bulbocastanum, joalo-joalo) e na le lihlopha tse fapaneng tsa liphatsa tsa lefutso tsa ho hanyetsa tse otlolohileng tse kopantsoeng le boemo bo phahameng ba ho hanyetsa ka mokhoa o sa tobang, e leng se bontšang ho iphetola ha nako e telele le likokoana-hloko. Mefuta ea Amerika Boroa, ho kenyelletsa le litapole tse lengoang, ha e na liphatsa tsa lefutso tsa ho hanyetsa.
2. Phuleng ea Toluca, libaka tse ka thōko tse nang le mefuta ea ho nyalana ea A1 le A2 li fumanoa, ka lebaka leo palo ea baahi ba mefuta e fapaneng ea P. infestans e atileng; ha e ntse e le naheng ea habo ea litapole tse lengoang, Amerika Boroa, likokoana-hloko li ata ka clonally.
3. Phula ea Toluca e ba le mafu a seoa a matla a selemo le selemo. Ka hona, har'a bafuputsi ba Amerika Leboea (Univesithi ea Cornell), ho na le maikutlo a tiileng mabapi le Mesoamerica (Amerika Bohareng) e le naha ea habo ea litapole tsa morao-rao (Goodwin et al., 1994).
Bafuputsi ba Amerika Boroa ha ba kopanele maikutlo ana. Ba lumela hore litapole tse lengoang le likokoana-hloko tsa bona tsa P. infestans li na le naha e tloaelehileng - Amerika Boroa Andes. Ba tšehelitse maikutlo a bona ka lithuto tsa limolek'hule mabapi le tlhahlobo ea DNA polymorphisms ea mitochondrial genome (mtDNA) le liphatsa tsa lefutso tsa nyutlelie RAS le β-tubulin (Gomez-Alpizar et al., 2007). Ba ile ba bontša hore mefuta e neng e bokelloa libakeng tse sa tšoaneng ho pota lefatše e simolohile melokong e meraro e fapaneng ea baholo-holo, eo kaofela ha eona e neng e fumanoa Andes ea Amerika Boroa. Li-haplotype tsa Andes ke litloholo tsa meloko e 'meli: lihlopha tsa moloko oa khale ka ho fetisisa oa mtDNA li fumanoa li-nightshades tse hlaha ho tloha karolong ea Anarrhicomenum naheng ea Ecuador, lihlopha tsa leloko la bobeli li tloaelehile ho litapole, tamati le li-nightshades tse hlaha. Ho Toluca, esita le li-haplotype tse sa tloaelehang li nkiloe molokong o le mong feela, ka ho fetoha ha liphatsa tsa lefutso tsa mefuta ea Toluca (maqhubu a tlaase a allelic a libaka tse ling tse fapaneng) a fana ka maikutlo a matla a mothehi ka lebaka la ho hoholeha ha morao tjena.
Ho phaella moo, mofuta o mocha oa P. andina, morphologically le liphatsa tsa lefutso tse tšoanang le P. infestans, li ile tsa fumanoa Andes, eo, ho ea ka bangoli, e bontšang Andes e le sebaka se seholo sa ho khetholla mofuta oa Phytophthora. Qetellong, Europe le United States, baahi ba P. infestans ba kenyelletsa meloko ea Andes ka bobeli, ha Toluca e le 'ngoe feela.
Sengoliloeng sena se hlahisitse karabelo ho tsoa ho sehlopha sa bafuputsi ba tsoang linaheng tse fapaneng ba ileng ba etsa liteko tse ngata ho ntlafatsa thuto e fetileng (Goss et al., 2014). Mosebetsing ona, pele, tatelano ea DNA ea microsatellite e rutang haholoanyane e ile ea sebelisoa ho ithuta DNA polymorphisms; ea bobeli, bakeng sa tlhahlobo ea lihlopha, litsela tsa ho falla, nako ea ho fapana ha baahi, joalo-joalo. ho ile ha sebelisoa mehlala e tsoetseng pele haholo (lipalo-palo tsa F, likhakanyo tsa Bayesian, joalo-joalo) 'me, ea boraro, papiso e ne e sebelisoa eseng feela le mefuta ea Andean P. andina, eo tlhaho ea lebasetere e ileng ea thehoa (P. infestans x Phytophthora sp.) , empa hape le Mexican endemics P. mirabilis, P. Ipomoeae, le Phytophthora phaseoli, tse nang le liphatsa tsa lefutso tse haufi le P. infestans 'me ke tsa clade e tšoanang (Kroon et al., 2012). Ka lebaka la litlhahlobo tsena, ho ile ha bontšoa ka ho hlaka hore karolo ea motso oa sefate sa phylogenetic sa mefuta eohle ea mofuta oa Phytophthora e nkiloeng thutong, ntle le ea lebasetere P. andina, ke ea mefuta ea Mexico, 'me phallo ea ho falla e na le tataiso Mexico - Andes, 'me eseng ka tsela e fapaneng,' me qaleho ea eona e lumellana le bokolone ba Europe ba Lefatše le Lecha (lilemo tse 300-600 tse fetileng). Ka hona, ho hlaha ha mefuta ea P. infestans, e khethehileng bakeng sa tšenyo ea litapole, e etsahetse setsing sa bobeli sa liphatsa tsa lefutso tsa ho thehoa ha li-nightshades tsa tuberous, i.e. Amerika Bohareng.
Genome of P. infestans
Ka 2009, sehlopha sa machaba sa bo-rasaense se ile sa tatellana le genome e feletseng ea P infestans (Haas et al, 2009), eo boholo ba eona e neng e le 240 Mb. Sena se feta ka makhetlo a 'maloa ho feta mefuta e amanang haufi-ufi ea P. sojae (95 Mb), e bakang ho bola ha metso ea soya, le P. Ramorum (65 Mb), e amang mefuta ea lifate tsa bohlokoa joalo ka oak, beech le tse ling. Lintlha tse fumanoeng li bontšitse hore genome e na le palo e kholo ea likopi tsa ho pheta-pheta tatellano - 74%. Genome e na le liphatsa tsa lefutso tsa protheine tse 17797, tseo bongata ba tsona e leng liphatsa tsa lefutso tse amehang lits'ebetsong tsa cellular, ho kenyeletsoa replication ea DNA, transcript le phetolelo ea liprotheine.
Papiso ea liphatsa tsa lefutso tsa mofuta oa Phytophthora e senotse mokhatlo o sa tloaelehang oa genome, o nang le li-block tsa tatellano ea liphatsa tsa lefutso tse bolokiloeng, tseo ho tsona liphatsa tsa lefutso li batlang li le holimo 'me litaba tsa tatellano e pheta-phetoang li batla li le tlase, le libaka tse nang le tatelano ea liphatsa tsa lefutso tse sa bolokehang. , e nang le matla a tlaase a liphatsa tsa lefutso le litaba tse phahameng tsa libaka tse pheta-phetoang. Li-blocks tse bolokiloeng li etsa 70% (12440) ea liphatsa tsa lefutso tsa protheine-coding ho P. infestans. Ka har'a li-blocks tse bolokiloeng, liphatsa tsa lefutso hangata li fumaneha haufi le sebaka se mahareng sa 604 bp. Libakeng tse pakeng tsa li-blocks tse bolokiloeng, sebaka sa intergenic se seholoanyane (3700 bp) ka lebaka la keketseho ea sekhahla sa likarolo tse pheta-phetoang. Liphatsa tsa lefutso tse fetohang ka potlako li fumaneha libakeng tse nang le liphatsa tsa lefutso tse ngata.
Tlhahlobo ea tatelano ea liphatsa tsa lefutso tsa P. infestans e bontšitse hore hoo e ka bang karolo ea boraro ea genome ke ea likarolo tse fetisetsoang. P. infestans genome e na le malapa a transposon a fapaneng haholo ho feta a mang a tsejoang. Boholo ba li-transposon tsa P. infestans ke tsa lelapa la Gypsy.
P. infestans genome e senotse palo e kholo ea malapa a liphatsa tsa lefutso a amehang ho pathogenesis. Karolo ea bohlokoa ea tsona e kenyelletsa liprotheine tse sebetsang tse fetolang physiology ea semela se amohelang le ho kenya letsoho ho ts'oaetso ea eona. Li oela likarolong tse peli tse pharaletseng: li-apoplastic effect, tse sebetsang libakeng tsa intercellular (apoplasts) le li-cytoplasmic effectors, tse kenang liseleng ka haustoria. Li-apoplastic effect li kenyelletsa li-enzyme tsa hydrolytic tse patiloeng tse kang proteases, lipases le glycosylases tse senyang lisele tsa semela; li-inhibitors tsa li-enzyme tse sireletsang limela le chefo ea necrotizing e kang Nep1-like proteins (NPLs) le Pcf-like small cysteine-rich proteins (SCRs).
Liphatsa tsa lefutso tsa P. infestans li ngata 'me hangata li kholoanyane ka boholo ha li bapisoa le liphatsa tsa lefutso tsa nonpathogenic. Liphello tse tsebahalang haholo tsa cytoplasmic ke RXLR le Crinkler (CNR). Liphello tse tloaelehileng tsa cytoplasmic bakeng sa oomycetes ke liprotheine tsa RXLR. Liphatsa tsa lefutso tsohle tsa RXLR tse fumanoeng ho tla fihlela joale li na le sehlopha sa amino-terminal Arg-XLeu-Arg, moo X e emelang amino acid. Phuputso e ile ea fana ka maikutlo a ho ba teng ha liphatsa tsa lefutso tsa 563 RXLR genome ea P. infestans, e leng 60% ho feta ea P. sojae le P. ramorum. Hoo e ka bang halofo ea liphatsa tsa lefutso tsa RXLR ho P. infestans genome ke mefuta e khethehileng. Li-effectors tsa RXLR li na le mefuta e mengata e fapaneng ea tatellano. Har'a bona, malapa a le mong a maholo le a manyenyane a 150 a ile a khetholloa. Ho fapana le proteome ea mantlha, liphatsa tsa lefutso tsa RXLR hangata li fumaneha libakeng tse futsanehileng, tse pheta-phetoang tsa genome. Lintho tse ka fetisetsoang tse etsang hore libaka tsena li be matla li khothalletsa ho kopana hape liphatseng tsena tsa lefutso.
Cytoplasmic CRN effectors e ile ea qala ho khetholloa ho P. infestans transcripts encoding peptide e susumetsang necrosis ea lisele tsa semela. Ho tloha ha ba sibolloa, ha ho tsejoe hakaalo ka lelapa la baetsi bana. Tlhahlobo ea P. infestans genome e senotse lelapa le leholo la liphatsa tsa lefutso tsa 196 CRN, tse kholo haholo ho feta tsa P. sojae (100 CRN) le P. ramorum (19 CRN). Joalo ka RXLR, li-CRN ke liprotheine tsa modular 'me li na le sebaka se bolokiloeng haholo sa N-terminal LFLAK (50 amino acids) le sebaka se haufi sa DWL se nang le mefuta e fapaneng ea lefutso. Boholo ba li-CRN (60%) li na le peptide ea matšoao.
Bokhoni ba li-CRN tse fapaneng tsa ho senya lits'ebetso tsa cellular tsa semela bo ithutoa. Ha ho hlahlojoa necrosis ea limela, ho tlosoa ha liprotheine tsa CRN2 ho entse hore ho khonehe ho khetholla sebaka sa C-terminal se nang le 234 amino acid (maemo a 173-407, sebaka sa DXG) se bakang lefu la sele. Tlhahlobo ea liphatsa tsa lefutso tsa P. infestans CRN e senotse libaka tse 'nè tse fapaneng tsa C-terminal tse bakang lefu la lisele ka har'a semela. Tsena li kenyelletsa libaka tsa pele tse khethiloeng tsa DC (P. infestans e na le liphatsa tsa lefutso tse 18 le li-pseudogenes tse 49), hammoho le libaka tsa D2 (14 le 43) le DBF (2 le 1), tse tšoanang le protheine kinases. Liprotheine tsa marang-rang tsa CRN tse hlahisoang limela li tsoela pele (ha ho se na li-peptide tsa pontšo) ka seleng ea semela le ho susumetsa lefu la sele ka mochine oa intracellular. Tatelano e 'ngoe e nang le domain name ea 255 CRN e kanna ea se sebetse joalo ka liphatsa tsa lefutso.
Keketseho ea palo le boholo ba malapa a liphatsa tsa lefutso tsa RXLR le CRN ho ka etsahala hore ebe e ne e le ka lebaka la recombination e seng ea allelic homologous le phetiso ea liphatsa tsa lefutso. Ho sa tsotellehe 'nete ea hore genome e na le palo e kholo ea likarolo tse sebetsang tsa transposable, ho ntse ho se na bopaki bo tobileng ba phetisetso ea liphatsa tsa lefutso.
Mekhoa e sebelisoang ho ithuta sebopeho sa baahi
Boithuto ba sebopeho sa liphatsa tsa lefutso tsa baahi hajoale bo ipapisitse le tlhahlobo ea litso tse hloekileng tsa mefuta ea eona. Tlhahlobo ea baahi ntle le ho arola litso tse hloekileng le eona e etsoa ka sepheo se itseng, joalo ka, mohlala, ho ithuta mabifi a baahi kapa boteng ba mefuta e hananang le fungicide ho eona (Filippov et al., 2004, Derevyagina et al., 1999). Mofuta ona oa lipatlisiso o kenyelletsa tšebeliso ea mekhoa e khethehileng, eo tlhaloso ea eona e fetang tekanyo ea tlhahlobo ena. Bakeng sa tlhahlobo ea papiso ea mefuta, ho sebelisoa mekhoa e mengata, e ipapisitse le tlhahlobo ea sebopeho sa DNA le thutong ea lipontšo tsa phenotypic. Ha ho etsoa tlhahlobo ea papiso ea baahi, motho o tlameha ho sebetsana le palo e kholo ea batho ba itšehlang thajana, e fanang ka litlhoko tse itseng mekhoeng e sebelisoang. Ka nepo, ba lokela ho fihlela litlhoko tse latelang (Cooke, Lees, 2004, Mueller, Wolfenbarger, 1999):
- e be theko e tlaase, e bonolo ho e kenya ts'ebetsong, ha e hloke nako ea bohlokoa, 'me e itšetlehile ka theknoloji e fumanehang phatlalatsa (mohlala, PCR);
- e tlameha ho hlahisa palo e lekaneng ea likarolo tse ikemetseng tsa matshwao;
- ho ba le ho ikatisa ho hoholo;
- sebelisa palo e fokolang ea lisele tse ntseng li hlahlojoa;
- e-ba e tobileng ho substrate (tšilafalo e teng setso ha ea lokela ho ama liphello);
- ha ba hloke tšebeliso ea mekhoa e kotsi le lik'hemik'hale tse chefo haholo.
Ka bomalimabe, ha ho na mekhoa e lumellanang le litekanyetso tsohle tse ka holimo. Bakeng sa thuto ea papiso ea mefuta, mekhoa e se e sebelisoa ho ipapisitse le tlhahlobo ea litšobotsi tsa phenotypic: virulence ho mefuta ea litapole le tamati (peiso ea litapole le tamati), mofuta oa ho nyalana, spectra ea peptidase le glucose-6-phosphate isomerase isoenzymes, le tlhahlobo ea mali. DNA sebopeho: bolelele polymorphism restriction fragment analysis (RFLP), eo hangata e tlatselletsoang ka RG 57 hybridization probe, microsatellite repeat analysis (SSR le InterSSR), amplification with random primers (RAPD), restriction fragment amplification (AFLP), amplification with primers Homologous to transposable element (mohlala, Inter SINE PCR), boikemisetso ba mitochondrial DNA haplotypes.
Litlhaloso tse khutšoane tsa mekhoa ea ho ithuta ka papiso ea mefuta e sebelisoang ho sebetsa le P. infestans
Litšobotsi tsa phenotypic marker
Mabelo a "litapole".
Peiso ea "litapole" ke lesupa le sebelisoang khafetsa. Peiso ea "litapole tse bonolo" e na le lefutso le le leng la "tapole" e nang le lefu la "potata", merabe "e rarahaneng" e na le bonyane tse peli. Black et al. (1953), ha a akaretsa lintlha tsohle tse fumanehang ho bona, o ile a tiisa hore morabe oa Phytophthora o khona ho hlasela limela tse nang le lefutso la ho hanyetsa / liphatsa tsa lefutso tse tsamaellanang le lefutso / liphatsa tsa lefutso tsa P. infestans, le merabe e sibolotsoeng 1, 2 , 3 le 4 tse tšoaetsang limela ka liphatsa tsa lefutso tsa R1, R2, R3 le R4, ka ho latellana, i.e. Tšebelisano pakeng tsa likokoana-hloko le moeti e etsahala ho latela molao-motheo oa "gene to gene". Ho feta moo, Black, ka ho nka karolo ha Gallegly le Malcolmson, o ile a sibolla mefuta ea bohanyetsi R5, R6, R7, R8, R9, R10 le R11, hammoho le merabe ea bona e tsamaellanang (Black, 1954; Black & Gallegly, 1957; Malcolmson & Black , 1966 Malcolmson, 1970).
Ho na le letoto le pharaletseng la data mabapi le sebopeho sa merabe sa pathogen ho tsoa libakeng tse fapaneng. Ntle le ho sekaseka lintlha tsena ka botlalo, re tla bonts'a mokhoa o akaretsang: moo mefuta e nang le mefuta e mecha ea ho hanyetsa kapa metsoako ea eona e sebelisitsoeng, qalong ho ne ho e-na le ho fokola ho itseng ha blight ea morao-rao, empa merabe e nang le liphatsa tsa lefutso tse tsamaellanang e ile ea hlaha 'me ea khethoa. 'me ho qhoma ha blight ea morao ho ile ha qala hape. Boloetse bo khethehileng khahlanong le liphatsa tsa lefutso tse 4 tsa pele (R1-R4) bo ne bo sa bonoa ka seoelo likolekeng tse bokelletsoeng pele ho hlahisoa mefuta e nang le liphatsa tsa lefutso ho lengoa, empa palo ea likokoana-hloko e ile ea eketseha haholo ha kokoana-hloko e jara mefuta ena ea lefutso. Genes 5-11, ho fapana le hoo, e ne e fumanoa hangata ka har'a likoleke (Shaw, 1991).
Phuputso ea karo-karolelano ea merabe e fapaneng ho pholletsa le nako ea ho hōla, e entsoeng ho elella bofelong ba lilemo tsa bo-1980, e bontšitse hore qalong ea tsoelo-pele ea lefu lena, li-clones tse nang le bohale bo tlaase le liphatsa tsa lefutso tse 1-2 li atile har'a baahi.
Ho feta moo, ha blight ea morao e ntse e tsoela pele, palo ea li-clones tsa pele e fokotseha 'me palo ea mabelo a "rarahaneng" a nang le mabifi a phahameng a eketseha. Ketsahalo ea ho qetela e fihla ho 100% qetellong ea nako. Ha u boloka li-tubers, ho na le ho fokotseha ha mabifi le tahlehelo ea liphatsa tsa lefutso tsa motho ka mong. Matla a ho fetola clone a ka hlaha ka tsela e fapaneng ka mefuta e fapaneng (Rybakova & Dyakov, 1990). Leha ho le joalo, lithuto tsa rona tse entsoeng ka 2000-2010 li bontšitse hore merabe e rarahaneng e fumanoe ho tloha qalong ea epiphytoty har'a mefuta e arohaneng le litapole le tamati. Sena se ka etsahala ka lebaka la liphetoho ho baahi ba P. infestans Russia.
Ka 1988-1995, ketsahalo ea "merabe e phahameng ka ho fetisisa" e nang le mefuta eohle ea likokoana-hloko kapa hoo e batlang e le libakeng tse sa tšoaneng e fihlile ho 70-100%. Boemo bona bo ile ba hlokomeloa, ka mohlala, Belarus, libakeng tsa Leningrad, Moscow, North Ossetia le Jeremane (Ivanyuk et al., 2002a, 2002b; Polityko, 1994; Schober-Butin et al., 1995).
Mabelo a "tamati".
Mefuteng ea tamati, ho fumanoe liphatsa tsa lefutso tse peli feela tsa ho hanyetsa blight kamora nako: Ph2 (Gallegly & Marvell, 1) le Ph1955 (Al-Kherb, 2). Joaloka tabeng ea merabe ea litapole, tšebelisano pakeng tsa tamati le P. infestans e etsahala ho latela molao-motheo oa "gene for gene". Race T1988 e tšoaetsa mefuta e se nang liphatsa tsa lefutso (mefuta e sebelisoang haholo indastering), morabe T0 e tšoaetsa mefuta e fapaneng ka mofuta oa Ph1 (Ottawa), morabe oa T1 o tšoaetsa mefuta e fapaneng ka mofuta oa Ph2.
Naheng ea Russia, hoo e batlang e le T0 feela e fumanoe ka litapole; ka tamati, T0 e ne e le teng qalong ea sehla, empa hamorao e ile ea nkeloa sebaka ke peiso ea T1 (Dyakov et al., 1975, 1994). Kamora 2000, T1 ea litapole e ile ea qala ho hlaha lichabeng tse ngata qalong ea epiphytoty. Naheng ea Amerika, mefuta e seng kotsi ho tamati, hammoho le merabe ea T0, T1 le T2, e fumanoe litapole, ha T1 le T2 li le ngata ho tamati (Vartanian & Endo, 1985; Goodwin et al., 1995).
Mofuta oa ho nyalana
Ho khanna thuto, mefuta ea teko (referense) e nang le mefuta e tsebahalang ea ho nyalana - A1 le A2 - ea hlokahala. Teko e itšehla thajana e jaloa le bona ka bobeli lijana tsa Petri tse nang le oat agar medium. Kamora ho qeta matsatsi a 10, lijana li hlahlojoa bakeng sa ho ba teng kapa ho se be teng ha li-oospores sebakeng se bohareng ba sebaka sa ho kopana sa mefuta. Ho na le likhetho tse 4 tse ka khonehang: mofuta oa mofuta oa A1 haeba o theha oospores ka tester A2, ho A2 haeba e theha oospores ka tester A1, ho A1A2 haeba e etsa li-oospores ka litlhahlobo ka bobeli, kapa e nyopa (00) haeba ha e etse li-oospores eseng ka tester leha e le efe (lihlopha tse peli tsa ho qetela li fumaneha seoelo).
Ho tseba ka potlako mefuta ea ho nyalana, ho ile ha etsoa liteko tsa ho tseba libaka tsa genomic tse amanang le mofuta oa ho nyalana, ka sepheo sa ts'ebeliso ea tsona e tsoelang pele bakeng sa ho khetholla mofuta oa ho nyalana ka PCR. Bafuputsi ba Amerika e ne e le ba bang ba pele ba ho etsa teko e atlehileng ea ho tseba sebaka se joalo (Judelson et al., 1995). Ba sebelisa mokhoa oa RAPD, ba ile ba khona ho khetholla sebaka sa W16 se amanang le mofuta oa ho nyalana ho bana ba li-isolate tse peli tse arohaneng 'me ba rala para ea 24-nt primers ho e holisa (W16-1 (5'-AACACGCAAGGCATATAAATGTA-3') le W16-2 (5' -GCGTAATGTAGCGTAACAGCTTC-3') Ka mor'a ho thibeloa ha sehlahisoa sa PCR ho sebelisa enzyme ea thibelo ea HaeIII, ho ne ho khoneha ho arola li-isolate tse nang le mefuta ea ho nyalana ea A1 le A2.
Teko e 'ngoe ea ho fumana matšoao a PCR ho fumana mefuta ea ho nyalana e entsoe ke bafuputsi ba Korea (Kim le Lee, 2002). Ba ile ba tseba lihlahisoa tse itseng ba sebelisa mokhoa oa AFLP. Ka lebaka leo, ho ile ha ntlafatsoa para ea li-primers PHYB-1 (ho ea pele) (5'-GATCGGATTAGTCAGACGAG-3') le PHYB-2 (5'-GCGTCTGCAAGGCGCATTTT-3'), e leng ho lumellang hore ho khethoe katoloso ea sebaka sa genome se amanang le A2. mofuta oa ho nyalana. Ka mor'a moo, ba ile ba tsoela pele ka mosebetsi ona 'me ba etsa li-primers 5' AAGCTATACTGGGACAGGGT-3' (INF-1, pele) le 5'-GCGTTCTTTCGTATTACCAC-3' (INF-2), e leng ho lumellang ho holisoa ka mokhoa o ikhethileng oa sebaka sa Mat-A1, se khethollang mefuta e nang le mathata. ho kopanya mofuta oa A1. Tšebeliso ea tlhahlobo ea PCR ea mefuta ea ho nyalana e bontšitse liphello tse ntle thutong ea baahi ba P. infestans Czech Republic (Mazakova et al., 2006), Tunisia (Jmour, Hamada, 2006) le libaka tse ling. Ka laboratoring ea rona (Mytsa, Elansky, e sa hatisoang), mefuta e 34 ea P. infestans e arohaneng le litho tse amehileng tsa litapole le tamati libakeng tse fapaneng tsa Russia (Kostroma, Ryazan, Astrakhan, Moscow libaka) li ile tsa hlahlojoa. Liphetho tsa tlhahlobo ea PCR ho sebelisoa li-primers tse khethehileng li tsamaellana le ho feta 90% le liphetho tsa tlhahlobo ea mofuta oa ho nyalana ho sebelisoa mokhoa oa setso oa motsoako oa limatlafatsi.
Letlapa la 1. Phapang ea ho hanyetsa ka hare ho Sib 1 clone (Elansky et al., 2001)
Sebaka sa pokello ea mehlala | Palo ea li-isolates tse hlahlobiloeng | Palo ea mefuta e sa utloahaleng (S), e hanyetsanang ka matla (SR) le mefuta e hanyetsanang (R), likhomphutha (%). | ||
S | SR | R | ||
Vladivostok | 10 | 1 (10) | 4 (40) | 5 (50) |
G. Chita | 5 | 0 | 0 | 5 (100) |
G. Irkutsk | 9 | 9 (100) | 0 | 0 |
Krasnoyarsk | 13 | 12 (92) | 1 (8) | 0 |
Ekaterinburg toropo | 15 | 8 (53) | 1 (7) | 6 (40) |
O. Sakhalin | 66 | 0 | 0 | 66 (100) |
Sebaka sa Omsk | 18 | 0 | 0 | 18 (100) |
Metalaxyl resistance e le lesupa la baahi
Mathoasong a lilemo tsa bo-1980, ho ile ha tlalehoa ho qhoma ho matla ha blight ea morao-rao e bakoang ke mefuta e hanyetsanang le metalaxyl ea P. infestans libakeng tse fapaneng. Temo ea litapole linaheng tse ngata e bile le tahlehelo e kholo (Dowley & O'Sullivan, 1981; Davidse et al., 1983; Derevyagina, 1991). Ho tloha ka nako eo, linaha tse ngata lefatšeng ka bophara li 'nile tsa beha leihlo ho hlaha ha mefuta e hanyetsanang le phenylamide ho batho ba P. infestans. Ntle le tlhahlobo e sebetsang ea menyetla ea ho sebelisa lithethefatsi tse nang le phenylamide, ho haha tsamaiso ea mehato ea tšireletso le ho bolela esale pele li-epiphytoties, ho hanyetsa lithethefatsi tsena ho fetohile e 'ngoe ea litšobotsi tse sebelisoang haholo bakeng sa tlhahlobo ea papiso ea baahi ba pathogen ena. Leha ho le joalo, ts'ebeliso ea ho hanyetsa metalaxyl lithutong tsa palo ea baahi e lokela ho etsoa ho ela hloko taba ea hore: 1 - motheo oa liphatsa tsa lefutso oa ho hanyetsa ha o e-so tsejoe hantle, 2 - ho hanyetsa metalaxyl ke mokhoa o itšetlehileng ka mokhoa o ikhethileng, o khonang ho ho fetola ho itšetlehile ka tšebeliso ea phenylamides, 3 - fapaneng tekanyo ea kutloisiso ho metalaxyl ea mathata ka hare ho mola o le mong oa clonal (Letlapa la 1).
Sebopeho sa isoenzyme
Matšoao a isoenzyme hangata a ikemela ho maemo a kantle, a bonts'a lefa la Mendelian 'me a kopane, a lumella motho ho khetholla pakeng tsa homo- le heterozygote. Tšebeliso ea liprotheine joalo ka matšoao a liphatsa tsa lefutso e etsa hore ho khonehe ho tsebahatsa tlhophiso e kholo ea liphatsa tsa lefutso, ho kenyeletsoa liphetoho tsa chromosomal le genomic, le li-substitutes tse le 'ngoe tsa amino acid.
Lithuto tsa electrophoretic tsa liprotheine li bontšitse hore li-enzyme tse ngata li teng linthong tse phelang ka mokhoa oa likaroloana tse 'maloa tse fapaneng ka ho tsamaea ha electrophoretic. Likaroloana tsena ke phello ea ho kenyelletsa mefuta e mengata ea li-enzyme ka li-loci tse fapaneng (li-isozyme kapa li-isozyme) kapa li-allele tse fapaneng tsa locus e tšoanang (allozymes kapa allozymes). Ke hore, li-isozyme ke mefuta e fapaneng ea enzyme e tšoanang. Mefuta e fapaneng e na le ts'ebetso e ts'oanang ea catalytic, empa e fapana hanyane ka li-substitutes tse le 'ngoe tsa amino acid sebopehong sa peptide le taolong. Phapang e joalo e senoloa ke electrophoresis.
Ha u ithuta ka mefuta ea P. infestans, ho sebelisoa spectra ea isoenzymes ea liprotheine tse peli: peptidase le glucose-6-phosphate isomerase (enzyme ena ke monomorphic ho baahi ba Russia, kahoo mekhoa ea ho ithuta ha e fanoe mosebetsing ona). Ho li arola ka li-isozyme sebakeng sa motlakase, litokisetso tsa protheine tse arohaneng le lintho tse phelang tse ntseng li ithutoa li sebelisoa ho poleiti ea gel e behiloeng tšimong ea motlakase. Sekhahla sa ho ata ha liprotheine ka gel e itšetlehile ka tefiso le boima ba limolek'hule, ka hona, tšimong ea motlakase, motsoako oa liprotheine o arotsoe ka likaroloana tse arohaneng, tse ka bonoang ka li-dae tse khethehileng.
Boithuto ba li-isoenzymes tsa peptidase bo etsoa ka cellulose acetate, starch kapa polyacrylamide gels. Mokhoa o bonolo ka ho fetisisa o ipapisitse le ts'ebeliso ea li-gel cellulose acetate tse hlahisoang ke Helena Laboratories Inc. Ha e hloke lisebelisoa tse ngata tsa tlhahlobo, e lumella motho ho fumana lihlopha tse fapaneng ho gel ka mor'a electrophoresis bakeng sa loci ea enzyme ka bobeli, 'me ha e hloke nako le chelete e ngata (setšoantšo sa 2).
Karolo e nyane ea mycelium e fetisetsoa ho microtube ea 1,5 ml, 'me ho eketsoa marotholi a 1-2 a metsi a silafetseng. Ka mor'a sena, sampole e homogenized (mohlala, ka drill ea motlakase e nang le sephutheloana sa polasetiki se loketseng microtube) le sedimented bakeng sa metsotsoana e 25 ka centrifuge ka 13000 rpm. Ho tsoa ho microtube e 'ngoe le e' ngoe, 8 μl. matla a phahametseng a fetisetsoa poleiting ea kopo.
Selulose acetate gel e ntšoa ka har'a sets'oants'o sa "buffer", e hlakotsoe lipakeng tsa pampiri e 'meli ea sefa ebe e beoa lera le sebetsang le shebile holimo motheong oa polasetiki oa sesebelisoa. Tharollo e tsoang ho poleiti e fetisetsoa ke motho ea sebelisang gel ka makhetlo a 2-4. Gel e fetisetsoa ka kamoreng ea electrophoresis,
Letlapa la 2. Sebopeho sa tharollo e sebelisetsoang ho mebala cellulose acetate gel ha ho hlahlojoa li-isoenzymes tsa peptidase: lerotholi la pente (bromophenol blue) le behoa moeling oa gel.
TRIS HCl, 0,05M, Ph 8,0 2 ml
Peroxidase, 1000 U/ml marotholi a 5
o-dianisidine, 4 mg/ml marotholi a 8
MgCl2, 20 mg/ml 2 marotholi
Gly-Leu, 15 mg / ml marotholi a 10
L-amino-acid oxidase, 20 u / ml 2 marotholi
Electrophoresis e etsoa ka metsotso e 20. ka 200 V. Ka mor'a electrophoresis, gel e fetisetsoa tafoleng ea ho penta ebe e penta ka tharollo e khethehileng ea ho penta (Letlapa la 2). 10 ml ea 1,6% DIFCO agar e qhibilihisoa pele ka ontong ea microwave, e pholile ho 60 ° C, ka mor'a moo 2 ml ea agar e kopantsoe le motsoako oa pente ebe e tšeloa holim'a gel. Methapo e hlaha ka hare ho metsotso e 15-20. L-amino-acid oxidase reagent e eketsoa hang-hang pele o kopanya tharollo le agar e qhibilihisitsoeng.
Ho baahi ba Russia, sebaka sa Pep 1 se emeloa ke genotypes 100/100 le 92/100. Homozygote 92/92 ha e fumanehe haholo (hoo e ka bang 0,1%). Locus Pep 2 e emeloa ke li-genotype tse tharo 100/100, 100/112 le 112/112, 'me mefuta eohle ea 3 e tloaelehile haholo (Elanky, Smirnov, 2003, Fig. 2).
Patlisiso ea Genome
Restriction fragment length polymorphism e lateloang ke hybridization (RFLP-RG 57)
Kakaretso ea DNA e phekoloa ka enzyme ea thibelo ea Eco R1, likaroloana tsa DNA li aroloa ho sebelisoa agarose gel electrophoresis. Nuclear DNA e kholo haholo 'me e na le tatellano e mengata e iphetang, e leng ho etsang hore ho be thata ho hlahloba ka ho toba likaroloana tse ngata tse fumanoeng ka ho thibela li-enzyme. Ka hona, likaroloana tsa DNA tse arohaneng ka gel li fetisetsoa ho lera le khethehileng 'me le sebelisetsoa ho kopanya ka RG 57 probe, e nang le li-nucleotide tse ngotsoeng ka li-labels tsa radioactive kapa fluorescent. Ts'ebetso ena e nyalisitsoe ho latela tatellano e iphetang ka bongata ho genome (Goodwin et al., 1992, Forbes et al., 1998). Kamora ho bona liphetho tsa hybridization ka lisebelisoa tse bobebe kapa tse nang le mahlaseli a kotsi, ho fumanoa profil ea multilocus hybridization (fingerprinting) e emeloang ke likhechana tsa 25-29 (Forbes et al., 1998). Bana ba Asexual (clonal) ba tla ba le lintlha tse tšoanang. Ka sebaka sa lihlopha ho electropherogram, ho tšoana le ho se tšoane ha lintho tse phelang ho bapisoa ho ahloloa.
DNA ea Mitochondrial haplotypes
Liseleng tse ngata tsa eukaryotic, mtDNA e hlahisoa ka sebopeho sa molek'hule oa DNA e selikalikoe habeli, eo, ho fapana le li-chromosome tsa nyutlelie tsa lisele tsa eukaryotic, e iphetang ka mokhoa o ikhethileng 'me ha e amane le limolek'hule tsa protheine.
Jenome ea mitochondrial ea P. infestans e hlophisitsoe, 'me mesebetsi e mengata e nehetsoe tlhahlobo ea bolelele ba likhechana tsa thibelo (Carter et al, 1990, Goodwin, 1991, Gavino and Fry, 2002). Ka mor'a hore Griffith le Shaw (1998) ba hlahise mokhoa o bonolo le o potlakileng oa ho khetholla li-haplotype tsa mtDNA, letšoao lena le ile la fetoha e 'ngoe ea tse tummeng ka ho fetisisa liphuputsong tsa P. infestans Motheo oa mokhoa ona ke ho hōlisa ka tatellano ea likaroloana tse peli tsa DNA ea mitochondrial. genome e tloaelehileng) e nang le li-primers F2-R2 le F4-R4 (Letlapa la 3) le thibelo ea tsona e latelang ho sebelisa li-enzyme tsa thibelo MspI (1st fragment) le EcoR1 (sekhechana sa bobeli). Mokhoa ona o re lumella ho khetholla li-haplotype tse 2: Ia, IIa, Ib, IIb. Mofuta oa II o fapane le mofuta oa I ka ho ba teng ha 4 bp ho kenngoa le sebaka se fapaneng sa libaka tsa thibelo libakeng tsa P1881 le P2 (setšoantšo sa 4).
Ho tloha ka 1996, har'a mefuta e bokelitsoeng Russia, ke li-haplotypes tsa Ia le IIa feela tse ileng tsa hlokomeloa (Elansky et al., 2001, 2015). Li ka tsejoa ka mor'a ho arohana ha lihlahisoa tsa thibelo ka primer F2-R2 tšimong ea motlakase (setšoantšo sa 4, 5). Mefuta ea mtDNA e sebelisoa ha ho bapisoa le mefuta le palo ea batho. Liphuputsong tse 'maloa, mefuta ea DNA ea mitochondrial e ne e sebelisetsoa ho khetholla melapo ea clonal le ho netefatsa hore P. infestans isolates (Botez et al., 2007; Shein et al., 2009). Ho sebelisa mokhoa oa PCR-RFLP, ho ile ha etsoa qeto ea hore mtDNA e fapane ka mokhoa o ts'oanang oa P. infestans (Elansky, Milyutina, 2007). Maemo a ho matlafatsa: 1x (500 sec. 94 ° C), 40x (30 sec. 90 ° C, 30 sec. 52 ° C, 90 sec. 72 ° C); 1x (5 mets. 72°C). Motsoako oa ho itšoara: (20 µl): 0,2U Taq DNA polymerase, 1x 2,5 mM MgCl2-Taq buffer, 0,2 mM e 'ngoe le e' ngoe ea dNTP, 30 pM primer le 5 ng ea DNA ea teko, metsi a deionized - ho fihla ho 20 µl.
Thibelo ea sehlahisoa sa PCR e etsoa lihora tse 4-6 ka mocheso oa 37 ° C. Thibelo motsoako (20 µl): 10x MspI (2 µl), 10x lithibelo buffer (2 µl), metsi deionized (6 µl), PCR sehlahisoa (10 µl).
Letlapa la 3. Li-primers tse sebelisetsoang ho holisa libaka tsa mtDNA polymorphic
Locus | Primer | Bolelele ba pele le ho beoa | PCR bolelele ba sehlahisoa | Thibelo enzyme |
---|---|---|---|---|
P2 | F2: 5'- TTCCCTTTGTCCTCTCTACCGAT | 21; 13619-13639 | 1070 | MspI |
R2: 5'- TTACGGCGGTTTAGCACATACA | 22; 14688-14667 | |||
P4 | F4: 5'- TGGTCATCCAGAGGTTTATGTT | 22; 9329-9350 | 964 | EcoRI |
R4: 5 - CCGATACCGATACCAGCACCAA | 22; 10292-10271 |
Matlafatso ea pele e sa reroang (RAPD)
Ha ho etsoa RAPD, ho sebelisoa primer e le 'ngoe (ka linako tse ling li-primers tse' maloa ka nako e le 'ngoe) ka tatellano e sa reroang ea nucleotide, hangata li-nucleotide tse 10 ka bolelele, tse nang le litaba tse phahameng (ho tloha ho 50%) tsa GC nucleotide le mocheso o tlase o fokolang (hoo e ka bang 35ºC). Li-primers tse joalo li "lula" libakeng tse ngata tse tlatselletsang ho genome. Ka mor'a ho matlafatsa, ho fumanoa palo e kholo ea li-amplicon. Palo ea tsona e ipapisitse le li-primer(s) tse sebelisitsoeng le maemo a karabelo (concentration ea MgCl2 le mocheso oa annealing).
Li-amplicon li bonoa ka ho matha holim'a polyacrylamide kapa gel ea agarose. Ha ho etsoa tlhahlobo ea RAPD, hoa hlokahala ho shebella ka hloko bohloeki ba lintho tse hlahlobiloeng, hobane ho silafatsoa ke lintho tse ling tse phelang ho ka baka keketseho e kholo ea palo ea lintho tsa khale, tse ngata haholo le ha ho hlahlojoa lisebelisoa tse hloekileng (Perez et al, 1998). Tšebeliso ea mokhoa ona oa ho ithuta genome ea P. infestans e bonahala mesebetsing e mengata (Judelson le Roberts, 1999, Ghimire et al., 2002, Carlisle et al., 2001). Khetho ea maemo a karabelo le li-primers (li-primer tsa 51 10-nucleotide li ile tsa ithutoa) li fanoe sehloohong sa Abu-El Samen et al., (2003).
Tlhahlobo ea Microsatellite repeat (SSR).
Ho pheta-pheta ha microsatellite (ho pheta-pheta ha tatellano e bonolo, li-SSR) ke tandemly e phetoa ka tatellano e khutšoane ea 1-3 (ka linako tse ling ho fihla ho 6) li-nucleotide tse teng liphatseng tsa nyutlelie tsa eukaryote kaofela. Palo ea ho pheta-pheta ka tatellano e ka fapana ho tloha ho 10 ho isa ho 100. Microsatellite loci e etsahala ka maqhubu a phahameng a lekaneng 'me e ajoa ka ho lekana ho pholletsa le genome (Lagercrantz et al., 1993). Polymorphism ea tatellano ea microsatellite e amahanngoa le phapang ea palo ea pheta-pheto ea motif ea motheo. Lits'oants'o tsa Microsatellite li na le li-codominant, tse li lumellang hore li sebelisoe ho sekaseka sebopeho sa baahi, ho tseba kamano, litsela tsa ho falla tsa genotype, jj. Har'a melemo e meng ea li-marker tsena, motho o lokela ho ela hloko polymorphism ea bona e phahameng, ho ikatisa hantle, ho se nke lehlakore le monyetla oa ho itlhahloba le ho hlahloba ka mokhoa o itekanetseng oa polymorphism ea microsatellite e etsoa ke ho matlafatsa PCR ho sebelisa li-primers tse tlatsanang le tatellano e ikhethang ea microsatellite. Qalong, tlhahlobo e ne e etsoa ka ho arola lihlahisoa tsa karabelo ho gel ea polyacrylamide. Hamorao, basebetsi ba Applied Biosystems ba ile ba etsa tlhahiso ea tšebeliso ea li-primers tse ngotsoeng ka fluorescent ka ho lemoha lihlahisoa tsa karabelo ho sebelisa mochine o ikemetseng oa laser (Diehl et al., 1990), ebe li-sequencer tse tloaelehileng tsa DNA (Ziegle et al., 1992). Ho ngola li-primer tse nang le lidae tse fapaneng tsa fluorescent ho etsa hore ho khonehe ho sekaseka matšoao a 'maloa tseleng e le' ngoe ka nako e le 'ngoe, ka hona, ho eketsa tlhahiso ea mokhoa haholo le ho ntlafatsa ho nepahala ha tlhahlobo.
Lingoliloeng tsa pele tse nehetsoeng ts'ebelisong ea tlhahlobo ea SSR bakeng sa thuto ea P. infestans li hlahile mathoasong a lilemo tsa bo-2000. (Knapova, Gisi, 2002). Hase matšoao 'ohle a hlahisitsoeng ke bangoli a bontšitseng tekanyo e lekaneng ea polymorphism, leha ho le joalo, tse peli tsa tsona (4B le G11) li kenyelelitsoe sehlopheng sa matšoao a 12 SSR a hlahisitsoeng ke Lees et al (2006) 'me hamorao a amoheloa ho Eucablight marangrang a lipatlisiso (www.eucablight .org) joalo ka maemo a P. infestans. Lilemo tse 'maloa hamorao, thuto e ile ea hatisoa mabapi le ho thehoa ha multiplex DNA analysis system bakeng sa P. infestans e thehiloeng ho matšoao a robeli a SSR (Li et al., 2010). Qetellong, ka mor'a ho hlahloba matšoao 'ohle a neng a reriloe pele le ho khetha tse rutang ka ho fetisisa, hammoho le ho ntlafatsa li-primers, li-tag tsa fluorescent le maemo a amplification, sehlopha se le seng sa bangoli se ile sa hlahisa mokhoa o le mong oa tlhahlobo ea multiplex ho kenyeletsa le matšoao a 12 (Letlapa la 4; Li et al. ., 2013a). Li-primers tse sebelisoang tsamaisong ena li ile tsa khethoa 'me tsa ngoloa ka e' ngoe ea matšoao a fluorescent a mane (FAM, VIC, NED, PET) e le hore mefuta ea boholo ba allele ea li-primers tse nang le mangolo a tšoanang a se ke a kopana.
Bangoli ba ile ba etsa tlhahlobo ho PTC200 thermal cycler (MJ Research, USA) ba sebelisa QIAGEN multiplex PCR kapa QIAGEN Typeit Microsatellite PCR kits. Bophahamo ba motsoako oa karabelo e ne e le 12.5 μl. Maemo a ho holisa a ne a le tjena: bakeng sa QIAGEN multiplex PCR: 95ºC (15 min), 30x (95ºC (20 s), 58ºC (90 s), 72ºC (60 s), 72ºC (20 min); bakeng sa QIAGEN Type-it Microsatellite PCR : 95ºС (5 mets), 28х (95ºС (30 sec), 58ºС (90 sec), 72ºС (20 sec), 60ºС (30 mets).
Karohano le pono ea lihlahisoa tsa PCR li entsoe ho sebelisoa ABI3730 automated capillary DNA analyzer (Applied Biosystems).
Letlapa la 4. Litšobotsi tsa matšoao a 12 a tloaelehileng a SSR a sebelisetsoang genotyping P. infestans (Li et al., 2013a)
Sehlooho | Nomoro ea alleles | Boholo ba boholo alleles (bp) | Lintlha tse qalang |
PiG11 | 13 | 130-180 | F: NED-TGCTATTTATCAAGCGTGGG R: GTTTTCAATCTGCAGCCGTAAGA |
MOTSOALLE | 4 | 255-275 | F: NED-ACTTGCAGAACTACCGCCC R: GTTTGACCACTTTCCTCGGTTC |
PinfSSR11 | 4 | 325-360 | F: NED-TTAAGCCACGACATGAGCTG R: GTTTAGACAATTGTTTTGTGGTCGC |
D13 | 16 | 100-185 | F: FAM-TGCCCCCTGCTCACTC R: GCTCGAATTCATTTTACAGACTTG |
PinfSSR8 | 4 | 250-275 | F: FAM-AATCTGATCGCAACTGAGGG R: GTTTCAAGATACACACGTCGCTCC |
PinfSSR4 | 7 | 280-305 | F: FAM-TCTTGTTCGAGTATGCGACG R: GTTTCACTTCGGGAGAAAGGCTTC |
MOTSOALLE | 4 | 160-175 | F: VIC-AGCGGCTTACCGATGG R: GTTTCAGGCGGCTGTTTTTCGAC |
MOTSOALLE | 3 | 185-205 | F: VIC-ATGAAAATACGTCAATGCTCG R: CGTTGGATATTTCTATTTCTTCG |
PinfSSR6 | 3 | 230-250 | F: GTTTTGGTGGGGCTGAAGTTTT R: VIC-TCGCCACAAGATTTATTCCG |
MOTSOALLE | 3 | 265-280 | F: VIC-ATGACGAAGATGAAAGTGAGG R: CGTATTTCCTGTTTATCTAACACC |
PinfSSR2 | 3 | 165-180 | F: PET-CGACTTCTACATCACCGGC R: GTTTGCTTGGACTGCGTCTTTAGC |
Pi4B | 5 | 200-295 | F: PET-AAAATAAAGCCTTTGGTTCA R: GCAAGCGAGGTTTGTAGATT |
Mohlala oa pono ea liphetho tsa tlhahlobo o bontšoa ho Feiga. 6. Liphetho li ile tsa hlahlobjoa ho sebelisoa lenaneo la GeneMapper 3.7 ka ho bapisa lintlha tse fumanoeng le lintlha tse tsoang ho li-isolates tse tsejoang. Ho nolofatsa tlhaloso ea liphetho tsa liteko, thuto e 'ngoe le e 'ngoe e lokela ho kenyelletsa likarolo tse 1-2 tsa litšupiso tse nang le genotype e tsebahalang.
Mokhoa o reriloeng oa ho etsa lipatlisiso o ile oa lekoa ho palo e kholo ea lisampole tsa masimong, ka mor'a moo bangoli ba tiisa liprothokholo lipakeng tsa lilaboratori tsa mekhatlo e 'meli, The James Hutton Institute (UK) le Wageningen University & Research (Netherlands), eo, hammoho le monyetla oa ho e sebelisa. likarete tse tloaelehileng tsa FTA bakeng sa pokello e nolofalitsoeng le thomello ea lisampole tsa DNA ho tsoa ho P. infestans li ile tsa re lumella ho bua ka monyetla oa tšebeliso ea khoebo ea nts'etsopele ena. Ho phaella moo, mokhoa o potlakileng le o nepahetseng oa genotyping P. infestans o itšehla thajana o sebelisa multiplex SSR analysis e entse hore ho khonehe ho etsa lithuto tse tloaelehileng tsa palo ea batho ba kokoana-hloko ena lefatšeng ka bophara, le ho thehoa ha database ea lefats'e ea morao-rao ka har'a moralo oa Morero oa Eucablight (www.eucablight.org), ho kenyelletsa, ho kenyelletsa le liphetho tsa tlhahlobo ea microsatellite, e entse hore ho khonehe ho latela ho hlaha le ho ata ha liphatsa tsa lefutso tse ncha lefatšeng ka bophara.
Amplified restriction fragment length polymorphism (AFLP). AFLP (amplified fragment length polymorphism) ke theknoloji ea ho fumana li-marker tsa molek'hule tse sa sebetseng ka ho sebelisa li-primers tse itseng. Ho AFLP, DNA e phekoloa ka motsoako oa li-enzyme tse peli tsa thibelo. Li-adapter tse khethehileng li hokeletsoe lintlheng tse khomarelang tsa likhechana tsa thibelo.
Likaroloana tsena li matlafatsoa ka ho sebelisa li-primers tse tlatselletsang tatellano ea li-adapter le sebaka sa lithibelo, hape li nka motheo o le mong kapa ho feta tse khethiloeng ka mokhoa o sa reroang lipheletsong tsa tsona tsa 3'. Sehlopha sa likhechana tse hlahisoang se ipapisitse le li-enzyme tse thibelang le li-nucleotide tse khethiloeng ka mokhoa o sa reroang lipheletsong tsa 3' tsa li-primers (Vos et al., 1995). AFLP genotyping e sebelisetsoa ho ithuta ka potlako ho fetoha ha liphatsa tsa lefutso tsa mefuta e sa tšoaneng.
Tlhaloso e qaqileng ea mokhoa ona e fanoe ka Mueller, Wolfenbarger, 1999, Savelkoul et al., 1999. Mosebetsi o mongata o entsoe ke bafuputsi ba Machaena ho bapisa qeto ea mekhoa ea AFLP le SSR. Litšobotsi tsa phenotypic le genotypic tsa 48 P. infestans isolates tse bokelitsoeng ho tloha libakeng tse hlano tsa Northern China li ile tsa ithutoa. Pono ea AFLP e senotse liphatsa tsa lefutso tse robeli tse fapaneng tsa DNA, ho fapana le liphatsa tsa lefutso tsa SSR, tseo ho seng mefuta e sa tšoaneng e ileng ea khetholloa (Guo et al., 2008).
Ho holisa ka li-primers homologous to transposable element tatellano
Matšoao a tsoang ho tatellano ea retrotransposon e molemo haholo bakeng sa 'mapa oa liphatsa tsa lefutso, ho ithuta mefuta-futa ea liphatsa tsa lefutso le mekhoa ea ho iphetola ha lintho (Schulman, 2006). Haeba u etsa li-primers tse tlatselletsang tatellano e tsitsitseng ea likarolo tse itseng tsa mehala, o ka holisa libaka tsa genome tse lipakeng tsa tsona. Lithutong tsa sesosa sa sesosa sa lefu la morao-rao, mokhoa oa ho holisa likarolo tsa genome o sebelisa primer e tlatselletsang tatellano ea mantlha ea SINE ea retropazone (Short Interspersed Nuclear Elements) e sebelisitsoe ka katleho (Lavrova, Elansky, 2003). Ho sebelisoa mokhoa ona, ho ile ha fumanoa liphapang esita le ho bana ba sexual ba itšehla thajana. Tabeng ena, ho ile ha etsoa qeto ea hore mokhoa oa inter-SINE PCR o totobetse haholo 'me lebelo la ho tsamaea ha likarolo tsa SINE ho genome ea Phytophthora e phahame.
Malapa a leshome le metso e 'meli a li-retrotransposons tse khutšoanyane (SINEs) a khethiloe ho P. infestans genome; Kabo ea mefuta ea li-retrotransposons tse khutšoanyane e ile ea ithutoa, likarolo (SINEs) tse fumanoang ka genome feela ea P. infestans li ile tsa khetholloa (Lavrova, 12).
Likarolo tsa ts'ebeliso ea mekhoa ea ho ithuta ka papiso ea mefuta lithutong tsa baahi
Ha u rera thuto, ho hlokahala hore u utloisise ka ho hlaka lipakane tseo e li hahamallang le ho sebelisa mekhoa e loketseng. Ka hona, mekhoa e meng e etsa hore ho khonehe ho hlahisa palo e kholo ea litšoaneleho tse ikemetseng, empa ka nako e ts'oanang li na le ho ikatisa ho fokolang 'me li itšetlehile haholo ka li-reagents tse sebelisoang, maemo a ho itšoara, le tšilafalo ea thepa e ithutoang. Ka hona, thutong e 'ngoe le e' ngoe ea sehlopha sa mefuta e mengata ho hlokahala hore ho sebelisoe mekhoa e mengata e tloaelehileng (referense), empa leha ho le joalo liphello tsa liteko tse 'maloa li thata haholo ho li kopanya.
Sehlopha sena sa mekhoa se kenyelletsa RAPD, AFLP, InterSSR, InterSINE PCR. Ka mor'a ho holisoa, palo e kholo ea likaroloana tsa DNA tsa boholo bo fapaneng li fumanoa. Ho eletsoa ho sebelisa mekhoa e joalo ha ho hlokahala ho theha phapang pakeng tsa mefuta e amanang haufi-ufi (batsoali-bana, mefuta-futa e hlaha, joalo-joalo), kapa maemong ao ho hlokahalang tlhahlobo e qaqileng ea sampole e nyane. Ka hona, mokhoa oa AFLP o sebelisoa haholo ho liphatsa tsa lefutso tsa P. infestans (van der Lee et al., 1997) le lithutong tsa intrapopulation (Knapova, Gisi, 2002, Cooke et al, 2003, Flier et al, 2003). Ha ho kgothaletswe ho sebelisa mekhoa e joalo ha u theha li-database tsa mathata, hobane Ho batla ho sa khonehe ho kopanya ho rekota liphetho ha ho etsoa litlhahlobo lilaboratoring tse fapaneng.
Ho sa tsotellehe ponahalo e bonolo le lebelo la ho bolaoa (ho itšehla thajana ha DNA ntle le tlhoekiso e ntle, ho hōlisa, pono ea liphello), sehlopha sena sa mekhoa se hloka ho sebelisa mokhoa o khethehileng oa ho ngola liphello: distillation ka gel ea polyacrylamide e ngotsoeng (radioactive kapa luminescent) li-primers le khanya e latelang ea lisebelisoa tse bobebe kapa tse nang le mahlaseli a kotsi. Mokhoa o tloaelehileng oa ho nahana ka lijeli tsa ethidium bromide agarose hangata ha o lokele mekhoa ena hobane palo e kholo ea likhechana tsa DNA tsa boholo bo fapaneng li ka kopanya.
Mekhoa e meng, ho fapana le hoo, e lumella ho hlahisa palo e nyane ea likarolo ka ho ikatisa haholo. Sehlopha sena se kenyelletsa thuto ea li-haplotypes tsa DNA tsa mitochondrial (li-haplotype tse peli feela tsa Ia le IIa li hlokometsoe Russia), mofuta oa ho nyalana (boholo bo arohaneng bo arotsoe ka mefuta e 2: A1 le A2, SF e ikemelang ha e fumanehe ka seoelo) le spectra ea peptidase isoenzymes ( tse peli tsa loci Pep1 le Pep2, tse nang le li-isozyme tse peli ka 'ngoe) le glucose-6-phosphate isomerase (Russia ha ho na phapang ea tšobotsi ena, le hoja polymorphism ea bohlokoa e tsejoa linaheng tse ling tsa lefats'e). Ho eletsoa ho sebelisa likarolo tsena ha ho hlahlobisisoa likoleke le ho bokella datha ea sebaka le lefats'e. Tabeng ea tlhahlobo ea li-isoenzymes le li-haplotype tsa DNA ea mitochondrial, hoa khoneha ho etsa ntle le mefuta e tloaelehileng ka ho feletseng, ha ho hlahlojoa mefuta ea ho nyalana, ho hlokahala liteko tse peli tse arohaneng le mefuta e tsebahalang ea ho nyalana.
Maemo a ho itšoara le li-reagents li ka ama feela phapang ea sehlahisoa ho electropherogram;
Hona joale, boholo ba baahi ba karolong ea Europe ea Russia ba emeloa ke mefuta ea mefuta e 'meli ea ho nyalana (Letlapa la 6), har'a tsona ho na le lihlopha tse arohaneng le mefuta ea Ia le IIa ea DNA ea mitochondrial (mefuta e meng ea mtDNA e fumanoang lefatšeng ha e e-s'o fumanoe. Russia ka mor'a 1993). The spectra of peptidase isoenzymes e emeloa ke genotypes tse peli sebakeng sa Pep1 (100/100, 92/92 le heterozygote 92/100, le 92/92 genotype e sa tloaelehang haholo (<0,3%)) le li-genotype tse peli ho Pep 2 locus (100/100 , 112/112 le heterozygote 100/112, le genotype 112/112 ha e tloaelehe ho feta 100/100, empa hape e tloaelehile haholo).
Ha ho phapang pakeng tsa li-isomerase tsa glucose-6-phosphate isomerase e ileng ea hlokomeloa ka mor'a 1993 (ho nyamela ha li-isolates tsa US-1 tse ithutoang li ne li e-na le genotype ea 100/100 (Elansky le Smirnov, 2002).
Sehlopha sa boraro sa mekhoa se etsa hore ho khonehe ho fumana sehlopha se lekaneng sa litšoaneleho tsa matšoao a ikemetseng ka ho ikatisa ho hoholo. Kajeno, sehlopha sena se kenyelletsa sampuli ea RFLP-RG57, e hlahisang likaroloana tsa 25-29 tsa DNA tsa boholo bo fapaneng. RFLP-RG57 e ka sebelisoa ka bobeli ha ho hlahlojoa lisampole le ha ho etsoa li-database. Leha ho le joalo, mokhoa ona o theko e boima haholo ho feta o fetileng, o hloka nako e ngata, 'me o hloka palo e lekaneng ea DNA e hloekisitsoeng haholo. Ka hona, mofuputsi o qobelloa ho fokotsa boholo ba lintho tse lekiloeng.
Nts'etsopele ea RFLP-RG57 mathoasong a lilemo tsa bo-90 e matlafalitse haholo lithuto tsa baahi tsa pathogen ea morao-rao ea blight. E ile ea fetoha motheo oa mokhoa o thehiloeng ho itšehla thajana le tlhahlobo ea "Clonal Lines" (sheba ka tlase). Hammoho le RFLP-RG57, mofuta oa mating, DNA fingerprinting (RFLP-RG57 mokhoa), spectra ea peptidase le glucose-6-phosphate isomerase isoenzymes, le mofuta oa DNA oa mitochondrial li sebelisoa ho khetholla mela ea clonal. Ka lebaka la hae, e bonts'itsoe ke al., 1994), ho nkeloa sebaka ha baahi ba khale ke ba bacha (Drenth et al, 1993, Sujkowski et al, 1994, Goodwin et al, 1995a), le li-clonal lines tse atileng linaheng tse ngata tsa lefatshe le ile la tsebahatswa. Lithuto tsa mefuta ea Serussia e sebelisang mokhoa ona li bontšitse polymorphism e phahameng ea genotypic ea mefuta e karolong ea Europe le monomorphism ea baahi ba libakeng tsa Asia le Bochabela bo Hōle ba Russia (Elansky et al, 2001). 'Me joale mokhoa ona e ntse e le oona o ka sehloohong ha ho etsoa lithuto tsa baahi ba P. infestans. Leha ho le joalo, tšebeliso e pharalletseng ea eona e sitisoa ke litšenyehelo tsa eona tse phahameng haholo le ts'ebetsong e hlokang mosebetsi o boima.
Mokhoa o mong o ts'episang o sa sebelisoeng haholo lipatlisisong tsa P. infestans ke tlhahlobo ea microsatellite repeat (SSR). Hajoale, mokhoa ona o sebelisoa haholo ho arola mela ea clonal. Bakeng sa tlhahlobo ea mefuta, litšobotsi tsa matšoao a phenotypic joalo ka boteng ba liphatsa tsa lefutso tsa mefuta ea litapole (Avdey, 1995, Ivanyuk et al., 2002, Ulanova et al., 2003) le tamati li sebelisitsoe haholo (mme li ntse li tsoela pele ho sebelisoa. ). Ho fihla joale, liphatsa tsa lefutso tsa virulence bakeng sa mefuta ea litapole li lahlehetsoe ke boleng ba tsona e le matšoao a thuto ea baahi ka lebaka la ponahalo ea palo e kholo (kapa e haufi le eona) ea liphatsa tsa lefutso tse ngata tse ka thoko. Ka nako e ts'oanang, mofuta oa T1 oa likokoana-hloko bakeng sa mefuta ea tamati e nang le mofuta o tšoanang oa Ph1 o ntse o sebelisoa ka katleho e le letšoao (Lavrova et al., 2003, Ulanova et al., 2003).
Lithuto tse ngata li sebelisa khanyetso ea fungicide e le letšoao la matšoao. Tšobotsi ena ha e khothaletsoe ho sebelisoa liphuputsong tsa baahi ka lebaka la ketsahalo e batlang e le bonolo ea liphetoho tsa ho hanyetsa meleng ea clonal ka mor'a tšebeliso ea metalaxyl- (kapa mefenoxam-) e nang le likokoana-hloko tšimong. Ka mohlala, phapang e kholo ea boemo ba ho hanyetsa e bontšitsoe ka har'a moloko oa Sib1 clonal (Elansky et al., 2001).
Kahoo, litšoaneleho tse khethiloeng tsa matšoao bakeng sa ho theha libanka tsa data le mefuta ea ho ngola likolekeng ke mofuta oa ho nyalana, peptidase isoenzyme spectrum, mofuta oa DNA oa mitochondrial, RFLP-RG57, SSR. Ho bapisa mehlala e fokolang, haeba ho hlokahala ho sebelisa palo e kholo ea litšoaneleho tsa matšoao, u ka sebelisa AFLP, RAPD, InterSSR, Inter-SINE PCR (Letlapa la 5). Leha ho le joalo, ho lokela ho hopoloa hore mekhoa ena ha e khonehe hantle, 'me tekong e' ngoe le e 'ngoe ea motho ka mong (amplification electrophoresis cycle) hoa hlokahala ho sebelisa li-isolate tse' maloa tsa litšupiso.
Letlapa la 5. Papiso ea mekhoa e fapaneng ea ho ithuta mefuta P. Mokhachane
criterion | ТС | Li-isoenzymes | MtDNA | RFLP-RG57 | KOTSI | ISSR | SSR | AFLP | Tšen |
---|---|---|---|---|---|---|---|---|---|
Bongata ba tlhahisoleseding | Н | Н | Н | С | В | В | С | В | В |
Ho ikatisa | В | В | В | В | Н | Н | С | С | С |
Monyetla oa lintho tsa khale | Н | Н | Н | Н | В | С | Н | С | В |
ditjeho tsa ho | Н | С | Н | В | Н | Н | Н | С | Н |
Matla a mosebetsi | Н | Н | Н | В | NS* | NS* | Н | С | NS* |
Lebelo la tlhahlobo** | В | Н | Н | С | Н | Н | Н | Н | Н |
Tlhokomeliso: H - tlaase, C - bohareng, V - holimo; NS* - matla a tlase a mosebetsi ha o sebelisa gel ea agarose kapa othomathike
genotype, medium - ha e tšeloa ka gel ea polyacrylamide e nang le li-primers tse ngotsoeng,
** - ho sa bale nako e sebelisitsoeng ho holisa mycelium ho arola DNA.
Sebopeho sa baahi
Mela ea clonal
Ha ho se na recombination kapa tlatsetso ea eona e sa reng letho ho sebopeho sa baahi, palo ea baahi e na le palo e itseng ea li-clones, phapanyetsano ea liphatsa tsa lefutso lipakeng tsa eona e sa tloaelehang haholo.
Ho batho ba joalo, ho ruta haholo ho ithuta eseng maqhubu a liphatsa tsa lefutso ka bomong, empa maqhubu a genotypes a nang le tšimoloho e le 'ngoe (clonal lineages) mme a fapana feela ka liphetoho tsa ntlha. Liphuputso tsa palo ea baahi tsa kokoana-hloko ea morao-rao le tlhahlobo ea melapo ea clonal li eketsehile haholo ho tloha ha ho fihla mokhoa oa RFLP-RG57 mathoasong a lilemo tsa bo-90 tsa lekholo le fetileng. Hammoho le RFLP-RG57, mofuta oa mating, spectra ea peptidase le glucose-6-phosphate isomerase isoenzymes, le mofuta oa DNA oa mitochondrial li sebelisoa ho khetholla mela ea clonal. Litšobotsi tsa mela e tloaelehileng ea clonal li fanoe ho Lethathamo la 6.
US-1 clone e ne e laola baahi hohle ho fihlela bofelong ba lilemo tsa bo-80, ka mor'a moo e ile ea qala ho nkeloa sebaka ke li-clone tse ling 'me ea nyamela Europe le Amerika Leboea. Hona joale e fumaneha Bochabela bo Hōle (Philippines, Taiwan, China, Japan, Korea, Koh et al., 1994, Mosa et al., 1993), Afrika (Uganda, Kenya, Rwanda, Goodwin et al., 1994, Vega-Sanchez et al., 2000, Ochwo et al., 2002) le Amerika Boroa (Ecuador, Brazil, Peru, Forbes et al., 1997, Goodwin et al., 1994). Ha ho mefuta ea leloko la US-1 e fumanoeng Australia feela. Kamoo ho bonahalang kateng, P. infestans isolates e fihlile Australia ka leqhubu le leng la ho falla (Goodwin, 1997).
US-6 clone e ile ea tloha leboea Mexico ho ea California ho elella bofelong ba lilemo tsa bo-70 'me ea baka seoa sa litapole le tamati moo kamora lilemo tse 32 tsa bosieo ba lefu. Ka lebaka la mabifi a eona a phahameng, e ile ea nkela sebaka sa US-1 sebaka mme ea ba matla lebopong le ka bophirima la United States (Goodwin et al., 1995a).
Li-genotypes US-7 le US-8 li ile tsa sibolloa United States ka 1992 'me li se li atile United States le Canada ka 1994. Ka nako e le 'ngoe ea masimo, clone US-8 e khona ho nka sebaka sa US-1 ka botlalo libakeng tsa litapole tse neng li tšoaelitsoe ke li-clone ka bobeli ka bongata bo lekanang (Miller le Johnson, 2000).
Clones BC-1 ho BC-4 li ile tsa khetholloa British Columbia ka palo e fokolang ea libaka tse ka thōko (Goodwin et al., 1995b). Clone ea US-11 e ile ea ata haholo United States 'me ea nkela US-1 sebaka sa Taiwan. Clones JP-1 le EC-1, hammoho le clone US-1, li ajoa Japane le Ecuador, ka ho latellana (Koh et al., 1994; Forbes et al., 1997).
SIB-1 ke mofuta o neng o atile Russia sebakeng se seholo ho tloha sebakeng sa Moscow ho ea Sakhalin. E ile ea sibolloa sebakeng sa Moscow ka 1993, 'me ho na le baahi ba bangata ba nang le mefuta e mengata ea li-clonal line tse hanyetsanang haholo le metalaxyl. Ka mor'a 1993, ho ata ha clone ena ho ile ha fokotseha haholo. Ka ntle ho Urals ka 1997-1998, SIB-1 e ne e fumanoa hohle ntle le Khabarovsk Territory (clone ea SIB-2 e atile moo). Karohano ea sebaka sa li-clones tse nang le mefuta e fapaneng ea ho nyalana ha e kenyelle ts'ebetso ea thobalano sebakeng sa Siberia le Bochabela bo Hōle. Sebakeng sa Moscow, ho fapana le Siberia, baahi ba emeloa ke li-clones tse ngata; hoo e ka bang karolo e 'ngoe le e' ngoe e arohaneng e na le mofuta o ikhethang oa li-multilocus genotype (Elansky et al., 2001, 2015). Phapang ena e ke ke ea hlalosoa feela ka ho hlahisoa ha mefuta ea fungal e tsoang likarolong tse fapaneng tsa lefats'e ka thepa ea peo e tsoang kantle ho naha. Kaha mefuta e 'meli ea ho nyalana e etsahala har'a baahi, ho ka etsahala hore mefuta-futa ea eona e boetse e bakoa ke ho kopana hape. Kahoo, British Columbia, ho hlaha ha genotypes BC-2, BC-3 le BC-4 ho nahanoa ka lebaka la ho kopanngoa ha li-clones BC-1 le US-6 (Goodwin et al., 1995b). Ho ka etsahala hore mefuta e nyalisitsoeng e boetse e fumanoa ho baahi ba Moscow. Ka mohlala, mefuta ea MO-4, MO-8 le MO-11 heterozygous bakeng sa sebaka sa PEP e ka ba mefuta e nyalisitsoeng pakeng tsa mefuta ea MO-12, MO-21, MO-22, e nang le mofuta oa A2 oa ho nyalana 'me e le homozygous bakeng sa allele e le 'ngoe. PEP locus le strain MO-8, e nang le mofuta oa ho nyalana oa A1 le homozygous bakeng sa allele e 'ngoe ea locus. 'Me haeba ho joalo,' me ka baahi ba mehleng ea kajeno ea P. infestans ho na le tšekamelo ea ho eketsa karolo ea ts'ebetso ea thobalano, joale boleng ba tlhahisoleseding ea tlhahlobo ea li-clone tsa multilocus bo tla fokotseha (Elansky et al., 2001, 2015).
Phapang ka meloko ea clonal
Ho fihlela lilemong tsa bo-90 tsa lekholo la bo20 la lilemo, mohala oa US-1 oa clonal o ne o atile lefatšeng. Boholo ba baahi ba masimong le ba libaka ba ne ba ipapisitse le mefuta e fapaneng ea mofuta oa US-1. Leha ho le joalo, ho ile ha boela ha hlokomeloa liphapang pakeng tsa li-isolates, tse ka 'nang tsa bakoa ke phetoho ea phetoho. Liphetoho li etsahetse ho DNA ea nyutlelie le ea mitochondrial 'me, har'a lintho tse ling, li amme boemo ba ho hanyetsa lithethefatsi tsa phenylamide le palo ea liphatsa tsa lefutso tsa virulence. Methalo e fapaneng le mefuta ea mantlha ea liphatsa tsa lefutso e khethoa ka linomoro tse eketsehileng ka mor'a letheba le latelang lebitso la mofuta oa pele oa genotype (mohlala, US-1.1 mutant line ea US-1 clonal line). Mehato ea menoana ea DNA ea US-1.5 le US-1.6 lineages e na le li-accessory lineages tsa boholo bo sa tšoaneng (Goodwin et al., 1995a, 1995b); leloko la clonal US-6.3 le lona le fapane le US-6 ka leloko le le leng la tlatsetso (Goodwin, 1997, Lethathamo la 7).
Ha ho ntse ho ithuta DNA ea mitochondrial, ho ile ha fumaneha hore mofuta oa 1b feela oa DNA ea mitochondrial o fumanoe moleng oa US-1 oa clonal (Carter et al., 1990). Leha ho le joalo, ha ho hlahlojoa mefuta ea leloko lena la clonal ho tloha Peru le Philippines, ho ile ha hlokomeloa batho ba itšehlang thajana bao mefuta ea bona ea DNA ea mitochondrial e neng e fapane le 1b ka boteng ba ho kenya le ho hlakolwa (Goodwin, 1991, Koh et al., 1994).
Letlapa la 6. Multilocus genotypes ea meloko e meng ea clonal ea P. infestans
Sehlooho | Mofuta oa ho nyalana | Li-isoenzymes | DNA menoana | Mofuta oa MtDNA | |
GPI | PEP | ||||
US-1 | A1 | 86/100 | 92/100 | 1.0111010110011E + 24 | Ib |
US-2 | A1 | 86/100 | 92/100 | 1.0111010010011E + 24 | - |
US-3 | A1 | 86/100 | 92/100 | 1.0111000000011E + 24 | - |
US-4 | A1 | 100/100 | 92/92 | 1.0111010010011E + 24 | - |
US-5 | A1 | 100/100 | 92/100 | 1.0111010010011E + 24 | - |
US-6 | A1 | 100/100 | 92/100 | 1.0111110010011E + 24 | IIb |
US-7 | A2 | 100/111 | 100/100 | 1.0011000010011E + 24 | Ia |
US-8 | A2 | 100/111/122 | 100/100 | 1.0011000010011E + 24 | Ia |
US-9 | A1 | 100/100 | 83/100 | * | - |
US-10 | A2 | 111/122 | 100/100 | - | - |
US-11 | A1 | 100/111 | 92/100 | 1.0101110010011E + 24 | IIb |
US-12 | A1 | 100/111 | 92/100 | 1.0001000010011E + 24 | - |
US-14 | A2 | 100/122 | 100/100 | 1.0000000000011E + 24 | - |
US-15 | A2 | 100/100 | 92/100 | 1.0001000010011E + 24 | Ia |
US-16 | A1 | 100/111 | 100/100 | 1.0001100010011E + 24 | - |
US-17 | A1 | 100/122 | 100/100 | 1.0100010000011E + 24 | - |
US-18 | A2 | 100/100 | 92/100 | 1.0001000010011E + 24 | Ia |
US-19 | A2 | 100/100 | 92/100 | 1.0101010000011E + 24 | Ia |
EC-1 | A1 | 90/100 | 96/100 | 1.1111010010011E + 24 | IIa |
SIB-1 | A1 | 100/100 | 100/100 | 1.0001000110011E + 24 | IIa |
SIB-2 | A2 | 100/100 | 100/100 | 1.0001000010011E + 24 | IIa |
SIB-3 | A1 | 100/100 | 100/100 | 1.1001010100011E + 24 | IIa |
MO-1 | A2 | 100/100 | 100/100 | 1.0001000110011E + 24 | IIa |
MO-2 | A2 | 100/100 | 100/100 | 1.0001000010011E + 24 | Ia |
MO-3 | A1 | 100/100 | 100/100 | 1.0101000010011E + 24 | IIa |
MO-4 | A1 | 100/100 | 92/100 | 1.0101110110011E + 24 | IIa |
MO-5 | A1 | 100/100 | 100/100 | 1.0001010010011E + 24 | IIa |
MO-6 | A1 | 100/100 | 100/100 | 1.0101010010011E + 24 | Ia |
MO-7 | A1 | 100/100 | 92/100 | 1.0001000110011E + 24 | IIa |
MO-8 | A1 | 100/100 | 92/92 | 1.0101100010011E + 24 | IIa |
MO-9 | A1 | 100/100 | 92/100 | 1.0001000010011E + 24 | IIa |
MO-10 | A1 | 100/100 | 100/100 | 1.0101100000011E + 24 | Ia |
MO-11 | A1 | 100/100 | 92/100 | 1.0101010010011E + 24 | Ia |
MO-12 | A2 | 100/100 | 100/100 | 1.0101010010011E + 24 | Ia |
MO-13 | A1 | 100/100 | 100/100 | 1.0101010000011E + 24 | Ia |
MO-14 | A1 | 100/100 | 100/100 | 1.01010010011E + 22 | Ia |
MO-15 | A1 | 100/100 | 100/100 | 1.101110010011E + 23 | Ia |
MO-16 | A1 | 100/100 | 100/100 | 1.0001000000011E + 24 | IIa |
MO-17 | A1 | 86/100 | 100/100 | 1.0101010110011E + 24 | Ib |
MO-18 | A1 | 100/100 | 100/100 | 1.0101110010011E + 24 | IIa |
MO-19 | A1 | 100/100 | 100/100 | 1.0101010000011E + 24 | IIa |
MO-20 | A2 | 100/100 | 100/100 | 1.0101010000011E + 24 | IIa |
MO-21 | A2 | 100/100 | 100/100 | 1.0101010000011E + 24 | IIa |
Tlhokomeliso: * - ha ho na data.
Letlapa la 7. Multilocus genotypes le mela ea bona e feto-fetohang
Sehlooho | Mofuta oa ho nyalana | | DNA Menoana (RG57) | Lintlha | |
GPI | PEP-1 | ||||
US-1 | A1 | 86/100 | 92/100 | 1011101011001101000110011 | Genotype 1 ea mantlha |
US-1.1 | A1 | 86/100 | 100/100 | 1011101011001101000110011 | Phetoho ho PEP |
US-1.2 | A1 | 86/100 | 92/100 | 1011101010001101000110011 | Phetoho ho RG57 |
US-1.3 | A1 | 86/100 | 92/100 | 1011101001001101000110011 | Phetoho ho RG57 |
US-1.4 | A1 | 86/100 | 100/100 | 1011101010001101000110011 | Phetoho ho RG57 le PEP |
US-1.5 | A1 | 86/100 | 92/100 | 1011101011001101010110011 | Phetoho ho RG57 |
US-6 | A1 | 100/100 | 92/100 | 1011111001001100010110011 | Genotype 2 ea mantlha |
US-6.1 | A1 | 100/100 | 92 /92 | 1011111001001100010110011 | Phetoho ho PEP |
US-6.2 | A1 | 100/100 | 92/100 | 1011101001001100010110011 | Phetoho ho RG57 |
US-6.3 | A1 | 100/100 | 92/100 | 1011111001011100010110011 | Phetoho ho RG57 |
US-6.4 | A1 | 100/100 | 100/100 | 1011011001001100010110011 | Phetoho ho RG57 le PEP |
US-6.5 | A1 | 100/100 | 92/100 | 1011111001001100010010011 | Phetoho ho RG57 |
EA-1 | A2 | 100/100 | 100/100 | 1011101000001100001111011 | Genotype 3 ea mantlha |
EA-1.1 | A2 | 100/100 | 100/100 | 1010101000001100001110011 | Phetoho ho RG57 |
Ho boetse ho na le liphetoho ho spectra ea isoenzymes. E le molao, li bakoa ke ho senyeha ha ntho e phelang qalong ea heterozygous bakeng sa enzyme ena hore e be li-homozygous. Ka 1993, re ile ra fumana khatello ea litholoana tsa tamati e nang le litšobotsi tsa US-1: RG57 fingerprinting, mitochondrial DNA type le genotype 86/100 bakeng sa glucose-6-phosphate isomerase, empa e ne e le homozygous (100/100) bakeng sa peptidase ea pele. locus sebakeng sa heterozygotes e tloaelehileng bakeng sa mohala ona oa clonal ke 92/100. Re rehile mofuta oa mofuta oa mofuta ona MO-17 (Letlapa la 6). Mela e fetotsoeng US-1.1 le US-1.4 le eona e fapana le US-1 ka liphetoho sebakeng sa pele sa peptidase (Letlapa la 7).
Liphetoho tse lebisang phetohong ea palo ea liphatsa tsa lefutso tsa mofuta oa tamati le tamati li etsahala hangata. Li tlalehiloe har'a libaka tse ikhethileng tsa leloko la US-1 la clonal ho baahi ba tsoang Netherlands (Drenth et al., 1994), Peru (Goodwin et al., 1995a), Poland (Sujkowski et al., 1991), le leboea Leboea. Amerika (Goodwin et al., 1995 ., 7b). Phapang ea palo ea liphatsa tsa lefutso tsa litapole e boetse e hlokometsoe har'a li-clonal tsa US-8 le US-1995 tsa Canada le USA (Goodwin et al., 1a), har'a batho ba ikhethileng ba leloko la SIB-2001 karolong ea Asia. ea Russia (Elansky et al., XNUMX).
Li-Isolates tse nang le phapang e matla maemong a ho hanyetsa lithethefatsi tsa phenylamide li ile tsa khetholloa ho baahi ba masimo a monoclonal, bao kaofela ha bona e neng e le ba leloko la Sib-1 clonal (Elansky et al, 2001, Letlapa la 1). Hoo e ka bang mefuta eohle ea leloko la US-1 la clonal e kotsing haholo ho metalaxyl, empa likheo tse hanyetsanang haholo tsa leloko lena li arotsoe Philippines (Koh et al., 1994) le Ireland (Goodwin et al., 1996).
Baahi ba kajeno ba P. infestans
Amerika Bohareng (Mexico)
Palo ea baahi ba P. infestans Mexico e fapane haholo le baahi ba bang ba lefats'e, e leng haholo-holo ka lebaka la boemo ba eona ba histori. Liphuputso tse ngata tsa baahi bana le mefuta e amanang le eona ea Phytophthora clade ho P. infestans, hammoho le mefuta ea lehae ea mofuta oa Solanum, e lebisitse ho qeto ea hore ho iphetola ha likokoana-hloko bohareng ba Mexico ho etsahetse hammoho le ho fetoha ha limela tse amohelang batho. 'me e ne e amahanngoa le ho kopanya thobalano (Grünwald, Flier, 2005). Mefuta e 'meli ea ho nyalana e emeloa ho baahi,' me ka tekanyo e lekanang, le boteng ba oospores mobung, limela le li-tubers tsa litapole le mefuta e sa tšoaneng e amanang le naha ea Solanum e tiisa ho ba teng ha ts'ebetso ea thobalano ho baahi (Fernández-Pavía). le tse ling, 2002). Liphuputso tsa morao-rao le haufi le Toluca Valley (sebaka se belaelloang sa tšimoloho ea kokoana-hloko) li tiisitse mefuta e mengata ea liphatsa tsa lefutso tsa P. infestans ea sebakeng seo (134 multilocus genotypes ka mohlala oa 176 accessions) le ho ba teng ha lihlopha tse 'maloa tse arohaneng. sebaka (Wang et al., 2017). Lintho tse tlatselletsang phapanong ena li kenyelletsa karohano ea sebaka sa libaka tsa libaka tse lithaba tsa Mexico bohareng, phapang ea maemo a temo le mefuta ea litapole tse sebelisoang liphuleng le lithabeng, le ho ba teng ha mefuta ea solanum e nang le tuber e hlaha e ka sebetsang e le libaka tse ling. (Fry et al., 2009).
Leha ho le joalo, hoa lokela ho hlokomeloa hore baahi ba P. infestans karolong e ka Leboea ea Mexico ba na le tlhaho ea clonal haholoanyane 'me ba tšoana haholo le baahi ba Amerika Leboea, e leng se ka bontšang mefuta e mecha (Fry et al., 2009).
Amerika Leboea
Baahi ba Amerika Leboea ba P. infestans esale ba e-na le sebopeho se bonolo haholo 'me sebopeho sa bona sa clonal se thehiloe nako e telele pele ho sebelisoa tlhahlobo ea microsatellite. Ho fihlela 1987, leloko la US-1 la clonal le ne le busa United States le Canada (Goodwin et al., 1995). Bohareng ba lilemo tsa bo-70, ha li-fungicides tse thehiloeng ho metalaxyl li ne li fumaneha, clone ena e ile ea qala ho nkeloa sebaka ke mefuta e meng e hanyetsanang haholoanyane e ileng ea falla Mexico (Goodwin et al., 1998). Qetellong ea lilemo tsa bo-90. US-8 genotype e nkile sebaka sa US-1 genotype United States mme ea fetoha leloko le ka sehloohong la litapole (Fry et al., 2009; Fry et al., 2015). Boemo bo ne bo fapane le tamati, moo mela e mengata ea clonal e neng e lula e le teng, 'me sebopeho sa bona se fetoha selemo le selemo (Fry et al., 2009).
Ka 2009, ho ile ha qhoma seoa se seholo sa lefu la morao-rao la tamati United States. Ntho e ikhethang ea seoa sena e ne e le ho qala ha eona hoo e batlang e le ka nako e le 'ngoe libakeng tse ngata karolong e ka leboea-bochabela ea United States,' me e ne e amahanngoa le thekiso e kholo ea lipeo tsa tamati e nang le tšoaetso litsing tse kholo tsa jarete (Fry et al., 2013). Tahlehelo ea lijalo e ne e le khōlō haholo. Tlhahlobo ea Microsatellite ea lisampole tse amehileng e senotse hore mofuta oa seoa sena ke oa leloko la US-22 la clonal ea mofuta oa A2. Ka 2009, karolo ea genotype ena ho baahi ba Amerika ba P. infestans e fihlile ho 80% (Fry et al., 2013). Lilemong tse latelang, palo ea li-genotypes tse mabifi tsa US-23 (haholo-holo ho tamati) le US-24 (litapole) palo ea baahi e ile ea eketseha butle-butle, leha ho le joalo, ka mor'a 2011, palo ea ho fumanoa ha US-24 e fokotsehile haholo, 'me ho fihlela joale. , hoo e ka bang 90% ea baahi ba pathogen ba USA e emeloa ke genotype ea US-23 (Fry et al., 2015).
Canada, joalo ka USA, ho elella bofelong ba lilemo tsa bo-90. mofuta o ka sehloohong oa genotype US-1 e ile ea nkeloa sebaka ke US-8, eo boemo ba eona bo ka sehloohong bo ileng ba lula bo sa sisinyehe ho fihlela 2008. Ka 2009-2010. Canada, ho bile le mafu a seoa a tebileng a lefu la morao-rao le amanang le thekiso ea lipeo tsa tamati e tšoaelitsoeng, empa li bakiloe ke genotypes US-23 le US-8 (Kalischuk et al., 2012). Ho hlokomelehang ke tlhaloso e hlakileng ea libaka tsa mefuta ena ea liphatsa tsa lefutso: US-23 e ne e le matla liprofinseng tse ka bophirimela tsa Canada (68%), ha US-8 e ne e le matla liprofinseng tse ka bochabela (83%). Lilemong tse latelang, US-23 e ile ea hasana libakeng tse ka bochabela, empa ka kakaretso karolo ea eona ho baahi e ile ea fokotseha hanyenyane ka lebaka la ponahalo ea genotypes US-22 le US-24 naheng (Peters et al., 2014). Ho fihlela joale, US-23 e boloka boemo bo ka sehloohong ho pholletsa le Canada; US-8 e teng British Columbia, 'me US-23 le US-24 li teng Ontario (Peters, 2017).
Kahoo, baahi ba Amerika Leboea ba P. infestans ba emela haholo-holo meloko ea clonal. Lilemong tse 40 tse fetileng, palo ea li-clonal genotypes tse sibollotsoeng e fihlile ho 24. Ho sa tsotellehe taba ea hore mefuta ea mefuta e 'meli ea ho nyalana e teng har'a baahi, monyetla oa ho hlaha ha liphatsa tsa lefutso tse ncha ka lebaka la ho kopanela liphate hape o ntse o le tlaase haholo. Leha ho le joalo, lilemong tse 20 tse fetileng, ho tlalehiloe linyeoe tse 'maloa tsa ho hlaha ha ephemeral recombinant (Gavino et al., 2000; Danies et al., 2014; Peters et al., 2014), 'me ketsahalong e' ngoe sephetho sa ho tšela e ne e le mofuta oa US-11, o thehileng sebaka sa Amerika Leboea ka lilemo tse ngata (Gavino et al., 2000). Ho fihlela ka 2009, liphetoho sebopehong sa baahi li ne li amahanngoa le ho hlaha ha liphatsa tsa lefutso tse ncha, tse mabifi le ho falla ha batho ba pele ba neng ba busa pele. E etsahetse ka 2009-2010 USA le Canada, li-epiphytoties li bontšitse ka lekhetlo la pele hore nakong ea ho ikopanya ha lichaba tsa lefatše, ho qhoma ha mafu ho ka boela ha amahanngoa le ho ata ka mafolofolo ha li-genotypes tse ncha nakong ea thekiso ea lisebelisoa tsa ho jala tse nang le tšoaetso.
Amerika Boroa
Ho fihlela morao tjena, liphuputso tsa baahi ba Amerika Boroa ba P. infestans e ne e se kamehla kapa ka tekanyo e kholo. Hoa tsebahala hore sebopeho sa baahi bana se bonolo haholo 'me se kenyelletsa mela e 1-5 naheng ka 'ngoe (Forbes et al., 1998). Kahoo, ka 1998, li-genotypes US-1 (Brazil, Chile) BR-1 (Brazil, Bolivia, Uruguay, Paraguay), EC-1 (Ecuador, Colombia, Peru le Venezuela), AR-1, AR li ile tsa sibolloa ka litapole - 2, AR-3, AR-4 le AR-5 (Argentina), PE-3 le PE-7 (Peru e ka boroa). Mofuta oa ho nyalana oa A2 o ne o le teng Brazil, Bolivia le Argentina 'me ha oa ka oa fumanoa ka nģ'ane ho moeli oa Bolivia-Peruvia sebakeng sa Lake Titicaca, moo mofuta oa A1-mofuta oa EC-1 o neng o atile Andes. Ho tamati, US-1 e ile ea lula e le mofuta o ka sehloohong oa genotype ho pholletsa le Amerika Boroa.
Boemo bona bo ile ba tsoela pele ka bo-2000. Ntlha ea bohlokoa e ne e le ho sibolloa ha mohala o mocha oa mofuta oa A2 oa clonal EC-2 ho beng ka litapole tse hlaha (S. brevifolium le S. tetrapetalum) karolong e ka Leboea ea Andes (Oliva et al., 2010). Liphuputso tsa Phylogenetic li bontšitse hore leloko lena ha le tšoane ka ho feletseng le P. infestans, le hoja e amana haufi-ufi le eona, ka hona ho ile ha etsoa tlhahiso ea ho e nahana, hammoho le leloko le leng, EC-3, le arohaneng le S. betaceum tamati sefate se melang Andes mofuta o mocha o bitsoang P. andina; leha ho le joalo, boemo ba mofuta ona (mofuta o ikemetseng kapa lebasetere la P. infestans le meloko e seng e sa tsejoe) e ntse e sa hlaka (Delgado et al., 2013).
Hajoale, baahi bohle ba Amerika Boroa ba P. infestans ba clonal. Ho sa tsotellehe ho ba teng ha mefuta e 'meli ea ho nyalana, ha ho na mefuta e mengata e ileng ea fumanoa. US-1 genotype e fumaneha hohle ka litamati, ho hlakile hore e tlohile litapole ke mefuta ea lehae, eo hantle-ntle e ntseng e sa tsejoe. Brazil, Bolivia le Uruguay, mofuta oa BR-1 o teng; Peru, hammoho le US-1 le EU-1, ho na le mefuta e meng e mengata ea lehae. Andes, boemo bo ka sehloohong bo bolokiloe ke moloko oa clonal EC-1, kamano ea eona le P. andina e sa tsoa sibolloa. Sebaka se le seng feela se "sa tsitsang" moo bakeng sa nako ea 2003-2013. liphetoho tse kholo tsa baahi li etsahetse, Chile e ile ea fetoha (Acuña et al., 2012), moo ka 2004-2005. palo ea pathogen e ile ea qala ho tsebahala ka ho hanyetsa metalaxyl le haplotype e ncha ea DNA ea mitochondrial (Ia sebakeng sa Ib ea pele e neng e le teng). Ho tloha 2006 ho isa 2011 palo ea baahi e ne e laoloa ke genotype 21 (ho ea ka SSR), karolo ea eona e fihlileng ho 90%, ka mor'a moo palema e fetisetsoa ho genotype 20, eo hangata lilemong tse peli tse latelang e ileng ea lula e le hoo e ka bang 67% (Acuña, 2015).
Europe
Historing ea Europe, bonyane ho ne ho e-na le maqhubu a mabeli a ho falla ha P. infestans ho tloha Amerika Leboea: lekholong la bo1 la lilemo. (HERB-1) le qalong ea lekholo la mashome a mabeli la lilemo (US-70). Tšebeliso e atileng ea li-fungicides tse nang le metalaxyl lilemong tsa bo-1. e lebisitse ho tlosoeng ha mofuta o hlahelletseng oa US-XNUMX le ho nkeloa sebaka ke mefuta e mecha. Ka lebaka leo, linaheng tse ngata tsa Europe Bophirimela, palo ea likokoana-hloko e ne e emeloa haholo-holo ke meloko e mengata ea li-clonal.
Tšebeliso ea tlhahlobo ea microsatellite ho hlahloba palo ea batho ba nang le likokoana-hloko e entse hore ho khonehe ho tseba liphetoho tse tebileng tse ileng tsa etsahala Europe Bophirimela ka 2005-2008, ho ile ha sibolloa mohala o mocha oa clonal UK, o bitsoang 2005_A13 (kapa "Blue 2"). ka mofuta oa ho nyalana oa A13, mabifi a phahameng le ho hanyetsa phenylamides (Shaw et al., 2). Mofuta o tšoanang oa genotype o ile oa fumanoa lisampoleng tse bokelletsoeng ka 2007 Netherlands le Northern France, ho fana ka maikutlo a ho fallela UK ho tsoa k'honthinenteng ea Europe, mohlomong ka litapole tsa peo (Cooke et al., 2004). Phuputso ea genome ea baemeli ba mohala ona oa clonal e bontšitse tekanyo e phahameng ea polymorphism ka tatellano ea eona (ka 2007, palo ea liphapang tsa eona tse nyenyane li fihlile ho 2016) le tekanyo e kholo ea phapang boemong ba liphatsa tsa lefutso, ho kenyeletsa. liphatsa tsa lefutso nakong ea tšoaetso ea limela (Cooke et al., 340; Cooke, 2012). Likarolo tsena, hammoho le nako e ntseng e eketseha ea karolo ea biotrophic, li ka baka bohale bo ntseng bo eketseha ba 2017_A13 le bokhoni ba eona ba ho hlasela esita le mefuta e sa tsoaneng ea tapole e hananang le blight.
Lilemong tse 'maloa tse latelang, mofuta oa genotype o ile oa ata ka potlako ho pholletsa le linaha tsa Europe Leboea-Bophirima (Great Britain, Ireland, Fora, Belgium, Netherlands, Jeremane), ha ka nako e tšoanang e tlosa li-genotypes tse neng li busa pele 1_A1, 2_A1, 8_A1 (Montarry et al. ., 2010; Gisi et al., 2011; Ho ea ka websaeteng ea www.euroblight.net, karolo ea 2011_A2015 ho baahi ba linaha tse boletsoeng e fihlile ho 2017-13% le holimo; ho ba teng ha mofuta ona oa lefutso ho boetse ho tlalehiloe linaheng tse ling tsa Europe Bochabela le Boroa. Leha ho le joalo, ka 2-60. 80_A2009 e ile ea lahleheloa ke boemo ba eona bo ka sehloohong Great Britain le Fora, e fana ka tsela ea 2012_A13 line (Ireland - 2_A6), 'me Netherlands le Belgium e ile ea nkeloa sebaka ke genotypes 1_A8, 1_A1 le 1_A6 (Cooke et al., 1; Cooke , 33; Stellingwerf, 2).
Ho fihlela joale, hoo e ka bang 70% ea baahi ba Europe Bophirimela ba P. infestans ke monoclonal. Ho ea ka webosaete ea www.euroblight.net, li-genotypes tse hlahelletseng linaheng tsa Europe Leboea-bophirima (Great Britain, France,
Netherlands, Belgium) e ntse e le teng, ka tekanyo e batlang e lekana, 13_A2 le 6_A1, 'me ea morao-rao ha e fumanehe ka ntle ho sebaka se boletsoeng (ntle le Ireland), empa e se e na le bonyane li-subclones tse 58 (Cooke, 2017). Phapang 13_A2 e teng ka lipalo tse hlokomelehang Jeremane, hape e bonoa ka linako tse ling linaheng tsa Europe Bohareng le Boroa. Genotype 1_A1 e etsa karolo e kholo ea baahi ba Belgium le likarolo tsa Netherlands le Fora. 8_A1 genotype e tsitsitse ho baahi ba Europe boemong ba 3-6%, ntle le Ireland, moo e bolokang boemo bo ka sehloohong 'me e arotsoe ka li-subclone tse peli (Stellingwerf, 2017). Qetellong, ka 2016, ho ile ha e-ba le keketseho ea makhetlo a mangata a ho hlaha ha li-genotypes tse ncha 36_A2 le 37_A2, tse ngolisitsoeng pele ka 2013-2014; ho fihlela joale, liphatsa tsa lefutso tsena li fumanoa Netherlands le Belgium le likarolong tsa Fora le Jeremane, hammoho le karolong e ka boroa ea Great Britain (Cooke, 2017). Hoo e ka bang 20-30% ea baahi ba Europe Bophirimela selemo le selemo ba emeloa ke liphatsa tsa lefutso tse ikhethang.
Ho fapana le Europe Bophirimela, ha genotype 13_A2 e hlaha, baahi ba Europe Leboea (Sweden, Norway, Denmark, Finland) ba ne ba sa emeloa ke mela ea clonal, empa ke palo e kholo ea li-genotypes tse ikhethang (Brurberg et al.,
2011). Nakong ea ho ata ha 13_A2 ho pholletsa le Europe Bophirima, boteng ba genotype ena Scandinavia ha boa ka ba tsejoa ho fihlela ka 2011, ha e qala ho fumanoa Northern Jutland (Denmark), moo mefuta e mengata ea litapole tsa indasteri e lengoang ka ts'ebeliso e matla ea metalaxyl. -e nang le li-fungicides (Nielsen et al., 2014). Ho latela www.euroblight.net, genotype 13_A2 e ile ea boela ea fumanoa lisampoleng tse 'maloa tse tsoang Norway le Denmark ka 2014 le lisampole tse' maloa tsa Norwegian ka 2016; Ho phaella moo, ka 2013, ho ba teng ha genotype 6_A1 ka bongata bo fokolang ho ile ha hlokomeloa Finland. Lebaka le ka sehloohong la ho hlōleha ha 13_A2 le mela e meng ea clonal ho hapa Scandinavia ho nkoa e le phapang ea boemo ba leholimo ea sebaka sena ho tloha linaheng tsa Europe Bophirimela.
Ho phaella tabeng ea hore lehlabula le pholileng le mariha a batang li khothalletsa ho phela ha oospores ho e-na le mycelium ea limela (Sjöholm et al., 2013), leqhoa la mobu mariha (e leng hangata le sa hlaheng linaheng tse futhumetseng tsa Europe Bophirimela) Khokahano ea ho mela ha oospore le ho lema litapole, e matlafatsang karolo ea bona e le mohloli oa tšoaetso ea mantlha (Brurberg et al., 2011). Hape hoa lokela ho hlokomeloa hore maemong a ka leboea, nts'etsopele ea ts'oaetso e tsoang ho oospores e ka pele ho nts'etsopele ea ts'oaetso ea tuber, eo qetellong e thibelang ho busa le ho feta, empa hamorao e ile ea hlahisa mela ea clonal (Yuen, 2012). Sebopeho sa batho ba ithutoang ka ho fetisisa ba P. infestans linaheng tsa Europe Bochabela (Poland, linaha tsa Baltic) li tšoana haholo le tsa Scandinavia.
Mona, hape, mefuta e 'meli ea ho nyalana e teng 'me bongata bo boholo ba li-genotypes tse khethiloeng ho sebelisa tlhahlobo ea SSR li ikhethile (Chmielarz et al., 2014; Runno-Paurson et al., 2016). Joalo ka Europe Leboea, ho ata ha mela ea clonal (haholo-holo genotype 13_A2) ha hoa ka ha ama baahi ba lehae ba pathogen, e bolokang boemo bo phahameng ba mefuta-futa ka ho se be teng ha mela e hlahelletseng.
Ho ba teng ha 13_A2 ka linako tse ling ho bonoa masimong a nang le mefuta ea litapole tsa khoebo. Naheng ea Russia boemo boa tšoana. Tlhahlobo ea Microsatellite ea P. infestans isolates e bokelitsoeng ka 2008-2011. libakeng tse fapaneng tsa 10 tsa karolo ea Europe ea Russia, li bontšitse tekanyo e phahameng ea mefuta-futa ea liphatsa tsa lefutso le ho haella ka ho feletseng ho kopana le meloko ea Europe ea clonal (Statsyuk et al., 2014). Lilemo tse 'maloa hamorao, phuputso ea lisampole tsa P. infestans tse bokelitsoeng sebakeng sa Leningrad ka 2013-2014 li bontšitse phapang e khōlō pakeng tsa bona le liphatsa tsa lefutso tse tsoang sebakeng sena tse khethiloeng thutong e fetileng. Lithutong tsena ka bobeli, ha ho li-genotype tsa Europe Bophirimela tse fumanoeng (Beketova et al., 2014; Kuznetsova et al., 2016).
Phapang e phahameng ea liphatsa tsa lefutso ea baahi ba Europe Bochabela ba P. infestans le ho se be teng ha li-clonal lineages ho bona ho ka bakoa ke mabaka a 'maloa. Taba ea pele, joalo ka Europe Leboea, maemo a leholimo a linaha tse nkoang a kenya letsoho ho thehoeng ha oospores e le mohloli oa mantlha oa ts'oaetso (Ulanova et al., 2010; Chmielarz et al., 2014). Ea bobeli, karolo e kholo ea tlhahiso ea litapole linaheng tsena e lengoa mapolasing a manyenyane a ikemetseng, hangata a pota-potiloe ke meru kapa litšitiso tse ling tsa ho tsamaea ka bolokolohi ha lintho tse tšoaetsanoang (Chmielarz et al., 2014). E le molao, litapole tse lenngoeng maemong a joalo ha li phekoloe ka lik'hemik'hale, 'me khetho ea mefuta e itšetlehile ka ho hanyetsa ha bona ka morao, i.e. ha ho na khatello e khethollang ea ho ba mabifi le ho hanyetsa metalaxyl, e thibelang li-genotype tse hanyetsanang, tse kang 13_A2, melemo ho feta mefuta e meng ea genotype (Chmielarz et al., 2014). Qetellong, ka lebaka la boholo bo fokolang ba masimo a mobu, beng ba bona hangata ha ba ikoetlise ho potoloha lijalo, ho lema litapole sebakeng se le seng ka lilemo tse ngata, e leng se tlatsetsang ho bokelloeng ha inoculum ea liphatsa tsa lefutso (Runno-Paurson et al., 2016; Elansky, 2015; Elansky et al., 2015).
Asia
Ho fihlela haufinyane tjena, sebopeho sa baahi ba P. infestans Asia se ne se ntse se sa utloisisoe hantle. Ho ne ho tsejoa hore e ne e emeloa haholo-holo ke mela ea clonal, 'me tšusumetso ea ho kopanela liphate hape ha ho hlaha liphatsa tsa lefutso tse ncha e ne e le nyenyane haholo. Kahoo, ho etsa mohlala, ka 1997-1998. karolong ea Asia ea Russia (Siberia le Bochabela bo Hōle), palo ea pathogen e ne e emeloa ke mefuta e meraro feela ea genotype e nang le boholo ba SIB-1 genotype (Elansky et al., 2001). Ho ba teng ha meloko ea kokoana-hloko ea pathogen ho bontšitsoe linaheng tse kang Chaena, Japane, Korea, Philippines le Taiwan (Koh et al., 1994; Chen et al., 2009). Leloko la US-1 la clonal le ne le laola sebaka se seholo sa Asia ho elella bofelong ba lilemo tsa bo-90 - mathoasong a bo-2000. hoo e ka bang hohle ho ile ha qala ho nkeloa sebaka ke mefuta e meng ea liphatsa tsa lefutso, eo le eona e ileng ea nkeloa sebaka ke e mecha. Maemong a mangata, liphetoho sebopeho le sebopeho sa baahi linaheng tsa Asia li ne li amahanngoa le ho falla ha liphatsa tsa lefutso tse ncha ho tsoa ka ntle. Kahoo, Japane, ntle le JP-3 genotype, mefuta eohle ea liphatsa tsa lefutso ea Majapane e hlahileng ka mor'a US-1 (JP-1, JP-2, JP-3) e na le tšimoloho ea ka ntle e netefalitsoeng kapa e fokolang (Akino et al. , 2011). Chaena, hona joale ho na le baahi ba bararo ba ka sehloohong ba pathogen, ba nang le likarohano tse hlakileng tsa libaka; ha ho na phallo ea lefutso kapa e fokolang haholo lipakeng tsa batho bana (Guo et al., 2010; Li et al., 2013b). Genotype 13_A2 e hlahile Chaena liprofinseng tsa eona tse ka boroa (Yunnan le Sichuan) ka 2005-2007, le 2012-1014. e boetse e bonoa ka leboea-bochabela ho naha (Li et al., 2013b). India, 13_A2 e hlahile ka nako e ts'oanang le ea Chaena, mohlomong ka litapole tsa peo tse tšoaelitsoeng (Chowdappa et al., 2015), le ka 2009-2010. e bakile ho phatloha ho hoholo ha morao ho tamati ka boroa ho naha, ka mor'a moo e ile ea ata ho litapole mme ka 2014 ea baka ho qhoma ha blight ea morao-rao Bengal Bophirima, e ileng ea lebisa tšenyong le ho ipolaea ha lihoai tse ngata tsa lehae (Fry, 2016).
Afrika
Ho fihlela 2008-2010. Ha ho so be le liphuputso tse hlophisitsoeng tsa P. infestans linaheng tsa Afrika. Hona joale, baahi ba Afrika ba P. infestans ba ka aroloa ka lihlopha tse peli, 'me karohano ena e amana ka ho hlaka le ho kenngoa ha litapole tsa peo ho tsoa Europe.
Afrika Leboea, e kenyang litapole tsa peo ka mafolofolo ho tsoa Europe, mofuta oa mating oa A2 o emeloa haholo hoo e batlang e le libakeng tsohle, e fanang ka monyetla oa ho hlaha ha liphatsa tsa lefutso tse ncha ka lebaka la kopano ea thobalano (Corbière et al., 2010; Rekad le tse ling, 2017). Ho phaella moo, Algeria, ho ba teng ha li-genotypes 13_A2, 2_A1 le 23_A1 ho tsejoa ka puso e totobetseng ea ba pele ba bona, hammoho le ho fokotseha butle-butle ha karolo ea li-genotypes tse ikhethang ho fihlela li nyamela ka ho feletseng (Rekad et al., 2017) . Ho fapana le libaka tse ling, Tunisia (ntle le leboea-bochabela ho naha) palo ea batho ba pathogen e emeloa haholo ke mofuta oa A1 (Harbaoui et al., 2014).
Mohala o ka sehloohong oa clonal mona ke NA-01. Ka kakaretso, karolo ea li-clonal lines sechabeng ke 43% feela. Afrika Bochabela le Boroa, moo peo e tsoang kantle ho naha e leng nyane ka mokhoa o nyamelang (Fry et al., 2009), P. infestans e emeloa ke mefuta e 'meli feela ea mofuta oa A1, US-1 le KE-1, mme ba morao-rao ba tlosa ba pele. ka litapole (Pule et al., 2012; Njoroge et al., 2016). Ho fihlela joale, liphatsa tsa lefutso tsena ka bobeli li na le palo e hlokomelehang ea mefuta e fapaneng ea subclonal.
Australia
Tlaleho ea pele ea lefu la morao-rao la litapole Australia e qalile ka 1907, 'me epiphytoty ea pele, eo ho ka etsahalang hore e bakiloe ke lipula tse matla likhoeling tsa lehlabula, e etsahetse ka 1909-1911. (Drenth et al., 2002). Leha ho le joalo, ka kakaretso, bothata ba morao-rao ha bo bohlokoa haholo moruong oa naha. Ho qhoma ha seretse ka mor'a nako, ho bakoang ke maemo a leholimo a fanang ka mongobo o phahameng, ho etsahala ha ho feta hanngoe ka mor'a lilemo tse ling le tse ling tse 5-7 'me li fumaneha haholo-holo karolong e ka leboea ea Tasmania le bohareng ba Victoria. Mabapi le se ka holimo, ha ho na lingoliloeng tse fanoeng thutong ea sebopeho sa baahi ba Australia ba P. infestans. Litaba tsa morao-rao tse fumanehang ke tsa 1998-2000. (Drenth et al., 2002). Ho ea ka bangoli, palo ea batho ba Victorian e ne e le leloko la US-1.3 la clonal, le ileng la tiisa ka mokhoa o sa tobang ho falla ha mofuta ona ho tloha United States. Lisampole tsa Tasmania li ile tsa abeloa mofuta oa AU-3, o fapaneng le mefuta ea genotype e neng e le teng ka nako eo likarolong tse ling tsa lefatše.
Likarolo tsa nts'etsopele ea bothata bo morao Russia
Europe, inoculum ea mantlha ea litapole e nkuoa e le ts'oaetso e hlahisitsoeng ka li-tubers tsa peo tse kulang, li-oospores tse koahetsoeng ke mariha mobung, hammoho le zoosporangia tse tlisoang ke moea ho tsoa ho limela tse holisitsoeng ho tsoa ho li-tubers tse nonneng masimong a selemo se fetileng (limela tsa "boithaopi"). , kapa ka liqubu tsa limela tse lahliloeng bakeng sa polokelo ea li-tubers. Ho tsena, mohloli o kotsi ka ho fetisisa oa tšoaetso o nkoa e le limela tse hōlileng ka liqubu tsa li-tubers tse lahliloeng, hobane moo palo ea li-tubers tse melang hangata e bohlokoa, 'me li-zoosporangia tse tsoang ho tsona li ka tsamaisoa libakeng tse telele. Mehloli e meng (li-oospores, "limela tsa boithaopo") ha li kotsi hakaalo, hobane Ha se tloaelo ho lema limela masimong a tšoanang ho feta hang ka mor'a lilemo tse 3-4. Tšoaetso e tsoang ho li-tubers tsa peō e kulang e boetse e fokola ka lebaka la mekhoa e metle ea ho laola boleng ba peo.
Ka kakaretso, palo ea inoculum ho baahi ba Europe e lekanyelitsoe, ka hona kholo ea seoa e lieha haholo 'me e ka laoloa ka katleho ka thuso ea lik'hemik'hale tsa fungicidal. Mosebetsi o ka sehloohong maemong a Europe ke ho loants'a ts'oaetso nakong ea ha ho qala ho hasana ho hoholo ha zoosporangia ho tsoa ho limela tse amehileng.
Naheng ea Russia, boemo bo fapane haholo. Bongata ba lijalo tsa litapole le tamati li lengoa lirapeng tse nyane tsa poraefete; mehato ea ts'ireletso e kanna ea se ke ea etsoa ho bona ho hang, kapa kalafo ea fungicidal e etsoa ka bongata bo sa lekaneng mme e qala kamora ho hlaha ha blight ea morao holimo. Ka lebaka leo, lirapa tsa poraefete li sebetsa e le mohloli o ka sehloohong oa tšoaetso; Sena se tiisoa ke maikutlo a rona a tobileng libakeng tsa Moscow, Bryansk, Kostroma le Ryazan: tšenyo ea limela lirapeng tsa botho e hlokomeloa le pele ho qala ho phekola limela tsa khoebo ka li-fungicides. Ka mor'a moo, seoa sena masimong a maholo se laoloa ke tšebeliso ea lithethefatsi tsa fungicidal, ha lirapeng tsa botho ho na le tsoelo-pele e potlakileng ea blight ea morao.
Tabeng ea mekhoa ea phekolo e fosahetseng kapa ea "tekanyetso" ea temo ea khoebo, ho hlaha hoa hlaha masimong; nakong e tlang ba nts'etsapele ka mafolofolo, ba hapa libaka tse kholoanyane (Elansky, 2015). Tšoaetso e hlahang lirapeng tsa poraefete e na le tšusumetso e kholo ho mafu a seoa libakeng tsa khoebo. Libakeng tsohle tse holang litapole tsa Russia, sebaka se tšoaretsoeng ke litapole lirapeng tsa poraefete se seholo ka makhetlo a 'maloa ho feta kakaretso ea masimo a bahlahisi ba baholo. Maemong a joalo, lirapa tsa meroho tse ikemetseng li ka nkuoa e le sesebelisoa sa lefats'e sa inoculum bakeng sa masimo a khoebo. A re leke ho khetholla thepa eo e leng tšobotsi ea li-genotypes tsa mathata lirapeng tsa botho.
Ho lema litapole tse sa kang tsa fetisa peo le taolo ea ho behelloa ka thoko, peo ea tamati e fumanoeng ho tsoa ho bahlahisi ba kantle ho naha ba belaetsang, temo ea nako e telele ea litapole le tamati libakeng tse tšoanang, kalafo e fosahetseng ka li-fungicides kapa ho ba sieo ha bona ka botlalo ho lebisa ho li-epiphytoties tse matla lekaleng la poraefete; ho fella ka ho tšela mahala, ho kopanngoa le ho thehoa ha li-oospores lirapeng tse ikemetseng. Ka lebaka leo, ho na le phapang e phahameng haholo ea genotypic ea pathogen, ha hoo e batlang e le mofuta o mong le o mong o ikhetha ho genotype ea oona (Elansky et al., 2001, 2015). Ho lema peo ea litapole tsa mefuta e fapaneng ea liphatsa tsa lefutso ho etsa hore ho se be monyetla oa hore mehala ea clonal e khethehileng ho hlasela mofuta o itseng e hlahe. Likokoana-hloko tse khethiloeng tabeng ena li khetholloa ke ho feto-fetoha ha tsona ho amanang le mefuta e mengata e amehang; Sena se fapane haholo le tsamaiso ea "clonal lines" e tloaelehileng ea masimo a maholo a mekhatlo ea temo e nang le mokhoa o nepahetseng oa ts'ireletso khahlanong le blight ea morao. "Clonal Lines" (ha mefuta eohle ea pathogen ea morao-rao tšimong e emeloa ke genotype e le 'ngoe kapa ho feta) e fumaneha hohle linaheng tseo ho tsona ho lemang litapole ho etsoang feela ke mapolasi a maholo: USA, Netherlands, Denmark, joalo-joalo. Engelane, Ireland, Poland, moo temo ea lehae e boetse e lema litapole ka moetlo, ho boetse ho na le mefuta e mengata e fapaneng ea genotypic lirapeng tse ikemetseng. "Clonal Lines" qetellong ea lekholo la bo20 la lilemo e ne e atile libakeng tsa Asia le Bochabela bo Hōle ba Russia (Elansky et al., 2001), eo ho bonahalang eka e bakoa ke tšebeliso ea mefuta e tšoanang ea ho lema e hlahisoang lapeng feela. litapole. Haufinyane tjena, maemo a libakeng tsena le ona a se a qalile ho fetoha molemong oa ho eketsa mefuta-futa ea batho.
Ho ba sieo ha liphekolo tse matla ka litokisetso tsa fungicidal ho na le phello e 'ngoe, e tobileng - ha ho na ho bokelloa ha mefuta e hanyetsanang lirapeng. Ehlile, liphetho tsa rona li bonts'a hore mefuta e hanyetsanang le metalaxyl e tsebahala hangata lirapeng tsa poraefete ho feta masimong a khoebo.
Sebaka se haufi sa ho lema litapole le tamati, se tloaelehileng bakeng sa lirapa tse ikemetseng, se thusa ho falla ha mefuta pakeng tsa lijalo tsena, ka lebaka leo, lilemong tse leshome tse fetileng, har'a mefuta e arohaneng le litapole, karolo ea mefuta e jereng mofuta oa ho hanyetsa mefuta ea tamati ea cherry (T1), eo pele e neng e le tšobotsi ea " mefuta ea tamati. Mefuta e nang le mofuta oa T1 maemong a mangata e fetoha e mabifi ho litapole le tamati.
Lilemong tsa morao tjena, blight ea morao-rao ea tamati e qalile ho hlaha maemong a mangata pejana ho feta litapole. Mohloli oa tšoaetso ea lipeo tsa tamati e ka ba li-oospores mobung kapa li-oospores tse teng kapa tse khomaretsoeng peo ea tamati (Rubin et al., 2001). Lilemong tse 15 tse fetileng, palo e kholo ea lipeo tse pakiloeng ka theko e tlase, haholo-holo tse tsoang kantle ho naha, li hlahile mabenkeleng, 'me bahlahisi ba bangata ba banyenyane ba fetohetse ho li sebelisa. Peo e kanna ea ba le mefuta e nang le li-genotypes tse tloaelehileng tsa libaka tseo li lenngoeng ho tsona. Ka mor'a moo, li-genotypes tsena li kenyelelitsoe ts'ebetsong ea thobalano lirapeng tsa botho, e leng se lebisang ho hlaha ha li-genotypes tse ncha ka ho feletseng.
Ka hona, ho ka boleloa hore lirapa tsa botho ke "pitsa e qhibilihisang" ea lefats'e moo, ka lebaka la phapanyetsano ea liphatsa tsa lefutso, li-genotype tse teng li ntse li sebetsa 'me tse ncha ka ho feletseng li hlaha. Ka nako e ts'oanang, khetho ea bona e etsahala tlas'a maemo a fapaneng haholo le a entsoeng bakeng sa litapole mapolasing a maholo: ho ba sieo ha mochine oa khatiso oa fungicidal, mefuta e fapaneng ea ho lema, ho ata ha limela tse anngoeng ke mefuta e fapaneng ea tšoaetso ea kokoana-hloko le baktheria, ho ba haufi le tamati le li-nightshades tse hlaha, ho tšela ka mafolofolo le ho thehoa ha oospore, monyetla oa hore li-oospores li sebetse e le mohloli oa tšoaetso bakeng sa selemo se tlang.
Sena sohle se lebisa ho phapano e phahameng haholo ea genotypic ea baahi ba malapeng. Tlas'a maemo a epiphytotic, blight ea morao-rao e ata ka potlako lirapeng tsa meroho 'me ho lokolloa spores tse ngata, li fofa ho ea masimong a haufi a khoebo. Leha ho le joalo, hang ha e se e le masimong a khoebo a nang le tsamaiso e nepahetseng ea thekenoloji ea temo le tšireletso ea lik'hemik'hale, li-spores tse fihlang ha li na monyetla oa ho qala epiphytoty tšimong, ka lebaka la khaello ea li-clonal tse thibelang likokoana-hloko tse khethehileng bakeng sa mefuta e sa tšoaneng e lengoang.
Mohloli o mong oa inoculum ea mantlha e ka 'na ea e-ba li-tubers tse kulang tse kentsoeng masimong a khoebo ka thepa ea peo. Li-tubers tsena li ne li hōlile, e le molao, masimong a nang le theknoloji e ntle ea temo le tšireletso e matla ea lik'hemik'hale. Li-genotypes tsa li-isolates tse amang li-tubers li ikamahanya le nts'etsopele ka mefuta ea tsona. Mefuta ena e kotsi haholo bakeng sa temo ea khoebo ho feta inoculum e tsoang lirapeng tse ikemetseng. Liphetho tsa lipatlisiso tsa rona le tsona li tšehetsa mohopolo ona. Baahi ba arohaneng le masimo a maholo a nang le ts'ireletso ea lik'hemik'hale hantle le theknoloji e ntle ea temo ha ba khethollehe ka mefuta e mengata ea genotypic. Hangata tsena ke mela e mengata ea clonal e mabifi haholo.
Lithapo tse tsoang peo ea khoebo li ka kenella ho batho ba lirapeng tsa meroho mme tsa kenella lits'ebetsong tse etsahalang ho tsona. Leha ho le joalo, serapeng sa meroho, tlholisano ea bona e tla ba tlase haholo ho feta lebaleng la khoebo, 'me haufinyane ba tla emisa ho ba teng ka mokhoa oa clonal line, empa liphatsa tsa bona tsa lefutso li ka sebelisoa ho batho ba "serapeng".
Tšoaetso e hlahang limeleng tsa "boithaopo" le liqubung tsa li-tubers tse lahliloeng nakong ea kotulo ha li bohlokoa hakaalo bakeng sa Russia, hobane Libakeng tse kholo tse holang litapole tsa Russia, serame se tebileng sa mariha se bonoa, 'me limela tse tsoang ho li-tubers tse koahetsoeng ke mobu ha li hole hangata. Ho feta moo, joalo ka ha liteko tsa rona li bonts'a, kokoana-hloko ea blight ea morao-rao ha e pholohe mochesong o ka tlase ho zero esita le li-tubers tse lulang li sebetsa. Sebakeng se omeletseng, moo ho lengoang litapole tsa pele, blight ea morao e fumaneha seoelo ka lebaka la nako e omileng le e chesang ea ho hola.
Kahoo, hajoale re ntse re shebile karohano ea batho ba P. infestans ho ba "masimong" le "lirapa". Leha ho le joalo, lilemong tsa morao tjena, lits'ebetso li hlokometsoe li lebisa ho kopaneng le ho kenella ha liphatsa tsa lefutso ho tsoa ho baahi bana.
Har'a tsona, motho a ka hlokomela keketseho e akaretsang ea ho bala le ho ngola ea bahlahisi ba banyane, ponahalo ea liphutheloana tse theko e tlase tsa litapole tsa peo, ho ata ha litokisetso tsa fungicidal lipaketeng tse nyane, le tahlehelo ea tšabo ea "chemistry" ke baahi.
Maemo a hlaha ha, ka lebaka la mosebetsi o mafolofolo oa mofani a le mong, metse eohle e iphumana e lenngoe ka li-tubers tsa peō tsa mefuta e sa tšoaneng le ho fuoa liphutheloana tse nyenyane tsa chefo e bolaeang likokoanyana. Hoa khoneha ho nahana hore litapole tsa mofuta o tšoanang le tsona li tla hlaha masimong a khoebo a haufi.
Ka lehlakoreng le leng, lik’hamphani tse ling tse hoebang ka chefo e bolaeang likokoanyana li buella merero ea phekolo ea lik’hemik’hale e “tšehang litšenyehelo tse tlaase”. Tabeng ena, palo ea liphekolo tse khothalelitsoeng ha li nyenyefatsoe 'me ho fanoa ka li-fungicides tse theko e tlaase,' me khatiso ha se ho thibela nts'etsopele ea lefu la morao-rao ho fihlela ho poma litlhōrō, empa ka ho lieha ho itseng ho epiphytoty e le ho eketsa Khefutsa. Merero e joalo e na le mabaka a utloahalang a moruo ha ho hōla litapole tsa ware ho tloha lihlahisoa tsa peo ea boemo bo tlaase, ha ka molao-motheo ho se na potso ea ho fumana chai e ngata. Leha ho le joalo, tabeng ena, ho fapana le baahi ba lirapa, liphatsa tsa lefutso tsa litapole li kenya letsoho tabeng ea ho khetha merabe e itseng ea fisioloji, e leng kotsi haholo bakeng sa mefuta e fapaneng.
Ka kakaretso, mekhoa ea ho kopana ha "serapa" le "tšimo" ea tlhahiso ea litapole e bonahala e le kotsi ho rona. Ho thibela litlamorao tsa bona tse mpe makaleng a malapa le a khoebo, ho tla hlokahala ho laola mefuta e fapaneng ea litapole tsa peo le mefuta e mengata ea li-fungicides tse fuoang barekisi ba ikemetseng ka liphutheloana tse nyane, le ho lekola merero ea ts'ireletso ea litapole le ts'ebeliso ea litokisetso tsa fungicidal. lefapha la khoebo.
Libakeng tsa lekala la poraefete, ho na le nts'etsopele e matla eseng feela ea ts'oaetso ea morao, empa le Alternaria. Boholo ba beng ba mapolasi a ikemetseng ha ba nke mehato e ikhethileng ho itšireletsa khahlanong le Alternaria, ba nka phoso ea nts'etsopele ea Alternaria bakeng sa ho pona ha makhasi a tlhaho kapa nts'etsopele ea bothata ba morao. Ka hona, ka kholo e kholo ea Alternaria ka mefuta e ka hlaseloang habonolo, merero ea malapa e ka sebetsa e le mohloli oa inoculum bakeng sa temo ea khoebo.
Mekhoa ea ho fetoha
Mokhoa oa phetoho
Kaha ketsahalo ea liphetoho e le ts'ebetso e sa reroang e etsahalang ka makhetlo a tlase, ho hlaha ha liphetoho sebakeng sefe kapa sefe ho ipapisitse le maqhubu a phetoho ea sebaka sena le boholo ba baahi. Ha u ithuta khafetsa ea liphetoho tsa mefuta ea P. infestans, palo ea li-colone tse hōlileng holima mecha ea litaba ea limatlafatsi ka mor'a ho phekoloa ka lik'hemik'hale kapa tsa 'mele tsa mutagen hangata li khethoa. Joalokaha ho ka bonoa ho data e hlahisitsoeng ho Lethathamo la 8, khafetsa ea phetoho ea mofuta o tšoanang libakeng tse fapaneng e ka fapana ka litaelo tse 'maloa tsa boholo. Maqhubu a phahameng a ho hanyetsa metalaxyl e ka ba le leng la mabaka a ho bokella mefuta e hanyetsanang le eona ka tlhaho.
Khafetsa ea liphatsa tsa lefutso tse iketsahallang feela kapa tse bakiloeng, tse baloang ho latela liteko tsa laboratori, ha se kamehla li tsamaellanang le lits'ebetso tse etsahalang ho batho ba tlhaho ka mabaka a latelang:
1. Ka likarohano tsa nyutlelie tse sa tsitsang, ha ho khonehe ho hakanya makhetlo a liphetoho tsa liphatsa tsa lefutso molokong oa nyutlelie. Ka hona, liteko tse ngata li fana ka tlhahisoleseding ka ho toba feela ka makhetlo a mangata a liphetoho, ntle le ho khetholla pakeng tsa liketsahalo tse peli tsa phetoho le ketsahalo e le 'ngoe e latelang mitosis.
2. Liphetoho tsa mohato o le mong hangata li fokotsa ho leka-lekana ha genome, ka hona, hammoho le ho fumana thepa e ncha, ho phela hantle ka kakaretso hoa fokotseha. Liphetoho tse ngata tse fumanoeng ka liteko li fokolitse bohale 'me ha lia tlalehoa ho batho ba tlhaho. Ka hona, coefficient corelation pakeng tsa tekanyo ea ho hanyetsa ha P. infestans mutants ho phenylamide fungicides le sekhahla sa kgolo ka seaplane ea maiketsetso e ne e le ka karolelano (-0,62), le ho hanyetsa fungicides le mabifi makhasi a litapole (-0,65) (Derevyagina et al., 1993), e bonts'ang ho fokola ha liphetoho tse fetotsoeng. Liphetoho tsa ho hanyetsa dimethomorph li ne li boetse li tsamaea le ho fokotseha ho matla ha katleho (Bagirova et al., 2001).
3. Bongata ba liphatsa tsa lefutso tse iketsahallang feela le tse susumetsoang li fetoha ka linako tse ling 'me ha li iponahatse ka mokhoa o makatsang litekong, empa li etsa pokello e patehileng ea ho fetoha ha batho ba tlhaho. Mathata a feto-fetohang a ikhethileng litekong tsa laboratori a na le liphetoho tse kholo kapa tse matla (Kulish le Dyakov, 1979). Kamoo ho bonahalang kateng, diploidity ea nuclei e hlalosa boiteko bo sa atleheng ba ho fumana liphetoho tlas'a tšusumetso ea mahlaseli a UV a neng a le kotsi ho mefuta e neng e hanyetsa pele (McKee, 1969). Ho ea ka lipalo tsa mongoli, liphetoho tse joalo li ka etsahala ka makhetlo a ka tlase ho 1: 500000. Phetoho ea liphetoho tse feteletseng ho ea ho homozygous, boemo bo hlalositsoeng ka mokhoa oa phenotypically bo ka etsahala ka lebaka la ho kopanya thobalano kapa ho kopanela liphate (sheba ka tlase). Leha ho le joalo, esita le tabeng ena, phetoho e ka 'na ea koaheloa ke li-alleles tse hlaheletseng tsa li-nuclei tse hlaha ka har'a coenotic (multinucleate) mycelium' me e be phenotypically e tsitsitseng feela nakong ea ho thehoa ha li-zoospores tsa mononuclear.
Letlapa la 8. Khafetsa ea liphetoho tsa P. infestans ho hōla lintho tse thibelang ho hōla tlas'a tšusumetso ea nitrosomethylurea (Dolgova, Dyakov, 1986; Bagirova et al., 2001)
Khahlano | Phetoho ea phetoho |
Oxytetracycline | 6,9 x 10-8 |
Blasticidin S | 7,2: x 10-8 |
Streptomycin | 8,3 x10-8 |
Trichothecin | 1,8 x 10-8 |
Cycloheximide | 2,1 x 10-8 |
Daconil | <4 x 10-8 |
Dimethomorph | 6,3 x 10-7 |
Metalaxyl | 6,9 x 10-6 |
Bongata ba baahi le bona bo phetha karolo ea bohlokoa ho hlaheng ha liphatsa tsa lefutso tse itlelang feela. Ho baahi ba bangata haholo, moo palo ea lisele e leng N> 1/a, moo a e leng sekhahla sa phetoho, phetoho e khaotsa ho ba ntho e sa tloaelehang (Kvitko, 1974).
Lipalo li bontša hore ka karolelano ea ts'oaetso ea tšimo ea litapole (matheba a 35 semeleng se le seng), li-spores tse 8x1012 li thehoa letsatsi le leng le le leng ka hektare e le 'ngoe (Dyakov, Suprun, 1984). Kamoo ho bonahalang kateng, lichabeng tse joalo liphetoho tsohle tse lumelloang ke mofuta oa phapanyetsano li etsahala sebakeng se seng le se seng. Esita le phetoho e sa tloaelehang e hlahang ka makhetlo a 10-9 e tla fumanoa ke batho ba sekete ho ba limilione ba phelang hektare e le 'ngoe ea tšimo ea litapole. Bakeng sa liphetoho tse etsahalang ka sekhahla se phahameng (mohlala, 10-6), mefuta e fapaneng ea liphetoho tse peli (ka nako e le 'ngoe) e ka etsahala letsatsi le leng le le leng ho baahi ba joalo, ke hore. mokhoa oa ho fetola liphatsa tsa lefutso o tla nkela sebaka sa ho kopanya hape.
Ho falla
Mefuta e 'meli e meholo ea ho falla e tsejoa ka P. infestans: ho feta libaka tse khutšoanyane (ka har'a tšimo ea litapole kapa masimo a haufi) ka ho qhala li-zoosporangia ka maqhubu a moea kapa lipula tsa pula, le libakeng tse telele - ka li-tubers tse lemang kapa litholoana tsa tamati tse tsamaisoang. Mokhoa oa pele o tiisa ho atolosoa ha sepheo sa lefu lena, ea bobeli - ho thehoa ha li-foci tse ncha libakeng tse hōle le tse ka sehloohong.
Ho ata ha tšoaetso ka li-tubers tsa langa le le lej le litholoana ha ho thuse feela ho hlaha ha lefu lena libakeng tse ncha, empa hape ke mohloli o ka sehloohong oa mefuta-futa ea liphatsa tsa lefutso tsa baahi. Litapole tse tsoang libakeng tse fapaneng tsa Russia le Europe Bophirimela li lengoa sebakeng sa Moscow. Litholoana tsa tamati li tlisoa libakeng tse ka boroa tsa Russia (sebaka sa Astrakhan, sebaka sa Krasnodar, North Caucasus). Peo ea tamati, e ka boela ea sebetsa e le mehloli ea tšoaetso (Rubin et al., 2001), e boetse e romelloa ho tsoa libakeng tse ka boroa tsa Russia, Chaena, linaheng tsa Europe le linaheng tse ling.
Ho ea ka lipalo tse entsoeng ke E. Mayr (1974), liphetoho tsa liphatsa tsa lefutso ho baahi ba sebaka seo li bakoang ke ho fetoha ha liphatsa tsa lefutso ha li fete ka seoelo 10-5 ka sebaka se seng le se seng, ha libakeng tse bulehileng phapanyetsano ka lebaka la phallo ea liphatsa tsa lefutso ha e ka tlaase ho 10-3 - 10-4. .
Ho falla ha li-tubers tse nang le tšoaetso ho ikarabella bakeng sa ho kena ha P. infestans Europe le ho hasana ho pholletsa le libaka tsohle tsa lefats'e moo litapole li lengoang; li baka liphetoho tse tebileng ka ho fetisisa tsa baahi. Blight ea morao-rao ea litapole e ile ea hlaha sebakeng sa Mmuso oa Russia hoo e ka bang ka nako e le 'ngoe le ponahalo ea eona Europe Bophirimela.
Kaha lefu lena le ile la hlokomeloa ka lekhetlo la pele ka 1846-1847 linaheng tsa Baltic 'me feela lilemong tse latelang li namela Belarus le libakeng tse ka Leboea-Bophirima tsa Russia, tšimoloho ea eona ea Bophirimela ea Europe e totobetse. Mohloli oa pele oa lefu la morao-rao Lefatšeng la Khale ha o totobetse hakaalo. Khopolo-taba e hlahisitsoeng ke Fry et al., 1992, Goodwin, 1995, Goodwin et al., 1994 e fana ka maikutlo a hore kokoana-hloko e tsoa Mexico ho ea Amerika Leboea, moo e ileng ea ata ka lijalo, eaba e isoa Europe Bophirimela (Fig. 7).
Ka lebaka la ho hoholeha khafetsa (phello e habeli ea botlolo), li-clones tse le 'ngoe li ile tsa kena Europe, bana ba bona ba ileng ba baka seoa sebakeng sohle sa Lefatše la Khale, moo ho lengoang litapole. E le bopaki ba khopolo-taba ena, bangoli ba qotsa, pele, ketsahalo e atileng ea mofuta o le mong feela oa ho nyalana (A1) mme, ea bobeli, homogeneity ea liphatsa tsa lefutso tsa mefuta e ithutoang e tsoang libakeng tse fapaneng (kaofela ha tsona ho latela matšoao a limolek'hule, ho kenyeletsoa 2 isoenzyme loci, lipaterone tsa DNA tsa ho hatisa menoana le sebopeho sa DNA ea mitochondrial lia ts'oana 'me li tsamaisana le clone ea US-1 e hlalositsoeng USA). Leha ho le joalo, lintlha tse ling li baka lipelaelo bonyane lipehelong tse ling tsa khopolo-taba e boletsoeng. Tlhahlobo ea DNA ea mitochondrial ea P. infestans, e arohaneng le lisampole tsa herbarium tsa litapole tse tšoaelitsoeng nakong ea epiphytoty ea pele ea 40s ea lekholo la bo1 la lilemo, li bontšitse hore ka sebopeho sa DNA ea mitochondrial li fapane le clone US-2001, eo ka hona e neng e le teng. bonyane eseng eona feela mohloli oa tšoaetso Europe (Ristaino et al, XNUMX).
Boemo ba lefu la morao-rao bo ile ba mpefala hape ka bo-80. Ho etsahetse liphetoho tse latelang:
1) Karolelano ea mabifi a baahi e eketsehile, e leng se lebisitseng, haholo-holo, ho ata ha mofuta o kotsi ka ho fetisisa oa blight ea morao-rao - tšenyo ea li-petioles le stems.
2) Ho bile le phetoho nakong ea ho hlaha ha blight ea morao ho litapole - ho tloha bofelong ba Phupu ho isa qalong ea Phupu esita le ho isa bofelong ba Phuptjane.
3) Mofuta oa ho nyalana oa A2, oo pele o neng o le sieo Lefatšeng la Khale, o ile oa qala ho fumanoa hohle.
Liphetoho li ile tsa etelloa pele ke liketsahalo tse peli: ts'ebeliso e atileng ea fungicide metalaxyl e ncha (Schwinn le Staub, 1980) le ho hlaha ha Mexico e le morekisi oa litapole lefatšeng ka bophara (Niederhauser, 1993). Ho ea ka sena, ho behiloe mabaka a mabeli a liphetoho tsa baahi: phetoho ea mofuta oa ho nyalana tlas'a tšusumetso ea metalaxyl (Ko, 1994) le kenyelletso e kholo ea mefuta e mecha e nang le li-tubers tse nang le tšoaetso tse tsoang Mexico (Fry and Goodwin, 1995). Le hoja li-interconversions tsa mefuta ea mating tlas'a tšusumetso ea metalaxyl ha lia ka tsa fumanoa feela ke Ko, empa hape le mosebetsing o etsoang laboratoring ea Moscow State University (Savenkova, Chherepennicova-Anikina, 2002), khopolo ea bobeli e molemo. Hammoho le ho hlaha ha mofuta oa bobeli oa ho nyalana, ho ile ha e-ba le liphetoho tse tebileng liphatseng tsa lefutso tsa mefuta ea Russia ea P. infestans, ho akarelletsa le liphatsa tsa lefutso tse sa nke lehlakore (isoenzyme le RFLP loci), hammoho le sebopeho sa DNA ea mitochondrial. Ho rarahana ha liphetoho tsena ho ke ke ha hlalosoa ke ketso ea metalaxyl; ho e-na le hoo, ho ne ho e-na le ho kenngoa ha mefuta e mecha e tsoang Mexico, eo, ho ba mabifi le ho feta (Kato et al., 1997), e nkile sebaka sa mefuta ea khale (US-1) , ho ba matla har'a baahi. Phetoho ea sebopeho sa baahi ba Europe e etsahetse ka nako e khuts'oane haholo - ho tloha 1980 ho isa 1985 (Fry et al., 1992). Sebakeng sa USSR ea mehleng, ho ile ha sibolloa "mefuta e mecha" likoleke tse tsoang Estonia ka 1985, ke hore, pejana ho Poland le Jeremane (Goodwin et al., 1994). Lekhetlo la ho qetela ha "sekhahla sa khale sa US-1" naheng ea Russia se qheletsoe ka thoko ho tamati e tšoaelitsoeng sebakeng sa Moscow e ne e le ka 1993 (Dolgova et al., 1997). Hape Fora, mefuta ea "khale" e ile ea fumanoa masimong a tamati ho fihlela mathoasong a lilemo tsa bo-90, ke hore, ka mor'a hore e nyamele nako e telele ho tloha litapole (Leberton, Andrivon, 1998). Liphetoho mefuteng ea P. infestans li ile tsa ama litšobotsi tse ngata, ho kenyelletsa le tsa bohlokoa ba bohlokoa, 'me tsa eketsa kotsi ea blight ea morao-rao.
Ho kopana hape ka thobalano
E le hore recombination ea thobalano e kenye letsoho ho feto-fetoha, hoa hlokahala, pele, bakeng sa ho ba teng ha mefuta e 'meli ea ho nyalana ho baahi ka karolelano e haufi le 1: 1,' me, ea bobeli, bakeng sa ho ba teng ha phapang ea pele ho baahi.
Karo-karolelano ea mefuta ea ho nyalana e fapana haholo ho batho ba fapaneng esita le lilemong tse fapaneng ka palo e le 'ngoe (Letlapa la 9,10, 90). Mabaka a liphetoho tse makatsang joalo ka maqhubu a mefuta ea ho nyalana ka bongata (ka mohlala, Russia kapa Iseraele mathoasong a lilemo tsa bo-2002 lekholong la ho qetela la lilemo) ha a tsejoe, empa ho lumeloa hore sena se bakoa ke ho hlahisoa ha tlhōlisano e eketsehileng. clones (Cohen, XNUMX).
Lintlha tse ling tse sa tobang li bonts'a ketsahalo ea thobalano lilemong tse itseng le libakeng tse ling:
1) Liphuputso tsa batho ba tsoang sebakeng sa Moscow li bontšitse hore ho baahi ba 13 moo karolo ea mofuta oa A2 ea ho nyalana e neng e le ka tlase ho 10%, kakaretso ea mefuta-futa ea liphatsa tsa lefutso e baloang ho tloha ho tse tharo tsa loci ea isoenzyme e ne e le 0,08, le ho baahi ba 14 moo karolo ea A2 e feta 30%, mefuta e fapaneng ea liphatsa tsa lefutso e ne e phahame habeli (0,15) (Elansky et al., 1999). Kahoo, ha monyetla oa ho kopanela liphate o phahame, ho ba le mefuta e mengata ea liphatsa tsa lefutso ea baahi.
2) Kamano lipakeng tsa karo-karolelano ea mefuta ea ho nyalana ka bongata le matla a sebopeho sa oospore e ile ea bonoa Israel (Cohen et al., 1997) le Holland.
(Flier et al., 2004). Liphuputso tsa rona li bonts'itse hore har'a batho ba arohaneng le mofuta oa A2 ba nkang karolo ea 62, 17, 9 le 6%, li-oospores li fumanoe ho 78, 50, 30 le 15% ea makhasi a hlahlobiloeng a litapole (a nang le matheba a 2 kapa ho feta), ka ho latellana.
Mehlala e nang le matheba a 2 kapa ho feta e ne e na le li-oospores hangata ho feta lisampole tse nang le letheba le le leng (1 le 32% ea lisampole ka ho latellana) (Apryshko et al., 14).
Li-Oospores li ne li atile haholo makhasi a karolo e bohareng le e tlase ea limela tsa litapole (Mytsa et al., 2015; Elansky et al., 2016).
3) Libakeng tse ling, li-genotype tse ikhethang li ile tsa sibolloa, ketsahalo ea eona e amanang le ho kopanya thobalano. Ka hona, Poland ka 1989 le Fora ka 1990, ho ile ha fumanoa mefuta ea homozygous bakeng sa glucose-6-locus.
phosphate isomerase (GPI 90/90). Kaha pele ke li-heterozygote tse 10/90 feela tse ileng tsa kopana le tsona nakong ea lilemo tse 100, homozygosity e bakoa ke ho kopana hape ka thobalano (Sujkowski et al., 1994). Colombia (USA), libaka tse ka thōko tse kopanyang A2 le GPI 100/110 le A1 le GPI 100/100 li tloaelehile, empa qetellong ea nako ea 1994 (August 16 le September 9), mathata a nang le li-genotypes tse ncha (A1 GPI 100/110). ) li ile tsa sibolloa le A2 GPI 100/100) (Miller et al., 1997).
4) Ho batho ba bang ba tsoang Poland (Sujkowski et al., 1994) le North Caucasus (Amatkhanova et al., 2004), kabo ea menoana ea DNA loci le allozyme protein loci e tsamaisana le kabo ea Hardy-Weinberg, e bonts'ang.
mabapi le tlatsetso e phahameng ea ho kopanya thobalano ho feto-fetoha ha baahi. Libakeng tse ling tsa Russia, ha ho ngollano ho kabo ea Hardy-Weinberg e ileng ea fumanoa ho baahi, empa boteng ba ho se leka-lekane ha likamano ho ile ha bontšoa, ho bontša boholo ba ho ikatisa ha clonal (Elansky et al., 1999).
5) Phapang ea liphatsa tsa lefutso (GST) lipakeng tsa mefuta e nang le mefuta e fapaneng ea ho nyalana (A1 le A2) e ne e le tlase ho feta lipakeng tsa batho ba fapaneng (Sujkowski et al., 1994), e bonts'ang ka mokhoa o sa tobang likamano tsa thobalano.
Ka nako e ts'oanang, tlatsetso ea ho kopanya thobalano ho mefuta e fapaneng ea baahi e ke ke ea e-ba holimo haholo. Monehelo ona o ne o baloa bakeng sa baahi ba sebaka sa Moscow (Elansky et al., 1999). Ho ea ka lipalo tsa Lewontin (1979), "recombination, e ka hlahisang mefuta e mecha ho tloha ho li-loci tse peli ka makhetlo a sa feteng sehlahisoa sa li-heterozygosities tsa bona, e sebetsa hafeela litekanyetso tsa heterozygosity bakeng sa alleles ka bobeli li se li phahame."
Ka karolelano e tloaelehileng ea mefuta e 'meli ea ho nyalana sebakeng sa Moscow e le 4: 1, khafetsa ea ho kopanya e tla ba 0,25. Monyetla oa hore mefuta e fapaneng e tla ba heterozygous bakeng sa tse peli ho tse tharo tse ithutoang tsa isoenzyme loci ho baahi ba ithutoang e ne e le 0,01 (2 mefuta ho tsoa ho 177). Ka hona, monyetla oa ho hlaha ha li-heterozygote tse peli ka lebaka la ho kopanya hape ha oa lokela ho feta sehlahisoa sa bona se atolositsoeng ke monyetla oa ho tšela (0,25x0,02x0,02) = 10-4, i.e. li-recombinants tsa thobalano hangata ha li kenyelelitsoe sampoleng e ithutoang ea mefuta. Lipalo tsena li entsoe bakeng sa baahi ba tsoang sebakeng sa Moscow, ba khetholloang ka phapang e batlang e phahame. Ho batho ba monomorphic joalo ka ba Siberia, ts'ebetso ea thobalano, leha e ka etsahala ho batho ka bomong, e ke ke ea susumetsa mefuta ea bona ea liphatsa tsa lefutso.
Ho phaella moo, P. infestans e khetholloa ke tlōlo ea khafetsa ea karohano ea chromosome ka meiosis, e lebisang ho aneuploidy (Carter et al., 1999). Litšitiso tse joalo li fokotsa ho nona ha mefuta e nyalisitsoeng.
Parasexual recombination, phetoho ea liphatsa tsa lefutso tsa mitotic
Litekong tsa ho kopanya P. infestans mefuta e nang le liphetoho tsa ho hanyetsa li-inhibitors tse fapaneng tsa kholo, ho ile ha sibolloa ho hlaha ha li-isolate tse hanyetsanang le li-inhibitors ka bobeli (Shattock, Shaw, 1975; Dyakov, Kuzovnikova, 1974; Kulish, Dyakov,
1979). Mefuta e hanyetsanang le li-inhibitors tse peli tsa kholo e hlahile ka lebaka la heterokaryotization ea mycelium, 'me tabeng ena e ile ea aroloa nakong ea ho ikatisa ke li-zoospores tse mononuclear (Judelson, Ge Yang, 1998), kapa ha lia ka tsa aroloa ka leloko la monozoospore, hobane li ne li e-na le tetraploid (kaha li-isolate tsa pele ke diploid) nuclei (Kulish, Dyakov, 1979). Heterozygous diploids e arotsoe ka lebelo le tlase haholo ka lebaka la haploidization, chromosome nondisjunction, le mitotic crossing over (Poedinok et al., 1982). Khafetsa ea lits'ebetso tsena e ka eketsoa ka litlamorao tse itseng ho li-heterozygous diploids (mohlala, mahlaseli a UV a li-spores tse melang).
Le ha ho etsoa ha li-hybrids tsa limela tse hanyetsanang habeli ha li hlahe feela ka har'a vitro, empa hape le ho li-tubers tsa litapole tse tšoaelitsoeng ke motsoako oa li-mutants (Kulish et al., 1978), ho thata haholo ho lekola karolo ea ho kopanya parasexual molokong oa batho. li-genotypes tse ncha ka bongata. Khafetsa ea ho thehoa ha lihlopha ka lebaka la haploidization, chromosome nondisjunction le mitotic crossing ntle le tšusumetso e khethehileng ha e na thuso (tlase ho 10-3).
Ho hlaha ha li-homozygous segregants tsa mefuta ea heterozygous li ka thehoa ho bobeli ba mitotic crossing over le mitotic gene conversion, eo ho P. sojae e hlahang ka makhetlo a 3 x 10-2 ho 5 x 10-5 ka locus, ho itšetlehile ka mofuta oa khatello () Chamnanpunt et al., 2001).
Le hoja khafetsa ea ho hlaha ha li-heterokaryone le li-diploid tsa heterozygous li ile tsa phahama ka mokhoa o sa lebelloang (ho fihlela liperesente tse mashome), ts'ebetso ena e etsahala feela ha litloaelo tse fetotsoeng tse fumanoang mofuteng o tšoanang li kopantsoe. Ha u sebelisa mefuta e fapaneng e arohaneng le tlhaho, heterokaryotization ha e hlahe (kapa e etsahala ka maqhubu a tlaase haholo) ka lebaka la ho ba teng ha limela tse sa lumellaneng le limela (Poedinok, Dyakov, 1981; Anikina et al., 1997b; Cherepennikova-Anikina et al., 2002) ). Ka lebaka leo, karolo ea parasexual recombination e ka fokotsoa feela ho intraclonal recombination in heterozygous nuclei le phetoho ea liphatsa tsa lefutso ho ea boemong ba homozygous ntle le ts'ebetso ea thobalano. Ts'ebetso ena e kanna ea ba ea bohlokoa ba lefu la mafu mefuteng e nang le phetoho e fetelletseng kapa e matla ea fungicide. Phetoho ea eona ho ea boemong ba homozygous ka lebaka la ts'ebetso ea parasexual e tla eketsa khanyetso ea mochine oa phetoho (Dolgova, Dyakov, 1986).
Gene selelekela
Mefuta ea Heterothallic Phytophthora e khona ho tsoalisana ho etsa li-oospores tse nyalisitsoeng (sheba Vorobyova le Gridnev, 1983; Sansome et al., 1991; Veld et al., 1998). Lebasetere la tlhaho la mefuta e 'meli ea Phytophthora le ne le le mabifi hoo le ileng la senya likete tsa li-alders Great Britain (Brasier et al., 1999). P. infestans ka etsahala le mefuta e meng ea genus (P. erythroseptica, P. nicotianae, P. Cactorum, joalo-joalo) ka limela tse tloaelehileng tse amohelang le mobung, empa ho na le boitsebiso bo fokolang ka lingoliloeng mabapi le monyetla oa ho interspecific poone e nyalisitsoeng etsahala. . Li-hybrids pakeng tsa P. infestans le P. Mirabilis li ile tsa fumanoa tlas'a maemo a laboratori (Goodwin le Fry, 1994).
Letlapa la 9. Palo ea mefuta ea P. infestans e nang le mofuta oa A2 oa ho nyalana linaheng tse fapaneng tsa lefats'e nakong ea ho tloha 1990 ho ea ho 2000 (ho ea ka mehloli e bulehileng ea lingoliloeng le liwebsaete www.euroblight.net, www.eucablight.org)
naha | 1990 | 1991 | 1992 | 1993 | 1994 | 1995 | 1996 | 1997 | 1998 | 1999 | 2000 |
---|---|---|---|---|---|---|---|---|---|---|---|
Belarus | 33 (12) | 34 (29) | |||||||||
Belgium | 15 (49 *) | 6 (66) | 20 (86) | ||||||||
Ecuador | 0 (13) | 0 (12) | 0 (19) | 0 (21) | 12 (41) | 25 (39) | 15 (75) | 22 (73) | 25 (68) | 0 (35) | |
Estonia | 8 (12) | ||||||||||
Engelane | 4 (26) | 3 (630) | 9 (336) | ||||||||
Finland | 0 (15) | 19 (117) | 12 (16) | 21 (447) | 6 (509) | 9 (432) | 43 (550) | ||||
France | 0 (35) | 0 (56) | 0 (83) | 0 (67) | 0 (86) | 2 (135) | 7 (156) | 6 (123) | 0 (73) | 0 (285) | 0 (135) |
Hungary | 72 (32) | ||||||||||
Ireland | 4 (145) | ||||||||||
Leboya Ireland | 10 (41) | 9 (58) | 1 (106) | 0 (185) | 0 (18) | 0 (56) | 0 (35) | 0 (26) | |||
Netherlands | 7 (41) | 5 (276) | 24 (377) | 44 (353) | 23 (185) | ||||||
Norway | 25 (446) | 28 (156) | 8 (39) | 18 (257) | 38 (197) | ||||||
Peru | 0(34, 1984 -86) | 0(287, 1997-98) | 0 (112) | 0 (66) | |||||||
Poland | 19 (180) | 21 (142) | 33 (256) | 26 (149) | 35 (70) | ||||||
Scotland | 25 (147) | 11 (163) | 22 (189) | 5 (22) | |||||||
Sweden | 25 (263) | 62 (258) | 49 (163) | ||||||||
Wales | 0 (16) | 7 (97) | 0 (48) | 0 (25) | |||||||
Korea | 36 (42) | 10 (130) | 15 (98) | ||||||||
China | 20(142, 1995-98) | 0 (6) | 0 (8) | 0 (35) | |||||||
Colombia | 0(40, 1994-2000) | ||||||||||
Uruguay | 100(25, 1998-99) | ||||||||||
Morocco | 60(108, 1997-2000) | 52 (25) | 42 (40) | ||||||||
Serbia | 76 (37) | ||||||||||
Mexico (Toluca) | 28(292, 1988-89) | 50(389, 1997-98) |
Letlapa la 10. Palo ea mefuta ea P. infestans e nang le mofuta oa A2 ea ho nyalana linaheng tse fapaneng tsa lefats'e ho tloha 2000 ho isa 2011.
naha | 2001 | 2002 | 2003 | 2004 | 2005 | 2006 | 2007 | 2008 | 2009 | 2010 | 2011 |
---|---|---|---|---|---|---|---|---|---|---|---|
Austria | 65 (83) | ||||||||||
Belarus | 42 (78) | ||||||||||
Belgium | 20 (102 *) | 4 (32) | 50 (14) | 25 (16) | 62 (13) | 54 (26) | 70 (54) | 30 (23) | 29 (35) | 62 (71) | 45 (49) |
Switzerland | 89 (19) | ||||||||||
Czech Republic | 35 (31) | 54 (64) | 38 (174) | 12 (80) | |||||||
Duitsland | 95 (53) | ||||||||||
Denmark | 48 (52) | ||||||||||
Ecuador | 5 (178) | 6 (108) | 9 (121) | 18 (94) | 2 (44) | 0 (66) | 5 (47) | ||||
Estonia | 54 (25) | 0 (24) | 33 (62) | 45 (140) | 25 (100) | 12 (103) | |||||
Engelane | 4 (47) | 10 (96) | 31 (55) | 55 (790) | 68 (862) | 70 (552) | 68 (299) | ||||
Finland | 47 (162) | 12 (218) | 42 | ||||||||
France | 0 (186) | 4 (108) | 8 (61) | 22 (103) | 33 (303) | 65 (378) | 74 (331) | 75 (125) | 75 (12) | ||
Hungary | 48 (27) | 48 (90) | 9 | 7 | |||||||
Leboya Ireland | 0 (38) | 0 (58) | 0 (40) | 0 (24) | 5 (54) | 0 (18) | 27 (578) | 45 (239) | 36 (213) | 82 (60) | 10 (80) |
Netherlands | 66 (24) | 93 (15) | 91 (11) | ||||||||
Norway | 39 (328) | 3 (115) | 12 (19) | ||||||||
Peru | 0 (36) | ||||||||||
Poland | 25 (46) | 10 (30) | 85 (20) | 38 (44) | 75 (66) | 55 (56) | 65 (35) | 72 (81) | 85 (21) | ||
Scotland | 3 (213) | 2 (474) | 24 (135) | 86 (337) | 88 (386) | 74 (172) | |||||
Sweden | 60 (277) | 39 (87) | |||||||||
Slovakia | 0 (36) | 14 (26) | 62 (26) | 0 (26) | |||||||
Wales | 25 (12) | 68 (106) | 80 (88) | 92 (143) | 75 (45) | ||||||
Korea | 46 (26) | ||||||||||
Brazil | 0 (49) | 0 (30) | |||||||||
China | 10 (30) | 0 (6) | 0 (6) | ||||||||
Вьетнам | 0(294, 2003-04) | ||||||||||
Uganda | 0 (8) |
Matla a sebopeho sa genotypic ea baahi
Liphetoho tsa sebopeho sa genotypic tsa P. infestans baahi ba ka etsahala tlas'a tšusumetso ea ho falla ha li-clone tse ncha ho tloha libakeng tse ling, mekhoa ea temo (ho fetoha ha mefuta e sa tšoaneng, tšebeliso ea li-fungicides) le maemo a leholimo. Litšusumetso tsa kantle li ama li-clones ka tsela e fapaneng maemong a fapaneng a potoloho ea bophelo;
Phello ea mefuta-futa
Mefuta e mecha e nang le liphatsa tsa lefutso tse matla tsa ho hanyetsa (R-genes) ke khetho e matla e khethang li-clone tse nang le liphatsa tsa lefutso tse tlatsetsang ho batho ba P. infestans. Haeba mefuta e mengata ea litapole e se na khanyetso e sa tobang e thibelang kholo ea palo ea batho ba pathogen, mokhoa oa ho fetola li-clones tse hlahelletseng sechabeng o etsahala kapele haholo. Ka hona, ka mor'a ho ata ha mefuta e sa tšoaneng ea Domodedovo, e nang le mofuta oa R3 oa ho hanyetsa, sebakeng sa Moscow, makhetlo a mangata a li-clones tse kotsi bakeng sa mefuta ena e fapaneng a eketsehile ho tloha ho 0,2 ho ea ho 0,82 ka selemo se le seng (Dyakov, Derevjagina, 2000).
Leha ho le joalo, liphetoho tsa maqhubu a liphatsa tsa lefutso tsa virulence (pathotypes) ho baahi ha li hlahe feela tlas'a tšusumetso ea mefuta e sa tšoaneng ea litapole. Ka mohlala, Belarus, ho fihlela ka 1977, li-clones tse nang le liphatsa tsa lefutso tsa 1 le 4 li ne li laola, tse neng li bakoa ke ho lengoa ha mefuta ea litapole tse nang le mefuta ea ho hanyetsa R1 le R4 (Dorozhkin le Belskaya, 1979). Leha ho le joalo, qetellong ea lilemo tsa bo-70 tsa lekholo la mashome a mabeli la lilemo, li-clone li ile tsa hlaha ka mefuta e fapaneng ea likokoana-hloko le motsoako oa tsona, 'me mefuta ea tsona ea ho hanyetsa ha e so ka e sebelisoa ho ikatisa litapole (liphatsa tsa lefutso tse eketsehileng) (Ivanyuk et al., 2002). Lebaka la ho hlaha ha li-clones tse joalo ka ho hlakileng ke ka lebaka la ho falla ha lintho tse tšoaetsanoang ho tloha Mexico ka li-tubers tsa litapole ho ea Europe. Lapeng, li-clone tsena ha lia ka tsa hlaha feela ka litapole tse lenngoeng, empa le ho mefuta e hlaha e nang le mefuta e fapaneng ea ho hanyetsa, ka hona, motsoako oa liphatsa tsa lefutso tse ngata ho genome o ne o hlokahala bakeng sa ho phela maemong ao.
Ha e le mefuta e sa tšoaneng e nang le khanyetso e sa tsitsang, ka ho fokotsa sekhahla sa ho ikatisa ha kokoana-hloko, e lieha ho fetoha ha baahi ba eona, eo, joalokaha ho se ho boletsoe, ke mosebetsi oa lipalo. Kaha aggressiveness ke polygenic, clones e nang le palo e kholo ea liphatsa tsa lefutso tsa "bohloeki" li bokellana ka potlako ho feta palo ea baahi. Ka hona, merabe e mabifi haholo ha se sehlahisoa sa ho ikamahanya le mefuta e lenngoeng e nang le khanyetso e sa tobang, empa, ho fapana le hoo, e fumanoa ho lenngoeng ha mefuta e kotsing haholo, e leng li-accumulators tsa likokoana-hloko.
Ka hona, Russia, baahi ba mabifi ka ho fetisisa ba P. infestans ba ile ba fumanoa libakeng tsa li-epiphytoties tsa selemo le selemo (baahi ba tsoang libakeng tsa Sakhalin, Leningrad, Bryansk). Bokhopo ba batho bana bo ile ba fetoha bo phahameng ho feta ba Mexico (Filippov et al., 2004).
Ntle le moo, ho thehoa li-oospores tse fokolang makhasi a mefuta e hanyetsanang ho feta ho tse hlaselehang habonolo (Hanson le Shattock, 1998), ke hore, khanyetso e sa tobang ea mefuta e boetse e fokotsa bokhoni ba ho kopanya likokoana-hloko le monyetla oa mekhoa e meng ea mariha.
Phello ea li-fungicides
Li-fungicides ha li fokotse feela palo ea li-fungus tsa phytopathogenic, i.e. susumetsa litšobotsi tsa bongata ba baahi ba bona, empa hape e ka fetola maqhubu a liphatsa tsa lefutso ka bomong, i.e. susumetsa sebopeho sa boleng ba baahi. Har'a matšoao a bohlokoa ka ho fetisisa a baahi ba fetohang tlas'a tšusumetso ea li-fungicides ke tse latelang: liphetoho tsa ho hanyetsa li-fungicides, liphetoho tse mabifi le likokoana-hloko, le liphetoho tsamaisong ea ho ikatisa.
Tšusumetso ea li-fungicides ho hanyetsa le ho ba mabifi ha baahi
Tekanyo ea tšusumetso e joalo e khethoa, pele ho tsohle, ka mofuta oa fungicide e sebelisitsoeng, e ka aroloa ka libaka tse ngata, oligo-site le mono-site.
Ea pele e kenyelletsa li-fungicides tse ngata tsa ho kopana. Ho hanyetsa ho bona (haeba ho khoneha ho hang) ho laoloa ke palo e kholo ea liphatsa tsa lefutso tse fokolang haholo. Lintho tsena li etsa qeto ea ho ba sieo ha liphetoho tse bonahalang khanyetso ea baahi ka mor'a ho e phekola ka li-fungicides (le hoja litekong tse ling ho ile ha fumanoa keketseho e fokolang ea ho hanyetsa). Palo ea fungal e bolokiloeng kamora ho fafatsa ka li-fungicides e na le lihlopha tse peli tsa mefuta:
1) Mathata a bolokiloeng libakeng tsa limela tse sa phekoloang ka lithethefatsi. Kaha ho ne ho se na ho kopana le fungicide, bohale le ho hanyetsa ha mefuta ena ha e fetohe.
2) Mathata a ileng a kopana le fungicide, eo mahloriso a eona lintlheng tsa ho kopana a neng a le ka tlase ho bolaeang. Joalokaha ho boletsoe ka holimo, khanyetso ea karolo ena ea baahi le eona ha e fetohe, leha ho le joalo, ka lebaka la phello e senyang ea fungicide, esita le maemong a mangata, ho metabolism ea sele ea fungal, ho ikoetlisa ka kakaretso le karolo ea eona ea likokoana-hloko - ho ba mabifi - ho fokotseha (Derevyagina, Dyakov, 1990).
Ka hona, esita le karolo e sa shoeng ea baahi ba pepesehetseng ho kopana le fungicide e na le bohale bo fokolang 'me e ke ke ea e-ba mohloli oa epiphytoty. Ka hona, kalafo e hlokolosi, e fokotsang khafetsa ea palo ea baahi ba sa kopane le fungicide, ke boemo ba katleho ea mehato ea ts'ireletso. Ho hanyetsa oligosite fungicides ho laoloa ke liphatsa tsa lefutso tse 'maloa tsa tlatsetso.
Phetoho ea liphatsa tsa lefutso e 'ngoe le e' ngoe e lebisa keketseho e fokolang ea khanyetso, 'me tekanyo ea kakaretso ea ho hanyetsa e khethoa ke ho eketsoa ha liphetoho tse joalo. Ka hona, ho eketseha ha khanyetso ho etsahala ka mekhahlelo. Mohlala oa ho eketseha ha mohato oa ho hanyetsa ke liphetoho tsa ho hanyetsa fungicide dimethomorph, e sebelisoang haholo ho sireletsa litapole ho tloha ho senya morao. Ho hanana le dimethomorph ke polygenic le tlatsetso. Phetoho ea mohato o le mong e eketsa khanyetso hanyane.
Phetoho e 'ngoe le e' ngoe e latelang e fokotsa boholo ba sepheo, ka lebaka leo, khafetsa ea liphetoho tse latelang (Bagirova et al., 2001). Keketseho ea khanyetso e tloaelehileng ea baahi ka mor'a ho phekoloa khafetsa ka oligosite fungicide e etsahala ka mohato le butle-butle. Sekhahla sa ts'ebetso ena se khethoa ke bonyane lintlha tse tharo: makhetlo a mangata a phetoho ea liphatsa tsa lefutso tsa khanyetso, coefficient ea ho hanyetsa (karolelano ea tekanyo e bolaeang ea mofuta o hanyetsanang o amanang le o bobebe) le phello ea liphetoho tsa liphatsa tsa lefutso ho ho phela hantle mmeleng.
Khafetsa ea ketsahalo ea phetoho e 'ngoe le e' ngoe e latelang e tlase ho feta e fetileng, kahoo ts'ebetso e fokotsehile (Bagirova et al., 2001). Leha ho le joalo, haeba mekhoa ea recombination (ho kopanela liphate kapa parasexual) e etsahala har'a baahi, joale hoa khoneha ho kopanya liphetoho tse fapaneng tsa batsoali ka mofuta o nyalisitsoeng le ho potlakisa ts'ebetso. Ka hona, palo ea batho ba panmix e fumana khanyetso ka potlako ho feta ea agamic, 'me qetellong, baahi ba se nang litšitiso tsa ho se lumellane ha limela, ka potlako ho feta baahi ba arotsoeng ke mekoallo e joalo. Ntlheng ena, ho ba teng ka bongata ba mefuta e fapaneng ka mefuta ea ho nyalana ho potlakisa mokhoa oa ho fumana khanyetso ho oligosite fungicides.
Lintlha tsa bobeli le tsa boraro ha li kenye letsoho ho bokellaneng ka potlako ha mefuta e hanyetsanang le dimethomorph ka bongata. Phetoho e 'ngoe le e' ngoe e latelang e batla e imena habeli ho hanyetsa, e seng ea bohlokoa, 'me ka nako e ts'oanang e fokotsa sekhahla sa kholo tikolohong ea maiketsetso le mabifi (Bagirova et al., 2001; Stem and Kirk, 2004). Ke ka lebaka lena har'a mefuta ea tlhaho ea P. infestans, esita le tse bokelletsoeng masimong a litapole a tšoaroang ka dimethomorph, ho hang ha ho na tse hanyetsanang.
Baahi ba phekoloang ka oligosite fungicide le bona ba tla ba le lihlopha tse peli tsa mefuta: tse neng li sa kopane le fungicide mme ka hona ha lia ka tsa fetola litšobotsi tsa pele (haeba mefuta e hanyetsanang e fumanoa har'a sehlopha sena, e ke ke ea bokellana ka lebaka la bohale bo phahameng le tlholisano ea mefuta e nang le kutloelo-bohloko), le mefuta e pepesehileng ka bongata bo sa bolaeang ba sebolaea-khuru. Ke har'a ba morao-rao moo ho ka khonehang ho bokella mefuta e sa tsitsang, hobane mona ba na le melemo ho feta tse mahlonoko.
Ka hona, ha u sebelisa oligosite fungicides, se bohlokoa ha se phekolo e hlokolosi haholo joalo ka khatello e phahameng ea lithethefatsi, makhetlo a 'maloa ho feta tekanyo e bolaeang, hobane ka stepwise mutagenesis khanyetso ea pele ea mefuta e fetotsoeng e tlase.
Qetellong, liphetoho tsa ho hanyetsa li-fungicides tsa monosite li hlalosa haholo, ho bolelang hore phetoho e le 'ngoe e ka fana ka maemo a phahameng a ho hanyetsa ho fihlela tahlehelo e feletseng ea kutlo. Ka hona, keketseho ea khanyetso ea baahi e etsahala kapele haholo.
Mehlala ea li-fungicides tse joalo ke phenylamides, ho kenyelletsa le metalaxyl e atileng haholo ea fungicide. Liphetoho tsa ho hanyetsa ho eona li etsahala ka makhetlo a mangata, 'me tekanyo ea ho hanyetsa liphatseng tsa lefutso e phahame haholo - e feta khatello ea maikutlo ka makhetlo a sekete kapa ho feta (Derevyagina et al., 1993). Le ha sekhahla sa kholo le ho halefa ha liphetoho tse hanyetsanang li fokotseha khahlano le semelo sa lefu la mefuta e bobebe ho tsoa ho fungicide ea systemic, palo ea batho ba hanyetsanang e ntse e hola ka potlako mme, ka nako e ts'oanang, mabifi a eona a ntse a eketseha. Ka hona, kamora lilemo tse 'maloa tsa ho sebelisa fungicide, bohale ba mefuta e hanyetsanang e ke ke ea lekana feela le bohale ba mefuta e bobebe, empa e bile e feta (Derevyagina, Dyakov, 1992).
Phello ho kopanyo ea thobalano
Kaha ho hlaha khafetsa ha mofuta oa A2 ho batho ba P. infestans ho tsamaellana le tšebeliso e matla ea metalaxyl khahlano le blight ea morao, ho ile ha etsoa tlhahiso ea hore metalaxyl e baka phetoho ea mefuta ea ho nyalana. Mofuteng oa P. parasitica, phetoho e joalo tlas'a tšusumetso ea chloroneb le metalaxyl e netefalitsoe ka liteko (Ko, 1994). Tsela e le 'ngoe e nang le motsoako o fokolang oa metalaxyl e lebisitse ho hlaha ha homothallic isolates ho tloha mofuteng oa metalaxyl-sensitive oa P. infestans e nang le mofuta oa A1 oa ho nyalana (Savenkova, Cherepnikova-Anikina, 2002). Nakong ea litemana tse latelang ho mecha ea litaba e nang le metalaxyl e phahameng, ho ne ho sa khonehe ho lemoha sekhetho se le seng se nang le mofuta oa A2, leha ho le joalo, boholo ba libaka tse arohaneng, ha li tšeloa ka li-isolates tsa A2, sebakeng sa oospores, li ile tsa theha lihlopha tse mpe tsa mycelium 'me li ne li nyopa. . Litemana tsa khatello e hanyetsanang e nang le mofuta oa A2 oa ho kopanya mecheng ea litaba e nang le metalaxyl e ngata e entse hore ho khonehe ho lemoha mefuta e meraro ea liphetoho mofuteng oa ho nyalana: 1) sterility e feletseng ha e tšeloa ka lihlopha tsa A1 le A2; 2) homothallism (sebopeho sa oospores monoculture); 3) phetoho ea mofuta oa A2 oa ho nyalana ho A1. Ka hona, metalaxyl e ka 'na ea e-ba sesosa sa liphetoho mefuteng ea ho nyalana ho batho ba P. infestans' me, ka lebaka leo, ketsahalo ea ho kopanya thobalano ho bona.
Phello ho recombination ea limela
Liphatsa tsa lefutso tse ling tsa lithibela-mafu li ile tsa eketsa khafetsa ea heterokaryotization ea hyphae le diploidization ea nuclei (Poedinok le Dyakov, 1981). Joalokaha ho boletsoe pejana, heterokaryotization ea hyphae nakong ea ho kopanngoa ha mefuta e sa tšoaneng ea P. infestans e etsahala ka seoelo haholo ka lebaka la ketsahalo ea ho se lumellane ha limela ho fungus ena. Leha ho le joalo, liphatsa tsa lefutso tsa ho hanyetsa lithibela-mafu tse ling li ka ba le litla-morao tse fellang ka ho hlōla ho se lumellane ha limela. Mofuta oa ho hanyetsa streptomycin oa 1S-1 o fetotsoeng o ne o e-na le thepa ena. Ho ba teng ha liphetoho tse joalo sebakeng sa Phytophthora ho ka eketsa phallo ea liphatsa tsa lefutso lipakeng tsa mefuta le ho potlakisa ho ikamahanya le maemo a batho bohle ho mefuta e mecha kapa li-fungicides.
Likokoana-hloko tse ling le lithibela-mafu li ka ama khafetsa ho kopanngoa ha mitotic, tse ka fetolang maqhubu a genotypes ka bongata. Benomyl ea fungicide e sebelisoang haholo e tlama ho beta-tubulin, protheine eo li-microtubules tsa cytoskeleton li hahiloeng ho eona, 'me ka tsela eo e senya mekhoa ea ho arohana ha chromosome ho anaphase of mitosis, ho eketsa khafetsa ea mitotic recombination (Hastie, 1970).
Sebolaya dikokwana-hloko sa para-fluorophenylalanine, se sebedisoang ho alafa elms ho tswa ho lefu la maDutch, se na le thepa e tshwanang. Para-fluorophenylalanine e boetse e ekelitse khafetsa ea ho kopanya hape ka heterozygous diploids ea P. infestans (Poedinok et al., 1982).
Liphetoho tsa cyclic ho sebopeho sa genotypic sa baahi nakong ea potoloho ea bophelo ba P. infestans
Potoloho ea khale ea nts'etsopele ea P. infestans sebakeng se futhumetseng e na le mekhahlelo e 4.
1) Mokhahlelo oa kholo ea kholo ea baahi (polycyclic phase) e nang le meloko e khutšoane. Hangata mohato ona o qala ka July mme o nka likhoeli tse 1,5-2.
2) Mokhahlelo oo keketseho ea baahi e emisang ka lebaka la ho fokotseha ho hoholo ha karolo ea lisele tse sa ameheng kapa ho qala ha maemo a leholimo a sa thabiseng. Mapolasing a tlosang litlhōrō pele ho kotuloa, karolo ena e oela ka ntle ho potoloho ea selemo le selemo.
3) Karolo ea overwintering ka li-tubers, e tsamaeang le ho fokotseha ho hoholo ha palo ea baahi ka lebaka la ts'oaetso ea kotsi ea li-tubers, nts'etsopele e liehang ea ts'oaetso ho tsona, ho ba sieo ha ts'oaetso ea li-tubers tlas'a maemo a tloaelehileng a polokelo, ho bola le ho lahla li-tubers tse amehileng. .
4) Karolo ea tsoelo-pele e liehang mobung le lipeo (monocyclic phase), eo nako ea moloko o ka fihlang khoeli kapa ho feta (bofelong ba May - mathoasong a July). Hangata ka nako ena, makhasi a kulang a thata ho lemoha esita le ka litebello tse khethehileng.
Mokhahlelo o moholo oa kholo ea baahi (polycyclic phase)
Litlhahlobo tse ngata (Pshedetskaya, Kozubova, 1969; Borisnok, 1969; Osha, 1969; Dyakov, Suprun, 1984; Rybakova, Dyakov, 1990) e bontšitse hore qalong ea epiphytoty, li-clones tse fokolang le tse fokolang hamorao li fetoha tse fokolang. Ho feta moo, sekhahla sa ho eketseha ha sekhahla sa baahi se phahame, 'me mefuta e sa tšoaneng ea semela e sa khone ho mamella.
Ha palo ea baahi e ntse e eketseha, ho kopana ha liphatsa tsa lefutso tsa bohlokoa ka ho khetheha tse kentsoeng mefuteng ea khoebo (R1-R4) le tse sa nkeng lehlakore (R5-R11) ho eketseha. Ka hona, ho baahi ba haufi le Moscow ka 1993, karolelano ea likokoana-hloko ho tloha bofelong ba July ho ea bohareng ba August e eketsehile ho tloha ho 8,2 ho ea ho 9,4, ka keketseho e kholo ka ho fetisisa e hlokometsoeng bakeng sa mofuta o sa nkeng lehlakore oa R5 (ho tloha ho 31 ho ea ho 86% ea li-clones tse kotsi) (Smirnov, 1996).
Ho fokotseha ha sekhahla sa ho hōla ha baahi ho tsamaisana le ho fokotseha ha mosebetsi oa likokoana-hloko oa baahi. Ka hona, lilemong tsa khatello ea maikutlo, palo eohle ea merabe le karolo ea merabe e kotsi haholo e tlase ho feta lilemong tsa epiphytotic (Borysenok, 1969). Haeba, bophahamong ba epiphytoty, maemo a leholimo a fetoha a sa thabiseng bakeng sa blight ea morao-rao mme ts'oaetso ea litapole e fokotseha, joale khatello ea li-clones tse kotsi haholo le tse mabifi le tsona lia fokotseha (Rybakova et al., 1987).
Keketseho ea maqhubu a liphatsa tsa lefutso tse susumetsang bohlasoa le mabifi a sechaba mohlomong e etsahala ka lebaka la khetho ea li-clone tse mabifi le tse mabifi ho batho ba tsoakaneng. Ho bonts'a khetho, ho entsoe mokhoa oa ho sekaseka liphetoho tse sa nke lehlakore mme o sebelisitsoe ka katleho ho chemostat populations of yeast (Adams et al., 1985) le Fusarium graminearum (Wiebe et al., 1995).
Khafetsa ea liphetoho tse hanyetsanang le blasticidin S tšimong ea baahi ba P. infestans e fokotsehile ho bapisa le ho eketseha ha mabifi a baahi, e leng se bontšang phetoho ea li-clones tse hlaheletseng nakong ea kholo ea baahi (Rybakova et al., 1987) .
Mokhahlelo oa overwintering ka li-tubers
Nakong ea mariha ka li-tubers tsa litapole, likokoana-hloko le mabifi a P. infestans li fokotseha, 'me ho fokotseha ha likokoana-hloko ho etsahala butle ho feta ka mabifi (Rybakova, Dyakov, 1990). Kamoo ho bonahalang kateng, tlas'a maemo a khothalletsang khōlo e potlakileng ea baahi (r-khetho), "e eketsehileng" liphatsa tsa lefutso le mabifi a phahameng a fetoha a molemo, ka hona, nts'etsopele ea epiphytoty e tsamaisana le khetho ea li-clones tse kotsi ka ho fetisisa le tse mabifi. Tlas'a maemo a ho tlala ha tikoloho, ha e se sekhahla sa ho ikatisa se phethang karolo e kholo, empa ho phehella ho ba teng maemong a sa thabiseng (K-khetho), "e eketsehileng" liphatsa tsa lefutso le mabifi li fokotsa 'mele,' me clones e nang le tsena. liphatsa tsa lefutso ke tsona tsa pele tse shoang, e le hore karolelano ea mabifi le likokoana-hloko tsa baahi li fokotsehe.
Mokhahlelo oa limela mobung
Mokhahlelo ona ke o makatsang ka ho fetisisa oa potoloho ea bophelo (Andrivon, 1995). Ho ba teng ha eona ho boletsoe ka mokhoa o inahaneloang - ka lebaka la khaello ea tlhahisoleseling mabapi le se etsahalang ho pathogen ka nako e telele (ka linako tse ling ho feta khoeli) - ho tloha ho hlaha ha lipeo tsa litapole ho ea ho ponahalo ea matheba a pele a lefu lena. bona. Ho itšetlehile ka maikutlo le liteko, boitšoaro ba fungus nakong ena ea bophelo bo ile ba tsosolosoa (Hirst, Stedman, 1960; Boguslavskaya, Filippov, 1976).
Sporulation ea fungus e ka theha li-tubers tse nang le tšoaetso mobung. Li-spores tse hlahisoang li mela ka hyphae, e khonang ho mela mobung nako e telele. E ka sehloohong (e entsoeng ka li-tubers) le ea bobeli (ka mycelium mobung) li-spores li nyolohela mobung ka maqhubu a capillary, empa li fumana bokhoni ba ho tšoaetsa litapole feela ka mor'a hore makhasi a tsona a tlase a theohe 'me a kopane le mobu. Makhasi a joalo (e leng, matheba a pele a lefu lena a fumanoa ho bona) ha a thehoe hang-hang, empa ka mor'a khōlo ea nako e telele le nts'etsopele ea litlhōrō tsa litapole.
Ka hona, karolo ea limela tsa saprotrophic e ka boela ea e-ba teng nakong ea bophelo ba P. infestans. Haeba mokhahlelong oa likokoana-hloko oa potoloho ea bophelo bo mabifi ke karolo ea bohlokoa ka ho fetisisa ea ho ikoetlisa, joale khethong ea saprotrophic phase e reretsoe ho fokotsa thepa ea likokoana-hloko, joalokaha ho bontšitsoe ka liteko bakeng sa li-fungus tse ling tsa phytopathogenic (bona Carson, 1993). Ka hona, karolong ena ea potoloho, thepa e mabifi e lokela ho lahleha ka matla haholo. Empa ho fihlela joale, ha ho liteko tse tobileng tse entsoeng ho tiisa maikutlo a entsoeng.
Liphetoho tsa nako ha li ame feela thepa ea pathogenic ea P. infestans, empa hape e hanyetsa li-fungicides, e ntseng e eketseha karolong ea polycyclic (nakong ea epiphytoties) le ho fokotseha nakong ea polokelo ea mariha (Derevyagina et al., 1991; Kadish, Cohen, 1992). Ho fokotseha ho matla haholo ha ho hanyetsa metalaxyl ho ile ha hlokomeloa nakong ea pakeng tsa ho lema li-tubers tse amehileng le ponahalo ea matheba a pele a lefu lena tšimong.
Intraspecific specialization le ho iphetola ha eona
P. infestans e baka mafu a seoa lijalong tse peli tsa bohlokoa khoebong, litapole le tamati. Li-epiphytoties ka litapole li qalile kapele ka mor'a hore fungus e kene libakeng tse ncha. Ho hlōloa ha langa le le lej ho ile ha boela ha hlokomeloa kapele ka mor'a hore tšoaetso e hlahe ka litapole, empa li-epiphytoties ka langa le le lej li ile tsa tsejoa lilemo tse lekholo hamorao - bohareng ba lekholo la mashome a mabeli la lilemo. Sena ke seo ba se ngolang mabapi le ho hloloa ha tamati USA ke Gallegly le Niederhauser
(1962): “Ka lilemo tse ka bang 100 ka mor’a hore ho be le epiphytoty e matla haholo ea 1845, ho ile ha etsoa boiteko bo fokolang kapa ha hoa ka ha etsoa ho hang ho hlahisa mefuta e manganga ea tamati. Le hoja lefu la sekoahelo la morao le ile la tlalehoa ka lekhetlo la pele litamati morao koana ka 1848, ha ea ka ea e-ba ntho eo balisanisi ba semela sena ba neng ba e ela hloko haholo ho fihlela ha lefu lena le qhoma ka matla ka 1946.” Naheng ea Russia, tomate ea morao-rao e ile ea ngolisoa lekholong la bo60 la lilemo. “Ka nako e telele, bafuputsi ba ne ba sa ele hloko lefu lena, kaha ha lea ka la baka tšenyo e khōlō moruong. Empa ka 70-1979s. Lekholong la boXNUMX la lilemo, li-epiphytoties tsa morao-rao tsa tamati li ile tsa boela tsa bonoa Soviet Union, haholo-holo sebakeng sa Lower Volga, Ukraine, North Caucasus le Moldova ... "(Balashova, XNUMX).
Ho tloha ka nako eo, tomate blight e fetohile ketsahalo ea selemo le selemo, e hasana hohle sebakeng sa liindasteri le lehae la temo le ho baka tšenyo e kholo ea moruo sejalong sena. Ho etsahetse eng? Ke hobane'ng ha ponahalo ea pele ea likokoana-hloko ho litapole le tšenyo ea epiphytotic sejalong see e ile ea etsahala hoo e ka bang ka nako e le 'ngoe, athe ho ile ha nka lilemo tse lekholo hore tšenyo ea epiphytotic e hlahe ka tamati? Liphapang tsena li rata motho oa Mexico ho e-na le mohloli oa tšoaetso oa Amerika Boroa. Haeba mefuta ea Phytophthora infestans e thehiloe e le likokoana-hloko tsa mefuta ea Mexican tuberous ea genus Solanum, joale ho hlakile hore na ke hobane'ng ha litapole tse lengoang, tse leng karolo ea mofuta o le mong le mefuta ea Mexico, li ile tsa ameha haholo, empa ka lebaka la khaello. ea coevolution le likokoana-hloko, ha baa ka ba hlahisa mekhoa ea khanyetso e tobileng le e sa tobang.
Tomate ke ea karolo e fapaneng ea mofuta, mofuta oa metabolism ea eona e na le phapang e kholo ho mefuta ea li-tuberous, ka hona, ho sa tsotellehe taba ea hore tamati ha e ka ntle ho lijo tse khethehileng tsa P. infestans, matla a tšenyo ea eona a ne a sa lekana. bakeng sa tahlehelo e tebileng ea moruo.
Ketsahalo ea li-epiphytoties ka langa le le lej e bakoa ke liphetoho tse tebileng tsa liphatsa tsa lefutso ho likokoana-hloko, tse eketsang boemo ba eona bo botle (pathogenicity) nakong ea likokoana-hloko. Re lumela hore mofuta o mocha, o khethehileng bakeng sa parasitism ka tamati, ke lebelo la T1 le hlalositsoeng ke M. Gallegly, e tšoaetsang mefuta ea tamati ea ciliegia (Red Cherry, Ottawa), e hanyetsanang le lebelo la T0 le tloaelehileng ho litapole (Gallegly, 1952). Kamoo ho bonahalang kateng, phetoho (kapa letoto la liphetoho) e ileng ea fetola lebelo la T0 peisong ea T1 e lebisitse ho hlaha ha li-clones tse ikamahanyang haholo le tšenyo ea tamati. Joalo ka ha ho etsahala hangata, ho eketseha ha pathogenicity ho moamoheli a le mong ho ne ho tsamaisana le ho fokotseha ha eona ho ea ho e mong, ke hore, ho ile ha hlaha tsebo ea pele, e e-so phethehe, ea intraspecific - ho litapole (peiso ea T0) le tamati (peiso ea T1).
Ke bopaki bofe bo tšehetsang khopolo ee?
- Ho etsahala ka litapole le tamati. Makhasing a tamati, lebelo la T1 ke lona le hlahelletseng, athe makhasi a litapole ha a fumanehe. Ho latela S.F Bagirova le T.A. Oreshonkova (e sa hatisoang) sebakeng sa Moscow ka 1991-1992, ketsahalo ea peiso ea T1 ha ho lema litapole e ne e le 0%, 'me ho lema tamati - 100%; ka 1993-1995 - 33% le 90%, ka ho latellana; ka 2001 - 0% le 67%. Lintlha tse tšoanang li ile tsa fumanoa Iseraele (Cohen, 2002). Liteko tsa ts'oaetso ea li-tubers tsa litapole tse nang le li-isolates tsa morabe oa T1 le motsoako oa li-isolates T0 le T1 li bonts'itse hore li-isolates tsa morabe oa T1 ha lia bolokoa hantle ka har'a li-tubers mme li nkeloa sebaka ke lihlopha tsa morabe T0 (Dyakov et al., 1975; Rybakova, 1988) .
2) Matla a peiso T1 ho lema tamati. Tšoaetso ea mantlha ea makhasi a langa le le lej e etsoa ke li-isolate tsa peiso ea T0, e laolang tlhahlobo ea tšoaetso libakeng tsa pele tse entsoeng makhasi. Sena se tiisa mokhoa o amoheloang ka kakaretso oa ho falla ha likokoana-hloko: 'mele o ka sehloohong oa tšoaetso e tsoang ho litapole ke lebelo la T0, leha ho le joalo, palo e nyane ea li-clones tsa peiso T1 e bolokiloeng ka litapole, hang ka tamati, e tlosa peiso ea T0 ebe e bokellana ho ea qetellong ea epiphytoty. Hape hoa khoneha hore ho na le mohloli o mong oa tšoaetso ea makhasi a tamati ka lebelo la T1, eseng matla joaloka li-tubers tsa litapole le makhasi, empa kamehla. Ka hona, mohloli ona o na le phello e fokolang ho sebopeho sa liphatsa tsa lefutso tsa baahi ba tšoaetsang langa le le lej, empa hamorao ba etsa qeto ea ho bokella lebelo la T1 (Rybakova, 1988; Dyakov et al., 1994).
3) Ho ba mabifi ho litapole le tamati. Tšoaetso ea maiketsetso ea makhasi a tamati le makhasi a litapole a arohaneng le merabe ea T0 le T1 e bontšitse hore tsa pele li mabifi ho feta litapole ho feta tamati, 'me tse qetellang li le mabifi bakeng sa tamati ho feta litapole. Phapang ena e bonahala ka ho falla ha batho ba merabe e fapaneng ho tloha ho batho ba tsoakaneng nakong ea litemana tse holim'a makhasi a sethopo (Dyakov et al., 1975) le libakeng tsa masimo (Leberton et al., 1999); phapang pakeng tsa boima bo fokolang ba tšoaetso, nako ea latency, boholo ba mabala a tšoaetsanoang le tlhahiso ea spore (Rybakova, 1988; Dyakov et al., 1994; Legard et al., 1995; Forbes et al., 1997; Oyarzun et al., 1998; Leberton et al., 1999; 2000;
Bokhopo ba peiso ea T1 bo itšehla thajana ho mefuta ea tamati e se nang liphatsa tsa lefutso tsa khanyetso bo phahame haholo hoo li-sporulate tse makhasing joalo ka setsing sa limatlafatsi ntle le necrosis ea lisele tse tšoaelitsoeng (Dyakov et al., 1975; Vega-Sanchez et al., 2000).
4) Virulence bakeng sa litapole le tamati. Race T1 e tšoaetsa mefuta ea tamati ea ciliegia e nang le mofuta oa ho hanyetsa oa Ph1, ha morabe oa T0 o sa khone ho tšoaetsa mefuta ena, ke hore. e na le virulence e fokolang. Mabapi le li-different
Liphatsa tsa litapole tsa R li bontša kamano e fapaneng, ke hore. mefuta e arohaneng le makhasi a tamati ha e kotsi ho feta "litapole" (Letlapa la 11).
5) Matšoao a sa nke lehlakore. Multidirectional intraspecific khetho e boetse e pakoa ke tlhahlobo ea matšoao a sa nke lehlakore ho baahi ba P. infestans parasitizing litapole le tamati. Ho baahi ba Brazil ba P. infestans, makhasi a tamati e ne e le a US-1 clonal lineage, le ho tloha makhasi a litapole ho ea lelokong la BR-1 (Suassuna et al., 2004). Florida (USA), ho tloha ka 1994, li-clone tsa US-90 li ile tsa qala ho busa litapole (ketsahalo e fetang 8%), 'me ho tamati, li-clones tsa US-11 le US-17,' me likaroloana tsa ho qetela li le mabifi ho feta tamati. ho feta bakeng sa litapole (Weingartner, Tombolato, 2004). Phapang e kholo ea maqhubu a genotype (menoana ea menoana ea DNA) lipakeng tsa litapole le tamati e arohaneng e thehiloe bakeng sa mefuta ea 1200 P. infestans e bokeletsoeng USA ho tloha 1989 ho isa 1995 (Deahl et al., 1995).
Tšebeliso ea mokhoa oa AFLP e entse hore ho khonehe ho arola mefuta e 74 e bokelletsoeng makhasi a litapole le tamati ka 1996-1997. Fora le Switzerland, ka lihlopha tse 7. Mefuta ea litapole le tamati ha ea ka ea fapana ka ho hlaka, empa mefuta ea "litapole" e ile ea fetoha liphatsa tsa lefutso ho feta mefuta ea "tamati". Ea pele e fumanoe ka lihlopha tse supileng, athe tsa morao-rao li ile tsa fumanoa ho tse 'nè feela, tse bontšang genome e khethehileng ea ho qetela (Knapova le Gisi, 2002).
6) Mekhoa ea ho itšehla thajana. Haeba likokoana-hloko tsa mefuta e 'meli ea limela tse amohelang li fetoha ka lehlakoreng la ho fokotsa tsebo ho "moamoheli" oa tsona, joale ho hlaha mekhoa e fapaneng ea pele le ea morao-rao e thibelang phapanyetsano ea liphatsa tsa lefutso (Dyakov le Lekomtseva, 1984).
Liphuputso tse 'maloa li hlahlobile tšusumetso ea mohloli oa mathata a batsoali ka katleho ea hybridization. Ha ho tšela mefuta e arohaneng le mefuta e fapaneng ea mofuta oa Solanum naheng ea Ecuador (Oliva et al., 2002), ho ile ha fumanoa hore mefuta e nang le mofuta oa A2 ho tsoa ho Solanaceae e hlaha (clonal line EC-2) e tšela hampe haholo ka mefuta ea tamati. line EC -3), 'me ka ho fetisisa ka katleho e tšela le mofuta oa litapole (EC-1).
Li-hybrids tsohle li ile tsa fetoha tse se nang pathogenic. Bangoli ba lumela hore liperesente tse tlase tsa hybridization le phokotso ea pathogenicity ho li-hybrids li bakoa ke mekhoa ea morao-rao ea ho ikarola hoa batho.
Litekong tsa Bagirova et al (1998), mefuta e mengata ea litapole le tamati e nang le thepa ea merabe ea T0 le T1 e ile ea tšela. Tse nonneng haholo e ne e le lifapano tsa mefuta ea T1xT1 e arohaneng le tamati (36 oospores sebakeng sa pono ea microscope, 44% ea limela tsa oospore), tse neng li sa sebetse hantle e ne e le lifapano tsa merabe ea T0xT1 e arohaneng le mabotho a fapaneng (palo e tlase ea ho theha le ho mela. li-oospores, karolo e phahameng ea li-oospores tse ntšang mpa le tse sa tsoelang pele). Ts'ebetso ea lifapano lipakeng tsa mabelo a T0 a arohaneng le litapole e bile a mahareng. Kaha mefuta e mengata ea peiso ea T0 e tšoaetsa litapole, e na le mohloli o ka tšeptjoang oa ho feta nako ea mariha - li-tubers tsa litapole, ka lebaka leo bohlokoa ba li-oospores e le likarolo tse tšoaetsanoang tse fetang mariha bakeng sa baahi ba litapole li tlase. "Sebopeho sa tamati" se fetotsoeng se khona ho feta mariha ka tomate ka mokhoa oa oospores (sheba ka tlase) 'me ka hona ho boloka tlhahiso e phahameng ea ts'ebetso ea thobalano. Ka lebaka la tsoalo e phahameng, T1 e fumana monyetla o ikemetseng oa tšoaetso ea mantlha ho tamati. Liphetho tse fumanoeng ke Knapova et al. (2002) li ka hlalosoa ka tsela e tšoanang. Lifapano tsa mefuta e arohaneng le litapole tse nang le mefuta e tsoang ho langa le le lej li fane ka palo e phahameng ka ho fetisisa ea oospores - 13,8 ka sq. tikoloho (e nang le mefuta e mengata ea 5-19) le peresente e mahareng ea ho mela ha oospore (6,3 e nang le mefuta e mengata ea 0-24). Lifapano tsa mefuta e arohaneng le tamati li hlahisitse liperesente tse tlase haholo tsa oospores (7,6 e nang le mefuta e fapaneng ea 4-12) e nang le liperesente tse phahameng ka ho fetisisa tsa ho mela (10,8). Lifapano lipakeng tsa mefuta e arohaneng le litapole li fane ka palo e mahareng ea li-oospores (8,6 e nang le data e ngata - 0-30) le liperesente tse tlase haholo tsa ho mela ha oospore (2,7). Ka hona, mefuta ea litapole ha e nonne haholo ho feta mefuta ea tamati, empa lifapano tsa popopulation ha lia hlahisa litholoana tse mpe ho feta lifapano tsa intrapopulation. Mohlomong liphapang le lintlha tse ka holimo ke Bagirova et al. li hlalosoa ke taba ea hore bafuputsi ba Marussia ba sebelitse ka mefuta e arohaneng mathoasong a lilemo tsa bo-90 tsa lekholo la mashome a mabeli la lilemo, 'me bafuputsi ba Switzerland ba sebelitse ka mefuta e arohaneng ho elella bofelong ba lilemo tsa bo-90.
Tsoalo e tlase e ka ipapisa le mefuta ea heteroploidy. Haeba ho baahi ba Mexico, moo ts'oaetso ea thobalano le tšoaetso ea mantlha e nang le bana ba oosporic e tloaelehileng, boholo ba mefuta e ithutoang ea P. infestans ke diploid, joale linaheng tsa Old World intrapopulation polymorphism ea ploidy e hlokomeloa (di-, tri- le mefuta ea tetraploid, hammoho le mefuta ea heterokaryotic e nang le li-nuclei tsa heteroploid) , le mathata a nang le mefuta e sa tšoaneng ea ho nyalana, i.e. e nonneng, e fapana ka ploidy ea nyutlelie (Therrien et al., 1989, 1990; Whittaker et al., 1992; Ritch le Daggett, 1995). The heteroploidy ea li-nuclei ho antheridia le oogonia e ka 'na ea e-ba sesosa sa tsoalo e tlaase.
Ha e le phapanyetsano ea nyutlelie pakeng tsa hyphae nakong ea anastomoses, sena se thibeloa ke ho se lumellane ha limela, tse arolang batho ba sexual ka li-clones tse ngata tse arohaneng le liphatsa tsa lefutso (Poedinok le Dyakov, 1987; Gorbunova et al., 1989; Anikina et al., 1997b).
7) Khokahano ea baahi. Lintlha tse ka holimo li bontša hore ho kopanya pakeng tsa "tapole" le "tamati" ea mefuta ea P. infestans hoa khoneha. Tšoaetso e ts'oanang ea batho ba fapaneng le eona e ka khoneha, le hoja ho fokotsehile ka mabifi.
Phuputso ea matšoao a baahi libakeng tse ka thoko ho tloha masimong a haufi a litapole le tamati ka 1993 e bonts'itse hore hoo e ka bang kotara ea likarolo tse arohaneng le makhasi a tamati li fetisitsoe ho tsoa tšimong ea litapole e haufi (Dolgova et al., 1997). Ha ho nahanoa, ho ka nahanoa hore phapang ea baahi ho lihlopha tse peli e tla eketseha 'me e lebise ho hlaha ha mefuta e khethehileng ea intraspecific (f.sp. potato le f.sp. tamati), haholo kaha oospores e ka bolokoa ka har'a maloanlahla a limela (Drenth). et al., 1995; Bagirova, Dyakov, 1998) le peo ea langa le le lej (Rubin et al., 2001). Ka lebaka leo, tamati hajoale e na le mohloli oa ntlafatso ea selemo ntle le li-tubers tsa litapole.
Leha ho le joalo, lintho tsohle li ile tsa etsahala ka tsela e fapaneng. Overwintering le oospores e ile ea lumella likokoana-hloko ho qoba mohato o moqotetsane ka ho fetisisa oa potoloho ea bophelo - sebaka sa monocyclic sa limela mobung, nakong eo thepa ea likokoana-hloko e fokotsehang, butle-butle e hlaphoheloa lehlabuleng la polycyclic.
Letlapa la 11. Maqhubu a liphatsa tsa lefutso tsa virulence bakeng sa mefuta e fapaneng ea litapole mefuteng ea P. infestans
naha | Год | Karolelano ea liphatsa tsa lefutso tsa virulence ka mefuta | mongoli | |
ho tloha litapole | ho tloha tamati | |||
France | 1995 | 4.4 | 3.3 | Leberton et al., 1999 |
1996 | 4.8 | 3.6 | Leberton, Andrivon, 1998 | |
Fora, Switzerland | 1996-97 | 6.8 | 2.9 | Knapova, Gisi, 2002 |
United States | 1989-94 | 5 | 4.8 | Goodwin et al., 1995 |
USA, Bophirimela Washington DC | 1996 | 4.6 | 5 | Dorrance et al., 1999 |
1997 | 6.3 | 3.5 | " | |
Ecuador | 1993-95 | 7.1 | 1.3 | Oyarzun et al., 1998 |
Iseraele | 1998 | 7 | 4.8 | Cohen, 2002 |
1999 | 6 | 5.7 | " | |
2000 | 6.7 | 6.1 | " | |
Russia, Moscow sebaka | 1993 | 8.9 | 6.7 | Smirnov, 1996 |
Russia, libaka tse fapaneng | 1995 | 9.4 | 8 | Kozlovskaya et al. |
1997 | 9.2 | 9.2 | " | |
2000 | 8.7 | 4.8 | " |
Li-zoosporangia tsa mathomo le li-zoospores tseo li-oospores li melang ka tsona li na le ts'ebetso e phahameng ea likokoana-hloko, haholo-holo haeba li-oospores li entsoe ka parthenogenetically tlas'a tšusumetso ea li-pheromone tse tsoang mofuteng o fapaneng oa ho nyalana. Ka hona, lintho tse tšoaetsanoang lipeong tsa tamati tse hōlileng ho tloha ho lipeo tse tšoaelitsoeng ke oospores li kotsi haholo bakeng sa tamati le litapole.
Liphetoho tsena li lebisitse tlhophisong e 'ngoe ea baahi, e hlahisitsoeng liphetohong tse latelang tse neng li le bohlokoa ho latela pono ea lefu la seoa:
- Lipeo tsa tamati tse tšoaelitsoeng li fetohile mohloli oa bohlokoa oa tšoaetso ea mantlha ea litapole (Filippov, Ivanyuk, puisano ea botho).
- Li-epiphytoties ka litapole li ile tsa qala ho bonoa ka Phuptjane, hoo e ka bang khoeli pele ho tloaelo.
- Ho lema litapole, peresente ea peiso ea T1, eo pele e neng e fumanoa moo ka bongata bo fokolang, e eketsehile (Ulanova et al., 2003).
- Likokoana-hloko tse arohaneng le makhasi a tamati ha li sa fapana le mefuta ea litapole tse nang le likokoana-hloko tse khethollang litapole tsa liphatsa tsa lefutso tsa likokoana-hloko 'me tsa qala ho feta mefuta ea "litapole" ka mabifi eseng ka tamati feela, empa le litapole (Lavrova et al., 2003; Ulanova et al. , 2003).
Ka hona, ho e-na le ho fapana, ho ile ha etsahala kopano ea baahi, ka ho hlaha ha palo e le 'ngoe ea limela tse peli tse nang le likokoana-hloko tse nang le likokoana-hloko tse phahameng le tse mabifi ho mefuta ka bobeli.
fihlela qeto e
Kahoo, ho sa tsotellehe lilemo tse fetang 150 tsa boithuto bo matla ba P. infestans, ho ntse ho sa tsejoe ho hongata ho baeloji, ho kenyelletsa le baeloji ea baahi ba sesosa sena sa mafu a bohlokoa ka ho fetisisa a limela tse lenngoeng tsa nightshade. Ha ho hlake hore na ho fetela ha mekhahlelo e le 'ngoe ea potoloho ea bophelo ho ama sebopeho sa baahi, ke mekhoa efe ea liphatsa tsa lefutso tsa canalized variability ea aggressiveness le virulence, kamano ke efe pakeng tsa mekhoa ea ho ikatisa ea thobalano le clonal ho batho ba tlhaho, ho se lumellane ha limela joang? e futsitsoe, ke karolo efe ea litapole le tamati tšoaetsong e ka sehloohong ea lijalo tsena le ho seo e leng tšusumetso ea bona ho sebopeho sa baahi ba likokoana-hloko. Litaba tse bohlokoa bakeng sa mesebetsi e sebetsang, joalo ka mekhoa ea liphatsa tsa lefutso tsa liphetoho tse mabifi tsa likokoana-hloko kapa khoholeho ea ho hanyetsa ka mokhoa o sa tobang oa litapole, ha li e-s'o rarolloe. Ha liphuputso tse mabapi le lefu la morao la litapole li ntse li teba le ho hola, kokoana-hloko ena e hlahisa mathata a macha ho bafuputsi. Leha ho le joalo, ntlafatso ea bokhoni ba liteko le ho hlaha ha mekhoa e mecha ea mekhoa ea ho laola liphatsa tsa lefutso le liprotheine li re lumella ho ba le tšepo ea ho fumana tharollo e atlehileng litabeng tse hlahisitsoeng.
Sengoloa se phatlalalitsoe koranteng ea "Ts'ireletso ea Litapole" (No. 3, 2017)