S.N. Elansky, L.Yu. Kokaeva, N.V. Statsyuk, Yu.T. Dyakov
Nhanganyaya
Oomycete Phytophthora infestans (Mont.) De Bary - chinokonzera caxative yekunonoka kunaya, chirwere chakakosha zvakanyanya mune zvoupfumi matapiri nematomatisi - chakwezva kutarisisa kwevatsvagiri vanobva kunyika dzakasiyana kwenguva inopfuura zana nehafu. Ichangoerekana yaonekwa muEurope pakati pezana ramakore rechiXNUMX, yakakonzera chirwere chembatatisi chakaramba chiri mundangariro dzezvizvarwa zvakawanda.
Kusvika ikozvino, inowanzozodaidzwa kuti "howa hwenzara yeIreland". Anenge makore zana mushure mekupararira kwekutanga kwezvirwere, mbatatisi dzemusango dzeMexico dzinopokana nehusiku hwekupedzisira dzakawanikwa, nzira dzekuyambuka nembatatisi dzakarimwa dzakagadzirwa (Muller, 1935), uye mhando dzekutanga dzisingadziviriri blight dzakawanikwa (Pushkarev, 1937) . Nekudaro, chinguva chidiki mushure mekutanga kwekurima kwavo kwekutengesa, marudzi eanononoka blight pathogen, ane huturu kumarudzi asingagadzirike, akaunganidzwa. uye matsva ekurwisa majini akaunzwa mumhando dzakasiyana kubva kumatapiri emusango eMexico akatanga kurasika nekukurumidza kushanda kwawo.
Kukundikana nekushandisa monogenic (yakamira) kuramba kumanikidza vafudzi kutsvaga nzira dzakaomarara dzekushandisa nonspecific polygenic (yakatwasuka) kuramba. Mumakore achangopfuura, mune vamwe vanhu vehutachiona, nhangemutange ine hukasha yakatanga kuunganidzwa, ichikonzera kukuvara kwekusada kunyatsogadzikana. Kuuya kwehutachiona husingaurayiwe nefungicide kwakonzera matambudziko mukushandiswa kwemakemikari ekudzivirira mbatatisi.
Nekuda kwekusiyana kwakakura pakati peoomycetes uye fungi mune zvemakemikari, ultrastructure uye metabolism, fungicides, kunyanya systemic inoshandiswa kuchengetedza zvirimwa kubva kune dzakawanda fungal hosha, haina basa kurwisa oomycetes.
Naizvozvo, mukuchengetedza kwemakemikari kubva kunonoka kunonoka, kuwanda (kusvika pagumi nemaviri pamwaka kana kupfuura) kupfapfaidza pamwe nekugadzirira kwekubata kwakawanda kwekuita kwakashandiswa. Nhanho yekuchinja yaive kushandiswa kwe phenylamides, iyo ine chepfu kune oomycetes uye inopararira zvine hurongwa muzvirimwa. Nekudaro, kushandiswa kwavo kwakapararira nekukurumidza kwakatungamira mukuunganidzwa kwehutachiona husinga tsanangurike muhuwandu hwefungus (Davidse et al., 12), iyo yakanyanya kuomesa dziviriro yemiti. P. infestans ndiyo chete hutachiona hwenzvimbo yakadzikama, kukuvara uko kurima kwehupenyu hakugone kudzikiswa pasina kushandisa nzira yemakemikari yekudzivirira (Van Bruggen, 1981).
Izvo zviri pamusoro zvinotsanangura kutarisisa kukuru kwakabhadharwa nevatsvagiri vanobva kunyika dzakasiyana siyana kuenda kunzverwa kweP infestans vanhu, simba rekuwanda kwavo uye kuumbwa kwemajini, pamwe nemagene ekusiyana.
Hupenyu kutenderera kwaR. INFESTANS
Oomycete Phytophthora infestans inogadzira intercellular mycelium ine haustoria mukati memashizha emazamu. Kudya pamashizha emashizha, kunokonzeresa kuumbwa kwemavara erima, ayo anoshanduka kuita matema uye kuora mune inonaya mamiriro ekunze. Nekukundwa kwakakomba, shizha rose rinofa. Mushure menguva yekupa chikafu, mabukira anoumbwa pane mycelium - sporangiophores - ayo anokura kunze kuburikidza nespata. Mumamiriro ekunze akanyorova, vanoumba maruva machena akakomberedza iwo mavara pazasi pemashizha. Panopera sporangiophores, zvakaumbwa semonmon zoosporangia, izvo zvinotsemuka uye zvinotakurwa nekusasa mvura (Fig. 1). Kupinda mumadonhwe emvura pamusoro peshizha rembatatisi, sporangia inomera ne 6-8 zoospores, iyo, mushure menguva yekufamba, yakatenderedzwa, yakafukidzwa negobvu uye ichimera nehubhu rinomera. Bukira rinopinda munyama yemashizha kuburikidza ne stomata. Pasi pemamwe mamiriro, sporangia inogona kukura mune yekukura chubhu yakanangana nemashizha emashizha. Mumamiriro ezvinhu akanaka, iyo nguva kubva kuhutachiona kusvika pakuumbwa kweiyo sporulation ingori mazuva 3-4 chete.
Kamwe pane pasi uye kusefa kuburikidza nevhu, sporangia anokwanisa kutapura tubers. Yakanyanya kukanganisa tubers inoora panguva yekuchengetedza; mune isina kusimba yakakanganiswa, hutachiona hunogona kuramba huripo kusvika mwaka unotevera. Uye zvakare, iyo inokonzeresa yekumhanyisa blight inogona kuenderera munguva yechando muchimiro che oospores (gobvu-rakakomberedzwa kuzorora pabonde spores) muvhu pane zvinomera zvezvirimwa uye pamadomasi mbeu. Oospores inoumbwa pane mhenyu mitezo yemiti kana marudzi emhando dzakasiyana dzekusangana achisangana nehunyoro hwakanyanya Muchirimo, asexual sporulation inoumbwa pane yakasimwa hutachiona hutachiona uye pamasara emiti ane oospores, zoospores inopinda muvhu uye inokonzeresa hutachiona hwemashizha ezasi emiti. Mune zvimwe zviitiko, iyo mycelium inogona kukura kubva kune hutachiona tuber padivi pegirini chikamu chemuti uye kazhinji inoonekwa muchikamu chepamusoro chehunde.
Musiyano wakakura pakati pe oomycetes uye mazhinji mafungi ari mune yakanyanya kudiplophase muhupenyu hwavo kutenderera ne gametic meiosis uye kumera kwe zygotes (oospores) isina kudzikisira yenyukireya fission. Ichi chimiro, pamwe nedipolar heterotallism, kutsiva bisexuality, zvinoita kunge zvinoita kuti zvikwanise kunyorera kune oomycetes nzira dzakagadziridzwa dzekufunda vanhu vepamusoro eukaryotes (kuongororwa kwepanmixia uye kupatsanurwa kwevanhu, mukati-uye kupindirana kwemajini kuyerera, nezvimwewo). Nekudaro, zvinhu zvitatu hazvibvumidze kuendesa zvizere nzira idzi kuchidzidzo cheP infestans vanhu.
1. Pamwe pamwe nehoribhurosi oospores, inozvitakura uye parthenogenetic oospores inoumbwa muhuwandu hwevanhu (Fife naShaw, 1992; Anikina et al., 1997a; Savenkova, Cherepnikoba-Anirina, 2002; Smirnov, 2003), uye kuwanda kwechimiro chavo inogona kunge yakaringana kukanganisa pamhedzisiro yekuedzwa.
2. Kuita bonde mu P. infestans kunoita mupiro usingakoshe kune simba rehukuru hwehuwandu, nekuti fungus inobereka zvakanyanya neanomera zvipuka, ichigadzira inopfuura 90% yemhedzisiro yeongororo yemhando yekukwirisa nenzira yechinyakare. pane svikiro svikiro ... mwaka wekukura muzvizvarwa zvinoverengeka zveasexual sporulation (polycyclic chirwere kukura). Oospores inoita basa rakakosha mukuchengetedza kweiyo muviri panguva iyo zvirimwa zvakasvibirira zvisipo (munguva yechando) uye mune hutachiona hwekutanga hwembeu. Zvino, mukati mezhizha, kubereka kwekroniki kunoitika uye kuwedzera kana, nekupesana, kuderera kwenhamba yemakonzo ega ega anomuka semhedzisiro yekurara pabonde, izvo zvinonyanya kutemerwa nekusarudzwa kweakanyanya kugadziriswa. Naizvozvo, iro reshiyo yemunhu clones muhuwandu pakutanga uye kumagumo epiphytotic inogona kuve yakasiyana zvachose.
3. Iyo yakatsanangurwa kutenderera inowanzoitika kune veruzhinji vagari veP. Infestans munyika yokumusha kwavo, Central America. Mune dzimwe nzvimbo dzepasi, kuita bonde kwaisazivikanwa kweanopfuura makore zana; iyo yekudyara mycelium muhutachiona hwembatatari tubers yaive nhanho yechando. Hupenyu hwekutenderera hwakanga hwazara zvachose, uye kupararira kwaive kwakatarisana nemasikirwo: hutachiona kubva kune imwechete hutachiona hwakasimwa tubers hwakapfuudzwa kumashizha, ichiumba nzvimbo yekutanga yechirwere, iyo inogona kusangana pamwe nekukura kukuru kwechirwere.
Nekudaro, mune mamwe matunhu panogona kuve nekuchinjika kwepabonde uye kweasexual macircuit, nepo kune vamwe - chete asexual kutenderera.
Kwakatangira P. INFESTANS
P. infestans akaonekwa muEurope pakupera kwehafu yekutanga yezana ramakore rechi1991. Sezvo mbatatisi inozvarwa kuchamhembe kwakadziva kumabvazuva kwechikamu cheSouth America, zvaifungidzirwa kuti utachiona hwakaunzwa kubva imomo kuenda kuEurope panguva yekukura kweChilean saltpeter. Zvisinei, zvidzidzo zvakaitwa paRockefeller Center chiteshi chembatatisi muToluca Valley, Mexico zvakaita kuti pfungwa iyi iongororwe (Niederhauser, 1993, XNUMX).
1. MuMupata weToluca, zvipuka zvembatatari zvemuno (Solanum demissum, S. bulbocastanum, nezvimwewo) zvine mageneti akasiyana emajeti ekumira akatwasanuka pamwe nehupamhi hwekupokana husingaenzaniswi, izvo zvinoratidza kubatana-kureba pamwe nehutachiona. MaSouth America marudzi, kusanganisira mbesa mbatatisi, kushomeka kwemajini.
2. MuMupata weToluca, unosarudzika nemhando dzekubatana A1 neA2 dzinowanikwa, semhedzisiro iyo huwandu hwevanhu vakapatsanurwa hweP infestans hwakapararira; uchiri munyika yekumusha yematata akarimwa, muSouth America, hutachiona hunopararira clonally.
3. MuMupata weToluca, mune hosha dzakaipisisa dzegore negore dzekunonoka kunaya. Naizvozvo, pakati pevaongorori vekuNorth America (Cornell University), iwo maonero nezveMesoamerica (Central America) senzvimbo yekuberekerwa mbatatisi phytophthora inosimbiswa (Goodwin et al., 1994).
Vatsvakurudzi vekuSouth America havagovani pfungwa iyi. Vanotenda kuti mbatatisi yakasimwa nehutachiona hwayo P. infestans vane nyika yakajairika - maSouth America Andes. Vakatsigira pfungwa yavo nemaonero emakemikari pane kuongororwa kweDNA polymorphisms ye mitochondrial genome (mtDNA) uye ma nyukireya majini RAS uye β-tubulin (Gomez-Alpizar et al., 2007). Vakaratidza kuti maronda akaunganidzwa kubva kumatunhu akasiyana epasi akaburuka kubva kumitsara mitatu yakasiyana yemadzitateguru ayo (ese matatu) anowanikwa muSouth America Andes. Andean haplotypes zvizvarwa zvemitsara miviri: yakasarudzika yeiyo yekare mtDNA dzinza inowanikwa pasango Solanaceae kubva kuAnarrhicomenum chikamu muEcuador, nepo kusarudzika kwemutsara wechipiri kwakajairika pamatatata, matomatisi, uye nightshades yemusango. MuToluca, nyangwe zvisingawanzo haplotypes zvakaburitswa kubva mumutsara mumwe chete, nekusiyana kwemajini kweToluca strains (yakaderera allelic frequency yemamwe masayiti akachinjika) inoratidza yakasimba muvambi mhedzisiro nekuda kwekufamba kwenguva.
Uye zvakare, imwe mhando nyowani P. andina yakawanikwa muAndes, morphologically uye genetiki yakafanana neP infestans, iyo, sekureva kwevanyori, inonongedzera kuAndes senzvimbo inopisa yekufungidzira mune genus Phytophthora. Chekupedzisira, muEurope neUnited States, P. Infestans huwandu hunosanganisira ese maAndean mitsara, nepo muToluca mumwe chete.
Chinyorwa ichi chakakonzera kumukira kubva kuboka revatsvakurudzi vanobva kunyika dzakasiyana, avo vakaita basa rakawanda rekuyedza kudzokorora chidzidzo chekare (Goss et al., 2014). Mune iri basa, chekutanga, yakawanda inodzidzisa microsatellite DNA kuteedzana kwakashandiswa kudzidza DNA polymorphisms; kechipiri, kuongororwa kwekubatanidzwa, nzira dzekutama, nguva dzakasiyana dzevanhu, nezvimwe. mamwe mamodheru epamberi akashandiswa (F-manhamba, fungidziro dzeBazesian, nezvimwewo) uye, chechitatu, fananidzo yaishandiswa kwete chete nemhando dzeAndean P. andina, umo maakasikwa hunhu (P. infestans x Phytophthora sp.) , asiwo nemhando dzekugara dzeMexico P. mirabilis, P. Ipomoeae, uye Phytophthora phaseoli - genetically close P. infestans inowanikwa mune imwechete clade (Kroon et al., 2012). Nekuda kweichi ongororo, zvakaratidzwa pachena kuti mudzi wemuti wephylogenetic wemarudzi ese erudzi rwePhytophthora wakatorwa muchidzidzo ichi, kunze kweiyo hybrid P. andina, ndewemhando dzeMexico, uye kuyerera kwekutama kune kutungamira Mexico - Andes, uye kwete zvinopesana, uye kutanga kwayo kunoenderana neEuropean colonization yeNyika Idzva (makore 300-600 apfuura). Nekudaro, kubuda kweP infestans marudzi, akasarudzika ekukundwa kwembatatisi, kwakaitika mune yechipiri genetiki nzvimbo yekuumbwa kwetuberous solanaceae, i.e. muCentral America.
Genome yaP. INFESTANS
Muna 2009, timu yepasi rese yemasayendisiti yakateedzera yakazara P infestans genome (Haas et al, 2009), saizi yaive 240 MB. Izvi zvakapetwa kakati wandei kupfuura mumhando dzakabatana dzepedyo P. sojae (95 Mb), izvo zvinokonzeresa kuora kwesoya, uye P. Ramorum (65 Mb), zvichikanganisa mhando dzakakosha dzemiti senge oak, beech uye mimwewo. Iyo data yakawanikwa yakaratidza kuti iyo genome ine nhamba hombe yemakopi akateedzana akateedzana - 74%. Iyo genome ine 17797 protein-coding genes, mazhinji acho ari mageneti anobatanidzwa muma cellular process, kusanganisira DNA kudzokorora, kunyora uye kushandura kwemapuroteni.
Kufananidzwa kwemajenome eiyo genus Phytophthora kwakaratidza kusarongeka kwesangano rejenome, riine mabhureki akateedzana emageneti akachengeterwa umo kuwanda kwemajini kuri kwakanyanya uye zvirimo zvekuteedzana zvakaderera, uye matunhu ega ega asina-akachengetedzwa geni. akateedzana, aine yakaderera geni kuwanda uye yakakwira zvemukati yekudzokorora matunhu. Conservative inovhara account ye70% (12440) yeese P. infestans protein-coding majini. Mukati memabhuroko akachengetedzwa, majini anowanzo kuve akapatsanurwa aine avhareji yepakati peiyo intergenic ye604 bp. Munzvimbo dziri pakati pemabhuroko anochengetedza, iyo intergenic daro yakakura (3700 bp) nekuda kwekuwedzera kwehuwandu hwezvinhu zvinodzokorora. Inokurumidza kuchinja inogadzira epachivande majini ari munzvimbo dzemhando-dzisina hurombo.
Kuongororwa kweiyo P. Infestans genome kuteedzana kwakaratidza kuti chinenge chikamu chimwe muzvitatu chejenome ndechezve zvinhu zvinochinja. Iyo genome yaP. Infestans ine yakanyanya kuwanda mhuri dzakasiyana dzevashanduri kupfuura mamwe magenome anozivikanwa. Mazhinji eP.Pestestans transposons ndewemhuri yechiGypsy.
Huwandu hukuru hwemhuri dzedzinza dzakabatanidzwa mupathogenesis dzakaratidzwa mu P. infestans genome. Chikamu chakakosha chazvo chinogadzira mapuroteni anoshandura ayo anoshandura mafizikisi echirimwa chinogashira uye nekupa hutachiona. Dzinowira mumapoka maviri akapamhamha: apoplastic maactor, ayo anoshanda munzvimbo dzecellular (apoplasts), uye cytoplasmic maactor, ayo anopinda maseru kuburikidza nehaustoria. Apoplastic maactor anosanganisira akavanzika hydrolytic enzymes senge mapuroteni, lipases uye glycosylases anoparadza masero emiti; inhibitors yeanogamuchira miti yekudzivirira enzyme; uye necrotizing chepfu yakadai seNep1-senge mapuroteni (NPLs) uye Pcf-senge diki cysteine-akapfuma mapuroteni (SCRs).
P. infestans inogadzira majini akawandisa uye kazhinji akakura pane asiri-pathogenic majini. Iwo anonyanya kuzivikanwa iwo ma cytoplasmic maactor RXLR uye Crinkler (CNR). Iwo chaiwo cytoplasmic maakisi e oomycetes ari RXLR mapuroteni. Ese maRXLR anogadzira mageneti akawanikwa kusvika parizvino ane amino-terminal boka Arg-XLeu-Arg, uko X iri amino acid. Nekuda kwekudzidza, zvakataurwa kuti pane mazana mashanu nemakumi matatu nematatu emaRXLR maP. Infestans genome, inova 563% kudarika muPojaja naP P. ramorum. Inenge hafu yemageneti eRXLR muP infestans genome ndiwo akasarudzika. RXLR maitirwo ane akasiyana siyana akateedzana. Pakati pavo, imwe mhuri hombe uye zana nemakumi mashanu akaonekwa Kusiyana neproteome yakakosha, iyo maRXLR anoshanda magene anowanzo kuwanikwa munzvimbo dzisina hurombo uye dzichidzokorodza hupfumi hwejenome. Iwo mafoni emahara anoona kushanduka kwenzvimbo idzi anosimudzira kuumbazve mune aya magene.
MaCytoplasmic CRN maactor akaonekwa pakutanga muP. Infestans zvinyorwa zvekukodha chirimwa chemukati necrosis peptides. Kubva zvavakawana, zvishoma zvave kuzivikanwa nezvemhuri yeava vanoita. Kuongororwa kwe P. Infestans genome kwakaburitsa mhuri hombe ye196 magene eCRN, ayo akakura zvakanyanya kupfuura muPojae (100 CRN) uye P. ramorum (19 CRN). SaRXLRs, maCRN ari modular mapuroteni uye ane anochengetedzwa zvakanyanya N-terminal LFLAK dura (50 amino acids) uye padyo neDWL domain ine akasiyana magene. Mazhinji maCRN (60%) ane chiratidzo pepeptidi.
Iko mukana weakasiyana maCRNs ekuvhiringidza maseru maitiro eiyo inomiririra chirimwa yakadzidzwa. Mune ongororo yemiti necrosis, kubviswa kweCRN2 mapuroteni kwakaita kuti zvikwanisike kuziva iyo C-terminal dunhu ine 234 amino acids (zvinzvimbo 173-407, DXG domain) uye zvichikonzera kufa kwesero. Kuongororwa kwemajini eCNN eP. Infestans kwakaratidza mana akasiyana maC-terminal matunhu, ayo anokonzeresa kufa kwemasero mukati memuti. Izvi zvinosanganisira madomaini eDC achangobva kuzivikanwa (P. Infestans ane mageneti gumi nemasere uye makumi mana nemapfumbamwe echirwere), pamwe neD18 (49 ne2) uye DBF (14 ne43) madomeni akafanana nemaprotein kinases. Mapuroteni eCRN domains anoratidzwa mune chirimwa anochengetedzwa (mukushayikwa kwemasaini peptides) mune chirimwa chemusero uye kukurudzira sero kufa neyakavanzika masisitimu. Imwe mitezo makumi maviri neshanu ine madomeni eCRN anogona kunge asingashande semajini.
Iko kuwedzera kwenhamba uye saizi yeiyo RXLR uye CRN inogadzira geni mhuri zvaifungidzirwa nekuda kweasina-allelic homologous kudzokorora uye kudzokorora kwemajini. Kunyangwe hazvo chokwadi chekuti genome ine huwandu hwakawanda hweanoshanda mafoni mafoni, hapachina humbowo chaihwo hwekuchinjiswa kweanogadzira magene.
Maitiro anoshandiswa mukudzidza kwehuwandu hwevanhu
Iko kudzidza kweiyo genetic chimiro chevanhu parizvino kwakavakirwa pakuongorora kwetsika dzakachena dzematunhu aro. Kuongororwa kwevanhu pasina kusiyanisa tsika dzakachena kunoitiswawo nekuda kwezvakanangana, senge, semuenzaniso, kudzidza hukasha hwevagari kana kuvapo kwematambudziko anopikisa fungicides mairi (Filippov et al., 2004; Derevyagina et al., 1999). Rudzi urwu rwekutsvagisa runosanganisira kushandiswa kwenzira dzakasarudzika, tsananguro yacho iri pamusoro pechiyero chekuongorora uku. Pakuenzanisa kwekuenzanisira kwematambudziko, nzira dzinoverengeka dzinoshandiswa, zvichibva pane kuongororwa kwechimiro cheDNA uye pakudzidza kwe phenotypic kuratidzwa. Kuenzanisa kuongorora kwevagari kunofanirwa kubata nenhamba yakakura yevasungwa, izvo zvinomisikidza zvimwe zvinodiwa panzira dzinoshandiswa. Sezvineiwo, vanofanirwa kuzadzisa zvinotevera zvinodiwa (Cooke, Lees, 2004, Mueller, Wolfenbarger, 1999):
- ive yakachipa, iri nyore kuitisa, haidi nguva yakakosha yemari, zvinoenderana neanowanikwa matekinoroji (semuenzaniso, PCR);
- inofanirwa kuburitsa huwandu hwakaringana hwakawanda hweakazvimirira macodomant marker maficha;
- vane reproducibility yakakwirira;
- shandisa hushoma huwandu hwetissue ichaongororwa
- ive yakanangana neiyo substrate (kusvibiswa kuripo mutsika hakufanirwe kukanganisa mhedzisiro);
- haidi kushandiswa kwemaitiro ane njodzi uye makemikari ane chepfu zvakanyanya.
Nehurombo, hapana nzira dzinoenderana neese ari pamusoro parameter. Zvekuenzanisa kwekutsvagisa kwematambudziko munguva yedu, nzira dzinoshandiswa zvichibva pakuongorora kwe phenotypic hunhu: huturu hwembatatisi uye madomasi marudzi (mbatatisi uye madomasi matomati), mhando yekukwikwidza, spectra ye peptidase isoenzymes uye glucose-6-phosphate isomerase, uye pane ongororo yeDNA mamiriro: kureba polymorphism kudzora chidimbu (RFLP), icho chinowanzo kuwedzerwa pamwe neiyo hybridization probe RG 57, kuongororwa kwemicrosatellite inodzokorora (SSR neInterSSR), kukwidziridzwa neyakajairika primers (RAPD), kukwidziridzwa kwezvimisikidzwa zvidimbu (AFLP) , kukwidziridzwa neyekutanga homologous kune akateerana enharembozha (semuenzaniso, Inter SINE PCR), kutsunga kwemitochondrial DNA haplotypes.
Rondedzero pfupi yemaitiro ekuenzanisa kwekuenzanisira kwematambudziko anoshandiswa mukushanda naP P. Infestans
Phenotypic mucherechedzo hunhu
"Mbatatisi" marudzi
"Mbatatisi" nhangemutange inowanzo kuongororwa uye kushandiswa chiratidzo. "Nyore mbatatisi" madzinza ane rimwe geni rembatatisi virulence, "yakaoma" - angangoita maviri. Black et al. (1953), muchidimbu data ravo rese, vakawana kuti iyo phytophthora nhangemutange inokwanisa kutapurira zvirimwa neinodzivirira geni / majini anoenderana neP infestans virulence gene / genes, uye akawana madzinza 1, 2, 3, uye 4 iyo inobata zvirimwa nemajini R1, R2, R3 uye R4, zvichiteerana, i.e. kudyidzana pakati pehutachiona nehutachiona kunoitika pane geni-ne-gene hwaro. Kupfuurirazve, Nhema, nekutora chikamu kwaGallegly naMalcolmson, vakawana ekuzvidzivirira majini R5, R6, R7, R8, R9, R10 uye R11, pamwe chete nemakwikwi anoenderana (Nhema, 1954; Nhema & Gallegly, 1957; Malcolmson & Nhema , 1966; Malcolmson, 1970).
Iko kune yakazara muviri wedata pane erudzi kuumbwa kwepathogen kubva kumatunhu akasiyana. Pasina kuongorora data iri zvakadzama, isu tinongoratidzira maitiro akajairika: uko kwakashandiswa mhando dzakasiyana nemajini ekurwisa kana musanganiswa wadzo, pakutanga pakanga paine kushomeka kwekupedzisira kwekutsvaira, asi ipapo madzinza ane anowirirana nehutachiona majeni akaonekwa uye akasarudzwa uye akaputika yekunonoka blight yakatanga zvakare. Hupombwe chaihwo hunopikisana neyekutanga hutachiona hwekudzivirira (R4-R1) hwakanga husingawanzo kucherechedzwa mumatanho akaunganidzwa isati yatanga kurimwa kwemhando dzakasiyana nemajini aya, asi huwandu hwehutachiona hunowedzera zvakanyanya apo hutachiona hwakagumburwa pamhando dzakatakura majini aya. MaGenesi 4-5, kune rimwe divi, aive akajairika mukuunganidzwa (Shaw, 11).
Kuongororwa kwehuwandu hwemarudzi akasiyana siyana mukati mekukura mwaka, kwakaitwa mukunopera kwema1980, kwakaratidza kuti pakutanga kwechirwere ichi, clones ine hukasha hwakanyanya uye 1-2 hutachiona hutachiona huzhinji muhuwandu.
Kupfuurirazve, sezvo blight yakanonoka ichikura, kuwanda kwema clones epakutanga kunodzikira uye huwandu hwemadzinza "akaomarara" ane hukasha hwakanyanya hunowedzera. Kuitika kwekupedzisira kunosvika zana muzana pakupera kwemwaka. Kana uchichengeta machubhu, pane kuderera kwehasha uye kurasikirwa kweumwe munhu hutachiona hutachiona. Simba rekutsiva kwemakoni rinogona kuitika mumhando dzakasiyana nenzira dzakasiyana (Rybakova & Dyakov, 100). Nekudaro, zvidzidzo zvedu muna 1990-2000 zvakaratidza kuti nhangemutange yakaoma inowanikwa kubva pakutanga kweepiphytotic pakati pematambudziko akaparadzaniswa nembatatisi nematomatisi. Izvi zvinogona kunge zvichikonzerwa neshanduko muhuwandu hwevane P. Infestans muRussia.
Na 1988-1995, kuitika kwe "superrace" neese kana anenge ese majini ehutachiona mumatunhu akasiyana akasvika 70-100%. Aya mamiriro akacherechedzwa, semuenzaniso, muBelarus, munzvimbo dzeLeningrad neMoscow, kuNorth Ossetia uye kuGerman (Ivanyuk et al., 2002a, 2002b; Polityko, 1994; Schober-Butin et al., 1995).
"Matomatisi" marudzi
Mumadomasi anorimwa, mageneti maviri chete ekurwisa kunaya kwakanonoka akawanikwa - Ph2 (Gallegly & Marvell, 1) uye Ph1955 (Al-Kherb, 2). Sezvazviri mumijaho yembatatisi, kudyidzana pakati pamadomasi naP P. infestans kunoitika pane geni-ne-gene hwaro. Rudzi rweT1988 runobata mhando dzisingave nemajini ekurwisa (mazhinji emhando dzinoshandiswa nemaindasitiri), iyo T0 nhangemutange inobata mhando neiyo Ph1 geni (Ottawa), uye mujaho weT1 unokanganisa mhando neiyo Ph2 geni.
MuRussia, ingangoita chete T0 yakawanikwa pamatatata; pamadomasi pakutanga kwemwaka, T0 yakakunda, asi gare gare yakatsiviwa neiyo T1 mujaho (Dyakov et al., 1975, 1994). Mushure ma2000, T1 pamatatata mune vakawanda vanhu yakatanga kuwanikwa pakutanga kweepiphytotic. MuUnited States, mbatatisi dzemhando dzemapatatisi dzaive dzisiri-pathogenic kumatomato, pamwe neT0, T1, uye T2 nhangemutange, nepo T1 neT2 zvakanyanyisa pamadomasi (Vartanian & Endo, 1985; Goodwin et al., 1995).
Kusangana mhando
Zvekudzidza, tester (chirevo) maronda ane anozivikanwa mating mhando - A1 uye A2 anodikanwa. Iyo bvunzo yekuzviparadzanisa yakaiswa jekiseni navo vari vaviri vaviri muPetri ndiro ine oat agar yepakati. Mushure mekumisidzana kwemazuva gumi, mahwendefa anoongororwa kuvepo kana kusavapo kwemaoospores mune yepakati munzvimbo yekubatana yematambudziko. Pane zvingasarudzwa zvina: kushushikana ndekwemhando yekukwikwidzana yeA10, kana ikaumba oospores neA4 tester, kuenda kuA1, kana ikaumba oospores neA2 tester, kuenda kuA2A1, kana ikaumba oospores ine vese testers, kana haina chena ( 1), kana ikasaumba oospores isina muedzo (mapoka maviri ekupedzisira haawanike).
Kuti ikurumidze kuona mhando dzekusvirana, kuyedza kwakaitwa kuti utarise matunhu ejenome anoenderana nemhando yekusvirana, nechinangwa chekushandisa kwavo zvekare kuona mhando yekusvirana nePCR. Imwe yekutanga yekubudirira kuyedza kutsvaga saiti yakadaro yakaitwa nevanoongorora veAmerica (Judelson et al., 1995). Vachishandisa nzira yeRAPD, ivo vakakwanisa kuziva iyo W16 dunhu inosanganisirwa nemhando yekukwikwidzana muvana vevaviri vakayambuka vari voga, uye dhizaina peya ye24-bp primers yekusimudzira kwayo (W16-1 (5'-AACACGCACAAGGCATATAAATGTA-3 ' ) uye W16-2 (5 '-GCGTAATGTAGCGTAACAGCTCTC-3') Mushure mekurambidzwa kwechigadzirwa chePCR nechirambidzo enzyme HaeIII, zvaigona kupatsanura kuzvipatsanura nemhando dzekubatanidza A1 neA2.
Kumwe kuyedza kwekutora mamaki ePDR ekuona mhando dzekubatana kwakaitwa nevanoongorora veKorea (Kim, Lee, 2002). Ivo vakaratidza zvimwe zvigadzirwa vachishandisa iyo AFLP nzira. Nekuda kweizvozvo, maviri emhando yekutanga PHYB-1 (kumberi) (5'-GATCGGATTAGTCAGACGAG-3 ') uye PHYB-2 (5'-GCGTCTGCAAGGCGCATTTT-3') yakagadziriswa, ichibvumira kusarudzika kwenzvimbo yegenome inosangana neA2 mhando yekusangana. Shure kwaizvozvo, vakaenderera mberi nebasa iri uye vakagadzira ekutanga 5 'AAGCTATACTGGGACAGGGT-3' (INF-1, kumberi) uye 5'-GCGTTCTTTCGTATTACCAC-3 '(INF-2), ichibvumira kusarudza kukwidziridzwa kwenzvimbo yeMat-A1 mamatambudziko ane mating mhando A1. Iko kushandiswa kwePCR kuongororwa kwemhando dzekuberekana kwakaratidza mhedzisiro yakanaka mukudzidza kweP infestans vanhu muCzech Republic (Mazakova et al., 2006), Tunisia (Jmour, Hamada, 2006) nedzimwe nzvimbo. MuLabhoritari yedu (Mytsa, Elansky, isina kuburitswa), 34 P. infestans masisitimu akasiyiwa kubva kurwara mbatatisi uye matomatisi matunhu munzvimbo dzakasiyana dzeRussia (Kostroma, Ryazan, Astrakhan, neMoscow matunhu) akaongororwa. Mhedzisiro yePCR yekuongorora ichishandisa chaiwo ma primers anopfuura makumi mapfumbamwe muzana akaenderana nemhedzisiro yekuongororwa kwerudzi rwekusvirana nenzira yechinyakare pane inosangana noutachiona.
Tafura 1. Kusiyana kwekushora mukati meSib 1 dombo (Elansky et al., 2001)
Semuenzaniso yekuunganidza nzvimbo | Huwandu hwevanosarudzika vakaongororwa | Nhamba yeyakaomarara (S), isina kusimba inodzivirira (SR) uye inodzivirira (R) matambudziko, pcs (%) | ||
S | SR | R | ||
G. Vladivostok | 10 | 1 (10) | 4 (40) | 5 (50) |
G. Chita | 5 | 0 | 0 | 5 (100) |
G. Irkutsk | 9 | 9 (100) | 0 | 0 |
G. Krasnoyarsk | 13 | 12 (92) | 1 (8) | 0 |
Yekaterinburg guta | 15 | 8 (53) | 1 (7) | 6 (40) |
O. Sakhalin | 66 | 0 | 0 | 66 (100) |
Dunhu reOmsk | 18 | 0 | 0 | 18 (100) |
Metalaxyl kuramba sehuwandu hwevanhu
Mukutanga kwema1980, kubuda kwakasimba kweshungu kwakanonoka kunokonzerwa nemetalaxyl-resistant P. infestans strains kwakacherechedzwa mumatunhu akasiyana siyana. Mapurazi embatatisi munyika zhinji akatambura kurasikirwa kukuru (Dowley & O'Sullivan, 1981; Davidse et al., 1983; Derevyagina, 1991). Kubva ipapo, munyika zhinji dzepasirese, kugara uchiongorora kuitika kwezve phenylamide-isingabatike masisitimu muP infestans vanhu vakaitwa. Pamusoro pekuongororwa kunoshanda kwetarisiro yekushandiswa kwemishonga ine phenylamide, kuvaka nzira yekudzivirira uye kufungidzira epiphytoties, kuramba kwemishonga iyi kwave chimwe chezvakaratidzwa zvinoshandiswa zvakanyanya pakuenzanisa kuongororwa kwehuwandu hweiyo pathogen. Zvisinei, kushandiswa kwekurwisa metalaxyl mukuenzanisira kwehuwandu hwevanhu kunofanirwa kuitiswa zvichitarisa chokwadi chekuti: 1 - iyo hwaro hwekuzvidzivirira haisati yanyatso tsanangurwa, 2 - kuramba metalaxyl hunhu hunosarudzika hunogona kuchinja. zvinoenderana nekushandiswa kwe phenylamides, 3 - dzakasiyana mwero wekunzwa kune metalaxyl masisitimu mukati meimwe clonal mutsetse (Tafura 1).
Isozyme spectra
Zviratidzo zveIsozyme zvinowanzozvimiririra nemamiriro ekunze, zvinoratidza nhaka yeMendelian uye zvine mukurumbira, zvichibvumidza kusiyanisa pakati pehomo- uye heterozygotes. Iko kushandiswa kwemapuroteni semagenera emakiki kunoita kuti zvikwanise kuona ese makuru marongero eiyo magene, kusanganisira chromosomal uye genomic mutation, uye imwechete amino acid kutsiva.
Zvidzidzo zveElectrophoretic zvemapuroteni zvakaratidza kuti ma enzyme mazhinji aripo mune zvipenyu nenzira yezvikamu zvinoverengeka zvakasiyana mukufamba kweelectrophoretic. Izvi zvidimbu mhedzisiro yekukodha akawanda mafomu enzyme nema loci akasiyana (isozymes kana isozymes) kana nema alleles akasiyana eiyo locus (allozymes kana alloenzymes). Ndokunge, isozymes marudzi akasiyana enzyme imwe. Mafomu akasiyana ane chiitiko chakafanana, asi anosiyana zvishoma mune imwechete amino acid kutsiva mu peptide uye anotarisira. Misiyano yakadaro inoratidzwa panguva yeelectrophoresis.
Mukudzidza kwe P. infestans strains, iwo masisitimu ezisoenzymes emapuroteni maviri, peptidase uye glucose-6-phosphate isomerase, anoshandiswa (iyi enzyme monomorphic muvanhu veRussia, saka, nzira dzekudzidza kwayo hadziuniswe mune iri basa) . Kuzvipatsanura kuva isozymes mune yemagetsi munda, gadziriro dzeprotein dzakasarudzika kubva kuzvinhu zvakafundwa dzinoiswa kune gel ndiro yakaiswa mumunda wemagetsi. Chiyero chekupararira kwemapuroteni ega ega mujeri chinoenderana nemutero uye uremu hwemaseru; Naizvozvo, mundima yemagetsi, musanganiswa wemapuroteni unopatsanurwa kuita zvikamu zvidiki, izvo zvinogona kutariswa uchishandisa madhayi akasarudzika.
Kudzidza kwepeptidase isoenzymes kunoitwa pane cellulose acetate, starch kana polyacrylamide gels. Iyo yakanyanya nyore ndiyo nzira inoenderana nekushandiswa kweserulose acetate gels akagadzirwa naHelena Laboratories Inc. Izvo hazvitsvake zviyero zvakakura zvezvinhu zvekuyedza, zvinobvumidza munhu kuti awane mabhendi akasiyana pajel mushure me electrophoresis kune ese enzyme loci, kuitiswa kwayo hakudi nguva yakakura uye mutengo wezvinhu (Fig. 2).
Chidimbu chemycelium chinotamisirwa mu 1,5 ml microtube, 1-2 madonhwe emvura yakasanganiswa yakawedzerwa pairi. Mushure meizvozvo, iyo sampuro inogadziriswa (semuenzaniso, ine chibooreso chemagetsi chine chinamatira chepurasitiki chakakodzera microtube) uye sedimented kwemasekondi makumi maviri neshanu pane centrifuge pa25 rpm. Kubva pane microtube yega yega, 13000 μl. iyo supernatant inoendeswa kune iyo yekushandisa ndiro.
Iyo cellulose acetate gel inobviswa mumudziyo wechidziro, yakadzimurwa pakati pemashizha maviri efirita pepa uye yakaiswa neinoshanda dura pamusoro peiyo yepurasitiki base yeiye anonyorera. Mhinduro kubva pahwendefa inoendeswa neanoshandisa kune iyo gel 2-4 nguva. Iyo gel inoendeswa kune ye electrophoresis kamuri,
Tafura 2. Kuumbwa kwemhinduro yakashandiswa kushatisa cellulose acetate gel mune ongororo ye peptidase isoenzymes, donhwe rependi (bromophenol bhuruu) inoiswa kumucheto kweiyo gel.
TRIS HCl, 0,05M, Ph 8,0 2 ml
Peroxidase, 1000 U / ml 5 madonhwe
o-dianisidine, 4 mg / ml 8 madonhwe
MgCl2, 20 mg / ml 2 madonhwe
Gly-Leu, 15 mg / ml 10 madonhwe
L-amino-acid oxidase, 20 u / ml 2 madonhwe
Electrophoresis inoitwa kwemaminitsi makumi maviri. pa20 V. Mushure me electrophoresis, iyo jeri inoendeswa patafura yekupenda uye yakasvibiswa neyakajeka pendi mhinduro (Tafura 200). 2 ml ye10% DIFCO agar inonyungudutswa muovheni yewa microwave, yatonhorera kusvika 1,6 ° C, mushure mezvo 60 ml yeagar inosanganiswa nemusanganiswa wependi uye ndokudururwa mujeri. Mitsetse inoonekwa mukati memaminetsi gumi nemashanu. Iyo L-amino-acid oxidase reagent inowedzerwa nguva pfupi usati wasanganisa mhinduro neyakaumbwa agar.
Mune vanhu veRussia, iyo Pep 1 locus inomiririrwa nemagenotypes 100/100 uye 92/100. Homozygote 92/92 inonyanya kushomeka (inenge 0,1%). Locus Rehr 2 inomiririrwa nematatu genotypes 100/100, 100/112, uye 112/112, uye ese matatu akasiyana akajairika (Elanky naSmirnov, 3, Fig. 2003).
Ongororo yeGenome
Kudzivisa chidimbu kureba kwepolymorphism pamwe neinotevera hybridization (RFLP-RG 57)
Yese DNA inorapwa neEco R1 yekudzivirira enzyme, zvidimbu zveDNA zvakapatsanurwa neagarose gel electrophoresis. Nyukireya DNA yakakura kwazvo uye iine kuwanda kwakateedzana; nekudaro, kunongedza kwakaringana kwezvimedu zvakawanda zvakawanikwa nekuita kwekudzivirira enzyme kunonetsa kuita. Naizvozvo, zvidimbu zveDNA zvakaparadzaniswa mujel zvinotamisirwa kune yakakosha membrane uye inoshandiswa kuomeswa neiyo RG 57 probe, iyo inosanganisira nucleotides yakanyorwa nemwaranzi inonzi radioactive kana fluorescent. Iyi probe inosanganisa nekudzokorora maitiro ejenomic (Goodwin et al., 1992, Forbes et al., 1998). Mushure mekuratidzwa kwemhedzisiro yehybrisization pane mwenje- kana radioactive zvinhu, yakawanda-locus hybridization profil (kudhinda kwemunwe) kunowanikwa, inomiririrwa nezvidimbu 25-29 (Forbes et al., 1998). Asexual (clonal) vana vanozove neakafanana ma profiles. Negadziriro yemabhanhire pane electrophoretogram, zvakafanana uye misiyano yezvisikwa zvakaenzaniswa zvinotongwa.
Mitochondrial DNA haplotypes
Mumasero mazhinji eukaryotic, mtDNA inoiswa muchimiro cheyakavharika denderedzwa reDNA mamorekuru, ayo, kusiyana nema chromosomes enyukireya emaseru eukaryotic, anodzokorora semi-achichengetedza uye haabatanidzwe nemapuroteni mamorekuru.
Iyo mitochondrial genome yaP. Infestans yakateedzana, uye akati wandei emabasa akatsaurirwa kuongororwa kwehurefu hwezvimedu zvezvimedu (Carter et al, 1990, Goodwin, 1991, Gavino, Fry, 2002). Mushure meGriffith naShaw (1998) vakagadzira nzira yakapusa uye inokurumidza yekumisikidza mtDNA haplotypes, mucherechedzo uyu wakazove mumwe weanonyanya kufarirwa muP.Pestestans zvidzidzo. Musimboti weiyi nzira unowedzerana mukusimudzira kwezvimedu zviviri zveMitochondrial DNA (kubva ku genome yakajairika) ine primers F2-R2 uye F4-R4 (Tafura 3) uye yavo inotevera kurambidzwa neanorambidza enzyme MspI (1st chidimbu) uye EcoR1 (2nd chidimbu). Iyo nzira inobvumira kuziva 4 haplotypes: Ia, IIa, Ib, IIb. Rudzi rwechipiri rwakasiyana nerudzi I nekuvapo kweinowisa 1881 bp muhukuru uye nenzvimbo yakasiyana yenzvimbo dzekurambidza munzvimbo dzeP2 neP4 (Fig. 3).
Kubva 1996, pakati pematambudziko akaunganidzwa mudunhu reRussia, chete haplotypes Ia na IIa vakaonekwa (Elansky et al., 2001, 2015). Vanogona kuzivikanwa mushure mekuparadzaniswa kwezvirambidzo zvigadzirwa neiyo yekutanga F2-R2 mumunda wemagetsi (Fig. 4, 5). Mhando dzemtDNA dzinoshandiswa mukuenzanisa kuongororwa kwematambudziko uye huwandu hwevanhu. Mune akati wandei emabasa, mhando dzeMitochondrial DNA dzakashandiswa kupatsanura clonal mitsetse uye paspassize P. infestans kuzviparadzanisa (Botez et al., 2007; Shein et al., 2009). Tichishandisa iyo PCR-RFLP nzira, zvakagumiswa kuti mtDNA yakasarudzika mune imwechete P. infestans strain (Elansky naMilyutina, 2007). Mamiriro ekusimudzira: 1x (500 sec. 94 ° C), 40x (30 sec. 90 ° C, 30 sec. 52 ° C, 90 sec. 72 ° C); 1x (5 min. 72 ° C). Reaction musanganiswa: (20 μl): 0,2 U Taq DNA polymerase, 1x 2,5 mM MgCl2-Taq buffer, 0,2 mM yega dNTP, 30 pM primer uye 5 ng yeiyo yakaongororwa DNA, deionized mvura - anosvika 20 μl.
Kurambidzwa kwechigadzirwa chePCR kunoitwa kwemaawa 4-6 panopisa 37 ° C. Kudzivirirwa musanganiswa (20 μl): 10x MspI (2 μl), 10x yekudzivirira buffer (2 μl), deionized mvura (6 μl), PCR chigadzirwa (10 μl).
Tafura 3. Primers anoshandiswa kukwidziridza mtDNA polymorphic matunhu
Locus | Primer | Primer kureba uye kuiswa | PCR chigadzirwa kureba | Dzora |
---|---|---|---|---|
P2 | F2: 5'- TTCCCTTTGTCCTCTACCGAT | 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 |
Random primer kuwedzera (RAPD)
Paunenge uchiita RAPD, imwe primer inoshandiswa (dzimwe nguva akati wandei ma primers panguva imwe chete) neanopokana nateuro yenoteotide, kazhinji gumi nucleotides pakureba, ine yakakwira yezvinyorwa (kubva ku10%) yeGC nucleotides uye yakadzika yekuwedzera tembiricha (ingangoita 50 ° C) . Aya ma primers "ivhu" pane akawanda ekuwedzera masayiti mugenome. Mushure mekusimudzira, huwandu hukuru hwemamplicons hunowanikwa. Huwandu hwavo hunoenderana neyekutanga (s) yakashandiswa uye nemamiriro ezvinhu (MgCl35 kusangana uye kupisa tembiricha).
Kufungidzira kwemaamplons kunoitwa ne distillation mu polyacrylamide kana agarose gel. Paunenge uchiita kuongorora kweRAPD, zvinodikanwa kuti unyatsoongorora kuchena kwezvinhu zvakaongororwa, nekuti kusvibiswa nezvimwe zvinhu zvipenyu kunogona kukonzera kuwedzera kwakanyanya muhuwandu hwezvigadzirwa, izvo zvakawandisa kunyangwe mukuongororwa kwezvinhu zvakachena (Perez et al, 1998). Kushandiswa kweiyi nzira mukudzidza kwe P. infestans genome kunoonekwa mumabasa mazhinji (Judelson, Roberts, 1999, Ghimire et al., 2002, Carlisle et al., 2001). Kusarudzwa kwemamiriro ezvinhu uye ma primers (51 10-nucleotide primers akafundwa) zvinopihwa muchinyorwa na Abu-El Samen et al., (2003).
Microsatellite Kudzokorora Kuongorora (SSR)
Microsatellite inodzokorora (yakapusa kuteedzana inodzokororwa, SSR) inodzokororwa nguva pfupi inoteedzana ye 1-3 (dzimwe nguva inosvika ku6) nucleotides inowanikwa mumagene enyukireya eukaryotes ese. Huwandu hwekudzokorora hunoteedzana hunogona kusiyana kubva pa10 kusvika 100. Microsatellite loci inoitika neyakaenzana yepamusoro uye yakawanda kana zvishoma zvakaenzana zvakagoverwa mukati meiyo genome (Lagercrantz et al., 1993). Polymorphism yekuteedzana kwema microsatellite inosanganiswa nemisiyano muhuwandu hwekudzokorora kwechikonzero chekutanga. Microsatellite mamaki anonyanya kukosha, izvo zvinoita kuti zvikwanise kuishandisa kuongorora chimiro chevanhu, kuona hukama, genotype migration pathways, nezvimwe. Pakati pezvimwe zvakanaka zvezvimakisi izvi, munhu anofanirwa kucherechedza yavo yakakura polymorphism, yakanaka kubereka, kusarerekera, uye kugona kuita otomatiki kuongorora uye kuongorora. . Pakutanga, kuongororwa kwakaitwa pamwe nekuparadzaniswa kwezvakaitwa zvigadzirwa pane polyacrylamide gel. Gare gare, vashandi veApplied Biosystems kambani yakaronga kushandisa fluorescently yakanyorwa ma primers pamwe nekuona kwekuita zvigadzirwa vachishandisa otomatiki laser detector (Diehl et al., 1990), uyezve zvakajairwa otomatiki DNA sequencers (Ziegle et al., 1992). Kunyora maprimer ane akasiyana fluorescent madhayi anotendera iwe kuti uongorore mamaki akati wandei pane imwe nzira uye, nekudaro, zvinowedzera zvakanyanya kugadzirwa kwenzira uye nekuvandudza kurongeka kweyekuongorora.
Iwo ekutanga mabhuku akanangidzirwa mukushandisa kweSSR ongororo yekuongorora P. infestans akaonekwa mukutanga kwema2000. (Knapova, Gisi, 2002). Haasi ese mamaki akarongwa nevanyori airatidza huwandu hwakaringana hwepolymorphism, zvisinei, maviri acho (4B neG11) akaverengerwa museti ye12 SSR mamaki akarongwa naLees et al. (2006) uye yakazogamuchirwa neEucablight research network (www.eucablight .org) sechiyero cheP infestans. Makore mashoma gare gare, chidzidzo chakaburitswa pamusoro pekugadzirwa kwesystem ye multiplex kuongororwa kweP infestans DNA yakavakirwa pamatanhatu eSSR maka (Li et al., 2010). Chekupedzisira, mushure mekuongorora mamaki ese akambofungidzirwa uye nekusarudza iwo anodzidzisa kwazvo, pamwe nekuvandudza ma primers, mavara e fluorescent, uye mamiriro ekusimudzira, timu rimwechete revanyori rakapa imwe-nhanho multiplex ongororo system yaisanganisira mamaki gumi nemaviri (Tafura 12; Li et al., 4a). Iwo ekutanga anoshandiswa mune ino sisitimu akasarudzwa uye akanyorwa nechimwe chezvina zvezvimisikidzo zvemagetsi (FAM, VIC, NED, PET) kuitira kuti iwo maratidziro ehukuru hwehukuru hwemaprimer ane iwo iwo mavara haana kuwiriranwa.
Vanyori vakaita ongororo pane PTC200 gudzazwi (MJ Research, USA) vachishandisa QIAGEN multiplex PCR kits kana QIAGEN Typeit Microsatellite PCR kits. Iyo vhoriyamu yemhinduro yekusanganisa yaive 12.5 μL. Mamiriro ekusimudzira aive akadai: eQIAGEN multiplex PCR: 95 ° C (15 min), 30x (95 ° C (20 s), 58 ° C (90 s), 72 ° C (60 s), 72 ° C ( 20 min); yeQIAGEN Type-it Microsatellite PCR): 95 ° C (5 min), 28x (95 ° C (30 sec), 58 ° C (90 sec), 72 ° C (20 sec), 60 ° C (30 min).
Kuparadzanisa uye kuona kwezvigadzirwa zvePCR kwakaitwa uchishandisa ABI3730 otomatiki capillary DNA analyzer (Applied Biosystems).
Tafura 4. Hunhu hwe12 standard SSR mamaki anoshandiswa kujenotyping P. Infestans (Li et al., 2013a)
Title | Nhamba yemarehole | Saizi renji alleles (bp) | Primers |
PiG11 | 13 | 130-180 | F: NED-TGCTATTTATCAAGCGTGGG R: GTTTCAATCTGCAGCCGTAAGA |
Pi02 | 4 | 255-275 | F: NED-ACTTGCAGAACTACCGCCC R: GTTTGACCACTTTCCTCGGTTC |
PinfSSR11 | 4 | 325-360 | F: NED-TTAAGCCACGACATGAGCTG R: GTTTAGACAATTGTTTTGTGGTCGC |
D13 | 16 | 100-185 | F: FAM-TGCCCCCTCTCTCTCTCT R: GCTCGAATTCATTTTACAGACTTG |
PinfSSR8 | 4 | 250-275 | FAM-AATCTGATCGCAACTGAGGG R: GTTTACAAGATACACACGTCGCTCC |
PinfSSR4 | 7 | 280-305 | FAM-TCTTGTTCGAGTATGGCGACG R: GTTTCACTTCGGGAGAAAGGCTTC |
Pi04 | 4 | 160-175 | F: VIC-AGCGGCTTTACCGATGG R: GTTTCAGCGGCTGTTTCGAC |
Pi70 | 3 | 185-205 | F: VIC-ATGAAAATACGTCAATGCTCG R: CGTTGGATATTTCTATTTCTTCG |
PinfSSR6 | 3 | 230-250 | F: GTTTTGGTGGGGCTGAAGTTTT R: VIC-TCGCCACAAGATTTATTCCG |
Pi63 | 3 | 265-280 | F: VIC-ATGACGAAGATGAAAGTGAGG R: CGTATTTTCCTGTTTATCTAACACC |
PinfSSR2 | 3 | 165-180 | F: PET-CGACTTCTACATCCACCGGC R: GTTTGCTTGGACTGCGTCTTTAGC |
Pi4B | 5 | 200-295 | F: PET-AAAATAAAGCCTTTGGTTCA R: GCAAGCGAGGTTTGTAGATT |
Muenzaniso wekutarisa iwo mhedzisiro yekuongorora unoratidzwa muFig. 6. Mhedzisiro yacho yakaongororwa uchishandisa iyo GeneMapper 3.7 software nekuenzanisa iyo yakawanikwa data neayo eanozivikanwa ega. Kugonesa kududzirwa kwemhedzisiro yekuongorora, zvakafanira kusanganisira 1-2 mareferenzi akapatsanurwa aine genotype inozivikanwa muchidzidzo chega chega.
Maitiro ekutsvagisa akarongwa akaedzwa pane akati wandei masampuli emunda, mushure mezvo vanyori vakaomesa zviga pakati pemarabhoritari emasangano maviri, The James Hutton Institute (UK) neWageningen University & Research (Netherlands), iyo, pamwe nemukana wekushandisa. akajairwa FTA makadhi eakareruka kuunganidzwa uye kutumirwa kweP infestans DNA masampuli akaita kuti zvikwanise kutaura nezve mukana wekushandisa kwekushambadzira kweichi chiitiko. Uye zvakare, yekutsanya uye chaiyo nzira yekuita genotyping P. infestans inoparadzanisa uchishandisa multiplex SSR ongororo yakaita kuti zvikwanisike kuitisa zviyero zvakaenzana zvevanhu veiyi pathogen pasirese pasirese, uye kugadzirwa kwepasirese yepasirese pane kunonoka blight mukati mehurongwa hwe Chirongwa cheEucablight (www.eucablight.org), kusanganisira, zvinosanganisira mhedzisiro ye microsatellite ongororo, zvakaita kuti zvikwanise kuteedzera kubuda uye kupararira kwemagenotypes matsva pasirese
Amplified kudzora chidimbu kureba polymorphism (AFLP). AFLP (yakawedzeredzwa chidimbu kureba polymorphism) hunyanzvi hwekugadzira zvisina kujairika mamorekuru mamaki vachishandisa chaiwo maprimer. Mune AFLP, DNA inorapwa pamwe nekubatanidzwa kwemaviri enzyme enzyme. Adapter chaiwo akasungirirwa kumatanho anonamira ezvimedu zvekudzivirira.
Izvi zvidimbu zvinobva zvawedzeredzwa zvichishandisa zvinowedzerana kune zvakateedzana seadapter uye saiti yekudzivirira uyezve zvichitakura imwechete kana zvimwe zvisina kujairika mabhesi kumagumo avo matatu. Iyo seti yezvimedu zvakawanikwa zvinoenderana nekuvharirwa ma enzyme uye zvakasarudzika zvakasarudzwa nucleotides pa3'-kumagumo ezvitangaridzo (Vos et al., 3). AFLP - genotyping inoshandiswa kukurumidza kudzidza kuchinja kwemagene ezvisikwa zvakasiyana.
Rondedzero yakadzama yenzira yacho inopihwa mumabasa aMueller, Wolfenbarger, 1999, Savelkoul et al., 1999. Basa rakawanda rekufananidza kugadzirisa kweAFLP neSSR nzira dzave kuitwa nevanoongorora veChinese. Iyo phenotypic uye genotypic hunhu hwe 48 P. infestans inoparadzanisa inounganidzwa kubva kumatunhu mashanu eNorth China yakadzidzwa. Iyo AFLP spectra yakaratidza masere akasiyana eDNA genotypes, kusiyana neiyo SSR genotypes, iyo isina kusiana kwakawanikwa (Guo et al., 2008).
Kuwedzeredzwa ne primers homologous kune akateerana enharembozha
Mamaki akatorwa kubva mukuteedzana kwetrotransposons ari nyore kwazvo kune mepu mepu, kudzidza kwemajini akasiyana uye shanduko maitiro (Schulman, 2006). Kana ma primers akagadzirwa anoenderana nekuteedzana kwakadzikama kwevamwe nharembozha, zvinokwanisika kukwidziridza matunhu egenome ari pakati pavo. Mune ongororo yeanokonzera causative ejoni yekupedzisira, nzira yekukudziridza zvikamu zvejenome uchishandisa primer inoenderana neyekuteedzana kweSINE (Pfupi Yakapeterwa Nuclear Elements) retropazone yakashandiswa zvinobudirira (Lavrova naElansky, 2003). Iyi nzira yakaburitsa misiyano kunyangwe mune yeasexual vana yeumwe wega. Panyaya iyi, zvakagumiswa kuti nzira yepakati - SINE - PCR yakanyatsojeka uye mwero wekufamba kwezvinhu zveSINE muPhytophthora genome yakakwira.
Mune genome yaP. Infestans, mhuri gumi nembiri dzemapfupi retrotransposons (SINEs) akaonekwa; kugoverwa kwemarudzi epfupi retrotransposons kwakaferefetwa, ma element (SINEs) akaonekwa ayo anowanikwa mune genome yeP chete infestans (Lavrova, 12).
Zvimiro zvekushandiswa kwenzira dzekuenzanisira kudzidza kwematambudziko muzvidzidzo zvevanhu
Paunenge uchironga kudzidza, zvinodikanwa kuti unzwisise zvakajeka zvinangwa zvainoteedzera uye kushandisa nzira dzakakodzera. Nekudaro, dzimwe nzira dzinobvumidza kuburitsa huwandu hukuru hwezviratidzo zvekumira zvakazvimirira, asi panguva imwe chete zvine yakaderera kubereka uye inotsamira zvakanyanya pama reagents anoshandiswa, mamiriro ekuita, uye kusvibiswa kwechinhu chiri kudzidzwa. Naizvozvo, muchidzidzo chega chega cheboka rematambudziko, zvinodikanwa kuti ushandise akati wandei (mareferenzi) akazviparadzanisa, asi kunyangwe mune ino kesi, mhedzisiro yezviedzo zvakati wandei zvakaoma kusanganisa.
Iri boka renzira dzinosanganisira RAPD, AFLP, InterSSR, InterSINE PCR. Mushure mekusimudzira, huwandu hwakawanda hwezvikamu zveDNA zvehukuru hwakasiyana hunowanikwa. Zvinokurudzirwa kushandisa matekiniki akadaro kana zvichidikanwa kumisikidza misiyano pakati pematambudziko ane hukama (mubereki-mwana, mhuka dzesango, nezvimwewo), kana muzviitiko panodiwa kuongororwa kwakadzama kwesampula diki. Nekudaro, iyo nzira yeAFLP inoshandiswa zvakanyanya mukubikwa kwemajini kweP infestans (van der Lee et al., 1997) uye muzvidzidzo zvekupinda mukati (Knapova, Gisi, 2002, Cooke et al, 2003, Flier et al, 2003). Nzira dzakadai hadzina kukodzera kushandisa paunenge uchigadzira dhatabhesi ematambudziko, nekuti zvinoita zvisingaite kubatanidza kuverenga kwemhedzisiro kana uchiita ongororo mumarabhoritari akasiyana.
Kunyangwe kuri pachena kupfava uye kumhanya kwekuurayiwa (DNA kusarudzika pasina kuchenesa kwakanaka, kuwedzera, kuona mhedzisiro), iri boka renzira rinoda kushandiswa kwenzira yakakosha yekunyora mhedzisiro: distillation mu polyacrylamide gel ine yakanyorwa kuti (radioactive kana luminescent) uye zvinotevera kuratidza kune mwenje kana radioactive zvinhu. Yakajairwa ethidium bromide agarose gel imaging kazhinji haina kukodzera kune idzi nzira nekuti nhamba hombe yezvimedu zveDNA zvehukuru hwakasiyana zvinogona kufiyiza.
Dzimwe nzira, pane zvinopesana, tendera kuburitsa huwandu hushoma hwezvinhu nekukwirisa kwavo kwakanyanya. Iri boka rinosanganisira kudzidza kwemitochondrial DNA haplotypes (maviri chete haplotypes Ia uye IIa anozivikanwa muRussia), mhando yekukwikwidza (mazhinji akapatsanurwa akapatsanurwa kuita marudzi maviri: A2 neA1, SF inozvitakura haiwanzo kuwanikwa) uye peptidase isozyme spectra ( maviri loci Pep2 naPep1, ane maviri maozimu ega ega) uye glucose-2-phosphate isomerase (muRussia hapana mutsauko wehunhu uhwu, kunyangwe kukosha kwepolymorphism kuchizivikanwa mune dzimwe nyika dzepasi). Zvinokurudzirwa kushandisa izvi maficha kana uchiongorora kuunganidzwa, kuunganidza matunhu nedunhu repasirese. Panyaya yekuongororwa kwesozymes uye haplotypes yeMitochondrial DNA, zvinokwanisika kuita pasina akasarudzika materu zvachose, nepo kuongororwa kwemhando dzekubatanidza kunoda maviri bvunzo akapatsanurwa ane anozivikanwa mating mhando.
Mamiriro ezvinhu uye ma reagents anogona chete kukanganisa mutsauko wechigadzirwa pa electrophoretogram; kuratidzwa kwezvigadzirwa mumhando idzi dzezvidzidzo hazvigoneke.
Parizvino, ruzhinji rwevanhu muchikamu cheEuropean cheRussia vanomiririrwa nematambudziko emhando mbiri dzekukwirana (Tafura 6), pakati pavo pane akazvipatsanura nemhando Ia uye IIa yemitochondrial DNA (mamwe marudzi emtDNA anowanikwa munyika aive haina kuwanikwa muRussia mushure megore ra1993). Iyo spectra ye peptidase isozymes inomiririrwa nemaviri genotypes paPep1 locus (100/100, 92/92 uye heterozygote 92/100, uye iyo 92/92 genotype inonyanya kushomeka (<0,3%)) uye maviri genotypes paPep 2 locus (100/100, 112/112 uye heterozygote 100/112, ine genotype 112/112 inoitika zvishoma zvishoma kupfuura 100/100, asi zvakare kazhinji).
Pakanga pasina misiyano mumitambo yezisoenzymes yeglucose-6-phosphate isomerase mushure megore ra1993 (kutsakatika kweiyo clonal mutsetse US-1); vese vakadzidza vakazviparadzanisa vaive ne100 / 100 genotype (Elansky naSmirnov, 2002).
Boka rechitatu renzira dzinoita kuti zvikwanisike kuwana rakaringana boka rezvazvimiririra mamaki maficha ane yakanyanya kubereka. Nhasi, boka iri rinosanganisira yekuongorora yeRFLP-RG57, iyo inogadzira 25-29 DNA zvidimbu zvehukuru hwakasiyana. RFLP-RG57 inogona kushandiswa zvese mukuongororwa kwemasampula uye mukuunganidzwa kwemadhatabhesi. Nekudaro, iyi nzira inodhura zvakanyanya kupfuura iyo yapfuura, inotora nguva, uye inoda yakakwana zvakakwana huwandu hwakanyanya hweDNA yakanatswa. Naizvozvo, muongorori anomanikidzwa kudzikisira huwandu hwezvinhu zvakaedzwa.
Kuvandudzwa kweRFLP-RG57 mukutanga kwema 90s ezana ramakore rapfuura kwakanyanyisa kuwedzeredza huwandu hwevaongorori veanokonzera causative mumiriri wekupedzisira kunaya. Yakave hwaro hwenzira inoenderana nesarudzo uye kuongororwa kwe "Clonal mitsetse" (ona pazasi). Pamwe chete neRFLP-RG57, mhando yekukwikwidza, DNA chigunwe (RFLP-RG57 nzira), spectra yepeptidase uye glucose-6-phosphate isomerase isoenzymes, uye mitochondrial DNA mhando inoshandiswa kuratidza clonal mitsetse. Kutenda kwaari, yakaratidzirwa al., 1994), kutsiviwa kwevanhu vekare nevanhu vatsva (Drenth et al, 1993, Sujkowski et al, 1994, Goodwin et al, 1995a), uye clonal mitsara inowanikwa munyika zhinji dze nyika yakazivikanwa. Zvidzidzo zvemitsara yeRussia vachishandisa nzira iyi zvakaratidza yakakwira genotypic polymorphism yematambudziko echikamu cheEuropean uye monomorphism yevagari veAsia neFar Eastern zvikamu zveRussia (Elansky et al, 2001). Uye ikozvino nzira iyi inoramba iri huru muzvidzidzo zvehuwandu hwevanhu P. infestans. Nekudaro, kugoverwa kwayo kwakakura kunodziviswa nemutengo wayo wakakwira uye kusimba kwevashandi mukuuraya.
Imwe nzira yekuvimbisa isingawanzoshandiswa muzvidzidzo zve P. infestans ndeye microsatellite dzokorora (SSR) kuongorora. Parizvino, iyi nzira inoshandiswa zvakanyanya kupatsanura clonal mitsara. Zvekuongororwa kwematambudziko, akadai phenotypic mucherechedzo hunhu sekuvapo kwehutachiona mageneti emhando dzembatatisi (Avdey, 1995, Ivanyuk et al., 2002, Ulanova et al., 2003) uye tomato zvakashandiswa zvakanyanya (uye ramba uchishandiswa) . Parizvino, magene ehutachiona kumhando dzembatatisi akarasikirwa nemutengo wawo sehunhu hunhu hwezvidzidzo zvevanhu nekuda kwechitarisiko cheiyo yakanyanya (kana padyo nayo) huwandu hwehutachiona hwehutachiona mune huwandu hwakawanda hwevasungwa. Panguva imwecheteyo, iyo T1 virulence geni yemadomasi mhando inotakura inoenderana Ph1 geni ichiri kubudirira kushandiswa sechiratidzo (Lavrova et al., 2003; Ulanova et al., 2003).
Mumabasa mazhinji, kuramba fungicides kunoshandiswa senzira yekumaka. Hunhu uhu haudiwe kushandiswa muzvidzidzo zvevanhu nekuda kwechitarisiko chiri nyore chekushomeka mumitsara yeclonal mushure mekushandisa metalaxyl- (kana mefenoxam-) ine fungicides mumunda. Semuenzaniso, misiyano yakakosha padanho rekurwisa yakaratidzirwa mukati meiyo Sib1 clonal mutsara (Elansky et al., 2001).
Nekudaro, mhando yekukwikwidza, peptidase isozyme spectrum, mitochondrial DNA mhando, RFLP-RG57, SSR ndiyo yakasarudzika mamaki ehunhu ekugadzira dhatabheki uye yekunyora maratidziro muzvikamu. Kuti uenzanise mashoma masampuli, kana zvichidikanwa kuisa yakanyanya kuwanda mamaki maficha, unogona kushandisa AFLP, RAPD, InterSSR, Inter-SINE PCR (Tafura 5). Nekudaro, zvinofanirwa kuyeukwa kuti nzira idzi hadzina kubereka zvakanaka, uye mune yega yega kuyedza (kukwidziridza electrophoresis kutenderera) kunofanirwa kushandisa akati wandei mareferenzi.
Tafura 5. Kufananidza kwenzira dzakasiyana dzekutsvaga kwekutambudzika P. infestans
chokuratidza | TS | Isofer mapurisa | MtDNA | RFLP-RG57 | RAPD | ISSR | SSR | AFLP | Zvak |
---|---|---|---|---|---|---|---|---|---|
Huwandu hwemashoko | Н | Н | Н | С | В | В | С | В | В |
Kuberekwazve | В | В | В | В | Н | Н | С | С | С |
Kubvira kwezvigadzirwa | Н | Н | Н | Н | В | С | Н | С | В |
mari | Н | С | Н | В | Н | Н | Н | С | Н |
Kushanda nesimba | Н | Н | Н | В | NS * | NS * | Н | С | NS * |
Kuongorora kumhanya ** | В | Н | Н | С | Н | Н | Н | Н | Н |
Tarira: H - yakaderera, C - yepakati, B - yakakwira; НС * - kusimba kwevashandi kwakaderera kana uchishandisa agarose gel kana otomatiki
genotyper, yepakati - kana yakadonhedzwa mu polyacrylamide gel ine mavara akanyorwa,
** - kwete kuverenga iyo nguva yakashandiswa pakukura kwemycelium yeDNA kusarudzika.
Chiumbwa chevagari
Clonal mitsara
Mukushayikwa kwekudzokororazve kana nemupiro wayo usina kukosha kuhurongwa hwehuwandu hwevanhu, huwandu hwevanhu hunosanganisira imwe nhamba yemakona, kuchinjaniswa kwemajini pakati payo zvisingawanzoitika.
Mune vanhu vakadai, zvinonyanya kudzidzisa kudzidza kwete kuwanda kwemajini, asi mafambiro egenotypes ane zvakafanana mavambo (clonal mitsetse kana clonal mitsetse) uye anosiyana chete nekuchinja poindi. Dzidzo yevagari vekunonoka blight pathogen uye kuongororwa kwemitsara yeklonal kwakawedzera zvakanyanya kubva pakauya nzira yeRFLP-RG57 mukutanga kwema90s ezana ramakore rapfuura. Pamwe chete neRFLP-RG57, mhando yekukwirisa, spectra ye peptidase uye glucose-6-phosphate isomerase isoenzymes, uye mitochondrial DNA mhando inoshandiswa kuzivisa clonal mitsetse. Hunhu hwemitsetse yakanyanya kuwanda inoratidzwa muTafura 6.
Clone US-1 yakatonga huwandu hwevanhu kwese kwese kusvika pakupera kwema80, mushure maizvozvo yakatanga kutsiviwa nemamwe macone uye yakanyangarika kubva kuEurope neNorth America. Ikozvino yawanikwa muFar East (Philippines, Taiwan, China, Japan, Korea, Koh et al., 1994, Mosa et al, 1993), mu Africa (Uganda, Kenya, Rwanda, Goodwin et al, 1994, Vega- Sanchez et al., 2000; Ochwo et al., 2002) uye muSouth America (Ecuador, Brazil, Peru, Forbes et al., 1997, Goodwin et al., 1994). Hapana mitsara iri yeUS-1 mutsara yakaonekwa muAustralia chete. Sezvineiwo, P. infestans vakazviparadzanisa vakauya kuAustralia nerimwe fungu rekutama (Goodwin, 1997).
Clone US-6 yakatama kubva kuchamhembe kweMexico ichienda kuCalifornia kuma70 apera uye yakakonzera denda imomo mumatezi nematomatisi mushure memakore makumi matatu nemaviri asina chirwere. Nekuda kwehukasha hwayo, yakatsiva iyo US-32 dombo ndokutanga kutonga kumahombekombe egungwa eUnited States (Goodwin et al., 1a).
Ma genotypes US-7 neUS-8 akawanikwa muUnited States muna 1992 uye atove muna 1994 akaparadzirwa muUnited States neCanada. Munguva imwe yemumunda mwaka, dombo reUS-8 rinokwanisa kubvisa zvachose US-1 munzvimbo dzembatatisi dzakatanga kutapukirwa nemakombo maviri akaenzana (Miller naJohnson, 2000).
Clones BC-1 kusvika BC-4 akaonekwa muBritish Columbia mune vashoma vanozviparadzanisa naGoodwin et al., 1995b). Clone US-11 yakapararira zvakanyanya muUnited States uye yakatsiva US-1 muTaiwan. Clones JP-1 uye EC-1, pamwe chete nekodhi US-1, zvakajairika muJapan neEcuador, zvichiteerana (Koh et al., 1994; Forbes et al., 1997).
SIB-1 idombo rakakunda muRussia pamusoro penzvimbo yakakura kubva mudunhu reMoscow kusvika kuSakhalin. Munharaunda yeMoscow, yakawanikwa muna1993, uye vamwe vanhu vemumunda vaiwanzova nematambudziko eiyi clonal mutsetse, unopesana zvakanyanya ne metalaxyl. Mushure me1993, kuwanda kweiyi clone kwakaderera zvakanyanya. Kunze kweUral muna 1997-1998, SIB-1 yakawanikwa kwese kwese, kunze kweKhabarovsk Territory (iyo SIB-2 dombo yakapararira ipapo). Kupatsanurwa kwenzvimbo kwema clones nemhando dzakasiyana dzekusvitsa hakubatanidzi kuita kwepabonde muSiberia neFar East. Munharaunda yeMoscow, kusiyana neSiberia, huwandu hwevanhu hunomiririrwa nemhando dzakawanda dzedombo; ingangoita yega yega yekuzviparadzanisa ine yakasarudzika multilocus genotype (Elansky et al., 2001, 2015). Iyi misiyano haigone kutsanangurwa chete nekutorwa kwemarudzi ehuswa kubva kunzvimbo dzakasiyana dzenyika nembeu dzekunze dzakatorwa. Sezvo mhando mbiri dzekusvirana dzinoitika muhuwandu, zvinokwanisika kuti kusiyana kwaro kuri zvakare nekuda kwekudzokorora. Nekudaro, muBritish Columbia, kumuka kwemagenotypes BC-2, BC-3, uye BC-4 kunofungidzirwa nekuda kwekusanganiswa kwematombo BC-1 uye US-6 (Goodwin et al., 1995b). Izvo zvinokwanisika kuti masanganiswa akasanganiswa anowanikwawo muMoscow. Semuenzaniso, kumanikidza MO-4, MO-8 uye MO-11 heterozygous yePEP locus inogona kuva mahybrids pakati pematambudziko MO-12, MO-21, MO-22, iine iyo A2 pairing mhando uye homozygous kune imwechete allele yePEP. locus uye kumanikidza MO-8, iine iyo A1 mating mhando uye homozygous yeimwe allele yeiyo locus. Uye kana izvi zviri izvo, uye muvazhinji vemazuvano e P. infestans pane maitiro ekuwedzera kune iro basa rekuita bonde, ipapo kukosha kweruzivo rwekuongororwa kwema multilocus clones kuchaderera (Elansky et al., 2001, 2015 ).
Kusiyanisa mumitsara yeclonal
Kusvikira kuma90 ezana ramakore rechimakumi maviri, iyo clonal mutsetse US-20 yakanga yakapararira pasirese. Mazhinji emunda uye enharaunda vanhu vaingova chete nematambudziko neUS-1 genotype. Nekudaro, mutsauko uripo pakati pevakazviparadzanisa vakaonekwawo, ungangove unokonzerwa nemaitiro ekuchinja. Shanduko dzakaitika mune zvese nyukireya uye mitochondrial DNA uye zvakakanganisa, pakati pezvimwe zvinhu, danho rekurwisa phenylamide zvinodhaka uye huwandu hwehutachiona hutachiona. Mitsara inosiyana neyekutanga genotypes nekuchinja inoratidzwa nenhamba dzekuwedzera mushure medheti rinotevera zita reiyo genotype yekutanga (semuenzaniso, iyo US-1 mutant mutsara weiyo clonal mutsetse US-1.1). Kugadzira zvigunwe zveDNA mitsara US-1 uye US-1.5 ine mitsara yekuwana yehukuru hwakasiyana (Goodwin et al., 1.6a, 1995b); iyo clonal mutsetse US-1995 inosiyanawo kubva kuUS-6.3 mune imwechete yekuwedzera mutsara (Goodwin, 6, Tafura 1997).
Muchidzidzo cheMitochondrial DNA, zvakawanikwa kuti chete mhando 1b yeDNA yemitochondrial inowanikwa mumutsetse weclonal US-1 (Carter et al., 1990). Nekudaro, mune ongororo yematambudziko eiyi clonal mutsetse kubva kuPeru nePhilippines, vakazviparadzanisa zvakacherechedzwa, mhando dzeMitochondrial DNA iyo yaisiyana kubva pa1b nekuvapo kwekuiswa nekubviswa (Goodwin, 1991, Koh et al., 1994) .
Tafura 6. Multilocus genotypes yemamwe P. infestans clonal mitsetse
Title | Kusangana mhando | Isozymes | DNA zvigunwe | MtDNA mhando | |
GPI | PEP | ||||
US-1 | А1 | 86/100 | 92/100 | 1.0111010110011E + 24 | Ib |
US-2 | А1 | 86/100 | 92/100 | 1.0111010010011E + 24 | - |
US-3 | А1 | 86/100 | 92/100 | 1.0111000000011E + 24 | - |
US-4 | А1 | 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 |
Ongorora: * - hapana data.
Tafura 7. Multilocus genotypes uye yavo mutant mitsetse
Title | Kusangana mhando | | DNA Zvigunwe (RG57) | Notes | |
GPI | PEP-1 | ||||
US-1 | А1 | 86/100 | 92/100 | 1011101011001101000110011 | Yekutanga genotype 1 |
US-1.1 | A1 | 86/100 | 100/100 | 1011101011001101000110011 | Kuchinja muPEP |
US-1.2 | A1 | 86/100 | 92/100 | 1011101010001101000110011 | Kuchinja muRG57 |
US-1.3 | A1 | 86/100 | 92/100 | 1011101001001101000110011 | Kuchinja muRG57 |
US-1.4 | A1 | 86/100 | 100/100 | 1011101010001101000110011 | Kuchinja muRG57 uye PEP |
US-1.5 | A1 | 86/100 | 92/100 | 1011101011001101010110011 | Kuchinja muRG57 |
US-6 | A1 | 100/100 | 92/100 | 1011111001001100010110011 | Yekutanga genotype 2 |
US-6.1 | A1 | 100/100 | 92 /92 | 1011111001001100010110011 | Kuchinja muPEP |
US-6.2 | A1 | 100/100 | 92/100 | 1011101001001100010110011 | Kuchinja muRG57 |
US-6.3 | A1 | 100/100 | 92/100 | 1011111001011100010110011 | Kuchinja muRG57 |
US-6.4 | A1 | 100/100 | 100/100 | 1011011001001100010110011 | Kuchinja muRG57 uye PEP |
US-6.5 | A1 | 100/100 | 92/100 | 1011111001001100010010011 | Kuchinja muRG57 |
BR-1 | A2 | 100/100 | 100/100 | 1011101000001100001111011 | Yekutanga genotype 3 |
BR-1.1 | A2 | 100/100 | 100/100 | 1010101000001100001110011 | Kuchinja muRG57 |
Iko kune zvakare shanduko mune yakatarwa yesozymes. Sezvo mutemo, ivo vanokonzerwa nekuparara kweiyo muviri pakutanga heterozygous yeiyi enzyme kuva homozygous. Muna 1993, pamichero yemadomasi, takacherekedza kusagadzikana nehunhu hunhu hweUS-1: RG57 kudhindwa kweminwe, mitochondrial DNA mhando, uye 86/100 genotype yeglucose-6-phosphatizomerase, asi yaive homozygous (100/100) yekutanga. peptidase locus pachinzvimbo che 92/100 heterozygote chaiyo yeiyi clonal mutsetse. Isu takatumidza iyo genotype yeiyi strain MO-17 (Tafura 6). Iyo mitsara inoshanduka US-1.1 uye US-1.4 zvakare inosiyana kubva kuUS-1 nekuchinja pama peptidase locus ekutanga (Tafura 7).
Shanduko dzinotungamira mukuchinjika kwehuwandu hwehutachiona hwemhando dzembatatisi nematomatisi zvakajairika. Ivo vaizivikanwa pakati pevamwe vari vega veiyo clonal mutsetse US-1 muhuwandu kubva kuNetherlands (Drenth et al., 1994), Peru (Goodwin et al., 1995a), Poland (Sujkowski et al., 1991), kuchamhembe kweNorth America ( Goodwin et al.,., 1995b). Misiyano muhuwandu hwembatatisi virulence majini akaonekwawo pakati pevamwe vari pamatanho eclonal mitsara US-7 uye US-8 muCanada neUnited States (Goodwin et al., 1995a), pakati pevamwe veSIB-1 mutsetse muAsia. chikamu cheRussia (Elansky et al, 2001).
Isolates ine yakasimba misiyano mumatanho ekurwisa phenylamide madhiragi akaonekwa mune monoclonal field populations, ese aive eiyo clonal mutsetse Sib-1 (Elansky et al, 2001, Tafura 1). Anenge ese maratidziro eiyo clonal mutsetse US-1 anonyanya kubatika kune metalaxyl; zvisinei, zvinodzivirirwa zvakanyanya zvega zvega mutsetse uyu zvakasarudzika muPhilippines (Koh et al., 1994) uye muIreland (Goodwin et al., 1996).
Mazuva ano vanhu veP. Infestans
Central America (Mexico)
Huwandu hwevanhu P. infestans muMexico hwakasiyana zvakanyanya kubva kune vamwe vanhu venyika, izvo zvinonyanya kukonzerwa nenzvimbo yaro yenhoroondo. Dzidzo dzakawanda dzehuwandu hwevanhu uye dzakabatana P. infestans marudzi e clade Phytophthora, pamwe nemhando dzenzvimbo dzerudzi rweSolanum, zvakatungamira mukufunga kuti shanduko yepathogen iri pakati peMexico yakaitika pamwe nekushanduka kwemiti inogarwa. uye yaibatanidzwa nekurara kwepabonde (Grünwald, Flier, 2005). Mhando mbiri dzekusvirana dzinomiririrwa muhuwandu hwevanhu, uye zvakaenzana, uye kuvepo kwemaoospores muvhu, pamiti uye machubhu embatatisi uye nemhando dzakabatana dzinokura musango Solanum inosimbisa kuvepo kwekuita bonde muhuwandu (Fernández- Pavía et al., 2002). Ongororo dzichangoburwa dzeToluca Valley nenzvimbo dzakapoteredzwa (iyo inofungidzirwa nzvimbo yekumusoro kwepathogen) yakasimbisa huwandu hwakasiyana hwemagariro evagari vemuP. mudunhu (Wang et al., 134). Zvinhu zvinopa mutsauko uyu kupatsanurana kwenzvimbo dzinowanikwa munzvimbo dzakakwirira dzepakati peMexico, misiyano mumamiriro ekurima nemhando dzembatatisi dzinoshandiswa mumipata nemakomo, uye kuvapo kwemhando dzesango dzinotyisa dzeSolanum dzinogona kuita sevamwe vanogamuchira (Fry et al. ., 176).
Nekudaro, zvinofanirwa kucherechedzwa kuti huwandu hwevanhu veP. Infestans kuchamhembe kweMexico vane hunyanzvi mune zvakasikwa uye hwakafanana zvakanyanya neNorth America, izvo zvinogona kuratidza kuti aya ndiwo matsva genotypes (Fry et al., 2009).
North America
Vanhu vekuNorth America veP. Infestans vagara vaine chimiro chakareruka uye hunhu hwavo hwakagadzwa nguva refu kusati kwashandiswa kuongorora microsatellite. Kusvika 1987, iyo clonal mutsetse US-1 yakatonga muUnited States neCanada (Goodwin et al., 1995). Pakati-ma70, apo metalaxyl-based fungicides yakaonekwa, dombo iri rakatanga kutsiviwa nemamwe, anodzivirira genotypes akatama achibva kuMexico (Goodwin et al., 1998). Pakupera kwema90s. iyo US-8 genotype yakatsiva zvachose iyo US-1 genotype muUnited States uye ikave yakakurisa clonal mutsetse pamatatata (Fry et al., 2009; Fry et al., 2015). Mamiriro akasiyana akakwenenzverwa nematomatisi, ayo aigara aine mitsara yakati wandei, uye maumbirwo avo akachinja gore rimwe nerimwe (Fry et al., 2009).
Mugore ra2009, denda rakakura kwazvo rekupedzisira kweblight rakatanga muUnited States pamadomasi. Chinhu cheiyi denda chaive kutanga kwayo panguva imwe chete munzvimbo zhinji kuchamhembe kwakadziva kumabvazuva kweUnited States, uye zvakazove zvakabatana nekutengesa kukuru kwematombo ematomato ane hutachiona munzvimbo huru dzebindu (Fry et al., 2013). Kurasikirwa kwezvirimwa kwaive kwakakura. Microsatellite ongororo yemasampuli akanganiswa yakaratidza kuti dambudziko rehosha iri raive rekukwirana US-22 A2-mhando kusangana. Muna 2009, mugove weiyo genotype muvanhu veAmerica ve P. infestans yakasvika 80% (Fry et al., 2013). Mumakore akatevera, huwandu hwehasha genotypes US-23 (kunyanya pamadomasi) uye US-24 (pamatatata) yakaramba ichiwedzera muhuwandu, zvisinei, mushure megore ra2011, huwandu hwekuonekwa kweUS-24 hwakadzikira zvakanyanya, uye kusvika nhasi, vangangoita 90% yehuwandu hwehutachiona muUnited States inomiririrwa neUS-23 genotype (Fry et al., 2015).
MuCanada, senge muUnited States, pakupera kwema90s. iro rinotungamira genotype US-1 rakadzoserwa neUS-8, chinzvimbo chikuru chakaramba chisina kuchinja kusvika 2008. MuCanada, maive nematambudziko akakomba anononoka kusangana nekutengesa kweanotapira matomatisi, asi akakonzerwa nemagenotypes US-2009 uye US-2010 (Kalischuk et al., 23). Kupatsanurwa kwakajeka kwenzvimbo kwemagenotypes aya kwakaratidza kuve kunoshamisa: US-8 yakatonga matunhu ekumadokero eCanada (2012%), nepo US-23 ichitonga matunhu ekumabvazuva (68%). Mumakore akatevera, US-8 yakapararira kumatunhu ekumabvazuva; zvisinei, kazhinji, mugove waro muvanhu wakadzikira zvishoma pakatarisana nekuonekwa kweiyo US-83 uye US-23 genotypes munyika (Peters et al., 22). Parizvino, US-24 inochengetedza chinzvimbo chakakura muCanada yese; US-2014 iripo muBritish Columbia, nepo US-23 neUS-8 varipo muOntario (Peters, 23).
Nekudaro, vanhu vekuNorth America veP. Infestans vanowanzo kuve clonal mitsetse. Mumakore makumi mana apfuura, huwandu hweakachekwa maclonal genotypes asvika makumi maviri nemana. Kunyangwe chokwadi chekuti mhando dzemhando mbiri dzekukwirisa dziripo muhuwandu, mukana wekuonekwa kwemajenotype matsva semhedzisiro yekurara pabonde unoramba uri pasi. Zvakangodaro, mumakore ekupedzisira makumi maviri, zviitiko zvinoverengeka zvekuonekwa kwevakawanda vanodzokorodza vanhu zvakanyorwa (Gavino et al., 40; Danies et al., 24; Peters et al., 20), uye mune imwe nguva mhedzisiro ye kuyambuka kwaive genotype US-2000, iyo yakadzika muNorth America kwemakore mazhinji (Gavino et al., 2014). Kusvikira 2014, shanduko mune chimiro chevanhu chaive chakabatanidzwa nekumuka kwezvitsva, zvine hukasha genotypes nekutevera kwavo kutama uye kusuduruka kwevaimbove vatungamiriri vekare. Chii chakaitika muna 11-2000 MuU.SA neCanada, epiphytotic kekutanga yakaratidza kuti munguva yekudyidzana kwenyika dzepasi rose, kuputika kwechirwere kunogona kusangana nekupararira kuri kuita kwemajenotypes matsva pakutengesa zvinhu zvekudyara zvine hutachiona.
South America
Kusvikira nguva pfupi yadarika, zvidzidzo zvevanhu veSouth America veP. Infestans zvaive zvisina kujairika kana hukuru. Izvo zvinozivikanwa kuti chimiro cheava vanhu chiri nyore uye chinosanganisira 1-5 clonal mitsetse pasirese (Forbes et al., 1998). Saka, na1998, genotypes US-1 (Brazil, Chile) BR-1 (Brazil, Bolivia, Uruguay, Paraguay), EC-1 (Ecuador, Colombia, Peru neVenezuela), AR-1, AR -2, AR- 3, AR-4 uye AR-5 (Argentina), PE-3 uye PE-7 (kumaodzanyemba kwePeru). Kusangana mhando A2 yaivepo muBrazil, Bolivia neArgentina uye haina kuwanikwa kupfuura mhiri kwemuganhu weBolivia nePeruvia munzvimbo yeLake Titicaca, kumashure uko EC-1 A1 genotype yaitonga muAndes. Pamadomasi, US-1 yakaramba iri mukurumbira genotype muSouth America yese.
Mamiriro acho akawedzera kana kushomeka muma 2000s. Chinhu chakakosha kwaive kuwanikwa kweiyo nyowani clonal mutsara EC-2 yerudzi rweA2 pane hama dzemusango dzembatatisi (S. brevifolium uye S. tetrapetalum) kuNorth Andes (Oliva et al., 2010). Zvidzidzo zvePhylogenetic zvakaratidza kuti mutsetse uyu hauna kunyatso kufanana neP infestans, kunyangwe iri yakabatana kwazvo nawo, zvine chekuita nezvawakarongedzerwa kuti uutarise, pamwe neumwe mutsetse, EC-3, yakasarudzika kubva pamuti wedomasi S. betaceum inokura muAndes, mhando nyowani inonzi P. andina; zvisinei, chimiro cherudzi urwu (yakazvimirira mhando kana musanganiswa weP infestans neimwe tambo isingazivikanwe) hazvisati zvanyatsojeka (Delgado et al., 2013).
Parizvino, vanhu vese vekuSouth America veP infestans vakaumbana. Kunyangwe paine kuvapo kwemhando mbiri dzekubatana, hapana vanhu vanodzokororwa vakaonekwa. Pamadomasi, iyo US-1 genotype iri kwese kwese, zvinoita kunge yabviswa pamatata nematambudziko emuno, iyo chaiyo mavambo ayo achiri kusazivikanwa. Iyo BR-1 genotype iripo muBrazil, Bolivia neUruguay; muPeru, pamwe neUS-1 uye EC-1, kune mamwe akawanda genotypes emuno. MuAndes, chinzvimbo chikuru chinochengeterwa nekonline mutsara EC-1, hukama hwayo neiyo ichangobva kuwanikwa P. andina inoramba isina kuongororwa. Iyo chete "isina kugadzikana" nzvimbo iko kune iyo nguva 2003-2013. pakave nekuchinja kukuru muhuwandu hwevanhu, vakave Chile (Acuña et al., 2012), iko muna 2004-2005. huwandu hwehutachiona hwakave nekushomeka kune metalaxyl uye nyowani haplotype yeMitochondrial DNA (Ia pachinzvimbo cheiyo yaivapo kare Ib). 2006 kusvika 2011 Muhuwandu hwevanhu, genotype 21 (sekureva SSR) yakatonga, mugove wacho wakasvika 90%, mushure mezvo chanza chakapfuudza kuna genotype 20, kuwanda kwechiitiko icho mumakore maviri akatevera kwakachengetwa kungangoita 67% (Acuña, 2015).
Europe
Mune nhoroondo yeEurope, kwaive neanenge maviri mafungu ekusimuka kweP infestans kubva kuNorth America: muzana ramakore rechi1. (HERB-1) uye kutanga kwezana ramakumi maviri (US-70). Kupararira kwakapararira kwemasitalaxyl-ine fungicides muma1s. kwakatungamira mukubviswa kweiyo yakakurisa genotype US-XNUMX uye kutsiva kwayo nema genotypes matsva. Nekuda kweizvozvo, munyika zhinji dzeWestern Europe, huwandu hwehutachiona hwakamiririrwa zvakanyanya nemitsetse yakati wandei.
Iko kushandiswa kwe microsatellite ongororo yekuongororwa kwehutachiona hwevanhu kwakaita kuti zvikwanisike kuona shanduko dzakakomba dzakaitika muWestern Europe muna 2005-2008. Muna 2005, imwe clonal lineage yakawanikwa muUK, inonzi 13_A2 (kana "Blue 13") uye inozivikanwa neiyo A2 mhando yekukwikwidza, kukwirisa kwakanyanya uye kuramba phenylamides (Shaw et al., 2007). Iyo genotype imwechete yakawanikwa mumasampuli akaunganidzwa muna2004 muNetherlands nekuchamhembe kweFrance, izvo zvakaratidza kuti yakatamira kuUK ichibva ku Continental Europe, pamwe nembesa mbatatisi (Cooke et al., 2007). Kuongororwa kwemajenome evamiriri veiyi clonal mutsetse kwakaratidza huwandu hwakakura hwepolymorphism yekuteedzana kwayo (na2016, huwandu hwayo hwakapatsanurwa hwasvika 340) uye hwakakosha misiyano iri padanho rejini, incl. zvinoshanda magene panguva yechirimwa hutachiona (Cooke et al., 2012; Cooke, 2017). Aya maficha, pamwe nekuwedzera kwenguva yeiyo biotrophic chikamu, zvinogona kunge zvakakonzera kuwedzera kwehasha kwe13_A2 uye kugona kwayo kutapura kunyangwe mhando dzematapiri dzinopokana nehusiku hunononoka.
Mumakore mashoma akatevera, genotype yakakurumidza kupararira nenyika dzekuchamhembe kwakadziva kumadokero kweEurope (Great Britain, Ireland, France, Belgium, Netherlands, Germany) pamwe nekubviswa panguva imwe chete kwema genotypes aimbove akakurisa 1_A1, 2_A1, 8_A1 (Montarry et al. , 2010; Gisi et al., 2011; Van den Bosch et al., 2011; Cooke, 2015; Cooke, 2017). Zvinoenderana newebsite www.euroblight.net, mugove we13_A2 muhuwandu hwevanhu venyika idzi hwasvika 60-80% uye nezvimwe; kuvepo kweiyo genotype kwakanyorwawo mune dzimwe nyika dzeEastern neSouthern Europe. Nekudaro, muna 2009-2012. 13_A2 yakarasikirwa nenzvimbo dzayo dzepamusoro muBritain neFrance, ichipa mutsara we6_A1 (8_A1 muIreland), uye muNetherlands neBelgium yakadzoserwa chikamu nemagenotypes 1_A1, 6_A1, uye 33_A2 (Cooke et al., 2012; Cooke, 2017; Stellingwerf, 2017).
Kusvika pari zvino, dzinosvika makumi manomwe kubva muzana (70%) yevagari vemuWestern Europe vaP. Zvinoenderana webhusaiti www.euroblight.net, akakurisa genotypes munyika dzeNorth West Europe (UK, France,
Netherlands, Belgium) inoramba iri, muzvikamu zvakaenzana zvakaenzana, 13_A2 uye 6_A1, iyo yekupedzisira haina kuwanikwa kunze kwenzvimbo yakatarwa (kunze kweIreland), asi inotova nematanho makumi mashanu nemasere (Cooke, 58). Misiyano 2017_A13 iripo muzviverengeka zvinoonekwa muGerman, uye inoonekwawo pano neapo munyika dzeCentral neSouthern Europe. Genotype 2_A1 inoumba chikamu chakakosha chevanhu veBelgium uye zvishoma neNetherlands neFrance. Genotype 1_A8 yakadzikama muhuwandu hweEurope padanho re1-3%, kunze kweIreland, uko inochengetedza nzvimbo yayo yekutungamira uye yakakamurwa kuita maviri madiki (Stellingwerf, 6). Pakupedzisira, muna 2017, pakanga paine kuwedzera kwehuwandu hwezviitiko zvema genotypes matsva 2016_A36 uye 2_A37, yakatanga kunyorwa muna 2-2013; parizvino, aya magenotypes anowanikwa muNetherlands neBelgium uye pamwe muFrance neGerman, pamwe nekuchamhembe kweGreat Britain (Cooke, 2014). Inenge 2017-20% yevevanhu vekuMadokero kweEurope vanomiririrwa gore rega rega neakasarudzika genotypes.
Kusiyana neWestern Europe, panguva iyo 13_A2 genotype payakaonekwa, vanhu vekuNorth Europe (Sweden, Norway, Denmark, Finland) havana kumiririrwa nemitsetse, asi nenhamba huru yeakasarudzika genotypes (Brurberg et al.,
2011). Munguva yekupararira kwe13_A2 kuWestern Europe, kuvapo kweiyo genotype muScandinavia hakuna kucherechedzwa kusvika muna2011, payakatanga kuwanikwa muNorth Jutland (Denmark), uko kunonyanya kurimwa mhando dzematapiri emabhizimusi nekushandiswa kunoitwa metalaxyl -ine fungicides (Nielsen et al., 2014). Sekureva kwa www.euroblight.net, genotype 13_A2 yakawanikwa zvakare mumasampuli akati wandei kubva kuNorway neDenmark muna2014 uye muzviverengero zvinoverengeka zveNorway muna2016; uyezve, mu2013 muFinland kuvepo kwechinguva chidiki cheiyo 6_A1 genotype kwakacherechedzwa. Chikonzero chikuru chekutadza kwe13_A2 uye mimwe mitsara yakakunda mukukunda kweScandinavia inoonekwa seyakasiyana mamiriro ekunze enharaunda ino kubva kunyika dzeWestern Europe.
Kuwedzera kune iyo yekuti inotonhora mwaka yechando uye kutonhora kwechando zvinobatsira kupona kwe oospores kwete pane vegetative mycelium (Sjöholm et al., 2013), ivhu rinotonhora muchando (izvo zvisingawanzo kuitika munyika dzinodziya dzeWestern Europe) zvinobatsira Kubvumirana kwe oospores kumera nekudyara. mbatatisi, izvo zvinowedzera basa ravo sosiyo yehutachiona hwehutachiona (Brurberg et al., 2011). Izvo zvinofanirwawo kucherechedzwa kuti, kuchamhembe, kukura kwehutachiona kubva kune oospores kunopfuura kukura kwehutachiona hutachiona, uhwo pakupedzisira hunodzivirira kutonga kweanotonyanya hasha, asi yakazogadzirwa mitsara yeclonal (Yuen, 2012). Maumbirwo eanonyanya kudzidza vanhu veP. Infestans muEastern Europe (Poland, iyo Baltic States) yakafanana neiyo muScandinavia.
Mhando mbiri dzekusvirana dziripowo pano, uye ruzhinji rwemagenotypes akatsunga kushandisa SSR ongororo akasiyana (Chmielarz et al., 2014; Runno-Paurson et al., 2016). Sezvakaita kuchamhembe kweEurope, kugoverwa kwemitsetse yekoniyoni (kunyanya ye13_A2 genotype) haina kukanganisa vanhu vemunharaunda yepathogen, iyo inochengetedza huwandu hwakawanda hwekusiyana nekushayikwa kwemitsara yakakurisa.
Iko kuvapo kwe13_A2 kunoonekwa pano neapo muminda ine mhando dzekutengesa mbatatisi. MuRussia, mamiriro ezvinhu ari kusimukira nenzira yakafanana. Microsatellite ongororo ye P. infestans inoparadzanisa inounganidzwa muna 2008-2011 mumatunhu gumi akasiyana echikamu cheEuropean cheRussia, yakaratidza mwero wakakura wegenotypic kusiyana uye kushomeka kwakakwana kwezviitiko pamwe neEuropean clonal mitsetse (Statsyuk et al., 10). Makore akati wandei gare gare, ongororo ya P. infestans sampuli yakaunganidzwa mudunhu reLeningrad muna2014-2013 yakaratidza misiyano yakakura pakati pavo nemajenotypes kubva mudunhu rino akaratidzwa muchidzidzo chekare. Mune zvese zvidzidzo, Western Europe genotypes haina kuwanikwa (Beketova et al., 2014; Kuznetsova et al., 2014).
Iyo yakakwira genet misiyano yeEastern Europe vanhu veP infestans uye kusavapo kwemasimba makuru emakondoni mavari anogona kunge ari nekuda kwezvikonzero zvakati wandei. Kutanga, senge kuchamhembe kweEurope, mamiriro ekunze enzvimbo dzinoonekwa anobatsira mukuumbwa kwe oospores sosi yekutanga yehutachiona (Ulanova et al., 2010; Chmielarz et al., 2014). Chechipiri, chakakosha chikamu chembatatisi dzinogadzirwa munyika idzi dzinorimwa pamapurazi madiki akazvimirira, anowanzo kukomberedzwa nemasango kana zvimwe zvipingamupinyi pakufamba kwemahara kwezvinhu zvinotapukira (Chmielarz et al., 2014). Semutemo, mbatatisi dzakasimwa pasi pemamiriro akadaro dzinenge dzisingarapwe nemakemikari, uye kusarudzwa kwemhando kunoenderana neyavo yekupedzisira blight kuramba, i.e. Iko hakuna kusarudzika kwekumanikidza uye kupokana ne metalaxyl, iyo inobvisa zvinodzivirira genotypes, senge 13_A2, yezvakanaka kupfuura mamwe magenotypes (Chmielarz et al., 2014). Chekupedzisira, nekuda kweiyo diki saizi yenzvimbo dzevhu, varidzi vadzo havawanzo kudzidzira kutenderera kwezvirimwa, kurima mbatatisi kwemakore munzvimbo imwechete, izvo zvinobatsira mukuunganidzwa kweinoculum yakasiyanasiyana (Runno-Paurson et al., 2016; Elansky , 2015; Elansky et al., 2015).
Asia
Kusvika nguva pfupi yadarika, chimiro cheP infestans vanhu muAsia chakaramba chisina kunzwisiswa. Zvaizivikanwa kuti zvinomiririrwa kunyanya ne clonal mitsetse, uye mhedzisiro yekurara pabonde pakuonekwa kwema genotypes matsva idiki kwazvo. Saka, semuenzaniso, muna 1997-1998. Muchikamu cheAsia cheRussia (Siberia neFar East), huwandu hwehutachiona hwakamiririrwa neatatu chete magenotypes ane mukurumbira weSIB-1 genotype (Elansky et al., 2001). Kuvapo kwemitsetse yeclonal pathogen yakaratidzirwa munyika dzakaita seChina, Japan, Korea, Philippines, neTaiwan (Koh et al., 1994; Chen et al., 2009). Iyo clonal mutsetse US-1 yakatonga pamusoro penzvimbo hombe yeAsia mukupera kwema90s - kutanga 2000s. dzinenge kwese kwese dzakatanga kutsiviwa nedzimwe genotypes, idzo, dzakazopa itsva. Muzviitiko zvakawanda, shanduko muchimiro uye kuumbwa kwevanhu munyika dzeAsia dzakabatana nekusimuka kwemagenotypes matsva kubva kunze. Nekudaro, muJapan, kunze kweiyo JP-3 genotype, mamwe ese maJapan genotypes akaonekwa mushure meUS-1 (JP-1, JP-2, JP-3) ane zvakawanda kana zvishoma zvakabvumidzwa mavambo ekunze (Akino et al. , 2011) ... Kune parizvino kune matatu makuru epathogen vanhu muChina, ayo ane nzvimbo yakajeka yenzvimbo; gene inoyerera pakati pevanhu ava isipo kana kushaya simba (Guo et al., 2010; Li et al., 2013b). Genotype 13_A2 yakaonekwa munharaunda yeChina mumatunhu ayo ekumaodzanyemba (Yunnan neSichuan) muna 2005-2007, uye muna 2012-1014. yakaonekwa zvakare kuchamhembe kwakadziva kumabvazuva kwenyika (Li et al., 2013b). KuIndia, 13_A2 yakaonekwa zvingangoitika panguva imwecheteyo kuChina, ingangove ine mbatatisi yembeu ine hutachiona (Chowdappa et al., 2015), uye muna 2009-2010. zvakakonzera chirwere chakakomba cheepiphytosis chekupedzisira kunaya padomasi kumaodzanyemba kwenyika, mushure mezvo chakazopararira kumatata uye muna 2014 chakakonzera kuputika kwekupedzisira kwekutsvaira muWest Bengal, izvo zvakakonzera kuparadzwa nekuzviuraya kwevarimi vazhinji vemuno (Fry, 2016 ).
Africa
Kusvikira 2008-2010. zvakarongeka zvidzidzo zveP. infestans munyika dzeAfrica hazvina kuitwa. Parizvino, vanhu vemu Africa ve P. infestans vanogona kukamurwa kuita mapoka maviri, uye kupatsanurwa uku kwakanyatsoenderana nechinhu chekuunzwa kwembeu mbatatisi kubva kuEurope.
KuNorth Africa, iyo inonyatsopinza mbatatisi yembeu kubva kuEurope, iyo A2 mating mhando inomiririrwa munzvimbo zhinji, izvo zvinopa fungidziro mukana wekubuda kwemajenotype matsva semhedzisiro yekurara pabonde (Corbière et al., 2010; Rekad et al., 2017). Uye zvakare, muAlgeria, kuvepo kwemagenotypes 13_A2, 2_A1, uye 23_A1 ine chirevo chekutanga cheavo, pamwe nekuderera zvishoma nezvishoma muhuwandu hwemagenotypes akasarudzika ekunyangarika, zvinoonekwa (Rekad et al., 2017). Kusiyana nedunhu rese, muTunisia (kunze kwekuchamhembe kwakadziva kumabvazuva kwenyika), huwandu hwehutachiona hunomiririrwa kunyanya neiyo A1 mhando yekukwikwidza (Harbaoui et al., 2014).
Iyo clonal mutsetse NA-01 wakanyanya pano. Pakati pese, chikamu chemitsetse yeclonal muhuwandu hwevanhu inongova 43%. KuEastern neSouthern Africa, uko huwandu hwembeu dzinounzwa kunze kwenyika huri kushoma zvishoma (Fry et al., 2009), P. infestans inomiririrwa nemitsara miviri chete yeA1-mhando, US-1 uye KE-1, uye yekupedzisira ichishingaira. inobvisa iyo yekutanga pamatezi (Pule et al., 2012; Njoroge et al., 2016). Parizvino, ese aya magenotypes ane nhamba inoonekwa yekusiyana kwema subclonal.
Australia
Chirevo chekutanga pamusoro pekuratidzwa kwekusviba kwekupedzisira kumatatata kuAustralia kwakadzokera muna 1907, uye epiphytotia yekutanga, inofungidzirwa kuti yakakonzerwa nemvura zhinji mumwedzi yezhizha, yakaitika muna 1909-1911. (Drenth et al., 2002). Muzhinji, zvisinei, kunonoka kunaya hakuna kukosha kwakakosha kwehupfumi hwenyika. Kupararira kwekushomeka kwekupedzisira kwekusviba, kunetswa nemamiriro ekunze anopa hunyoro hwakanyanya, hazviitike kazhinji kanopfuura kamwe pamakore mashanu kusvika kumashanu uye zvinowanikwa munharaunda yekuchamhembe kweTasmania nepakati peVictoria. Nezve izvo zvataurwa pamusoro apa, hapana zviripo zvinyorwa zvakapihwa kuti zviongororwe chimiro chevanhu veAustralia veP. Infestans. Razvino ruzivo rwuripo rwunobva muna 5-7. (Drenth et al., 1998). Sekureva kwevanyori, huwandu hweVictoria yaive clonal mutsetse US-2000, iyo zvisina kunangana nekusimbisa kutama kweiyo genotype kubva kuUnited States. Tasmanian specimens dzakarongedzwa senge mhando AU-2002, dzakasiyana nema genotypes aive aripo panguva iyoyo mune dzimwe nzvimbo dzepasi.
Zvimiro zvekusimudzirwa kwekupedzisira blight muRussia
MuEurope, inoculum yepambatiri inoonekwa sehutachiona hunounzwa nehutachiona hwembeu, maoospores akakwidibirwa muvhu, pamwe nezoosporangia inounzwa nemhepo kubva kuzvirimwa zvakasimwa kubva kune zvakawandisa tubers muminda yegore rapfuura ("vanozvipira" zvirimwa), kana pamirwi yeakaunganidzwa bhukumaki rekuchengetedza machubhu. Pakati peizvi, zvirimwa zvinokura pamirwi yemarubhi akaraswa zvinoonekwa senzvimbo inotyisa yehutachiona. ipapo huwandu hwamera tubers hunowanzo kuve hwakakosha, uye zoosporangia inogona kutakurwa kubva kwavari pamusoro penzvimbo refu. Mamwe masosi (oospores, "vanozvipira" zvirimwa) haana ngozi zvakanyanya, nekuti haisi tsika kurima zvirimwa muminda imwechete kazhinji kanopfuura kamwe pamakore 3-4. Kutapukirwa kubva kune anorwara mhodzi yematubhu zvakare kuri kushoma nekuda kweiyo yakanaka mbeu mhando yekudzora system.
Kazhinji, huwandu hwe inoculum muvanhu veEurope ishoma, uye nekudaro kukura kwedenda iri kunonoka uye kunogona kudzorwa zvinobudirira uchishandisa makemikari fungicidal gadziriro. Basa hombe mumamiriro eEuropean nderekurwisa hutachiona muchikamu apo kupararira kwemazhinji ezosposangia kubva kuzvirimwa zvine hutachiona kunotanga.
MuRussia, mamiriro acho akasiyana zvakanyanya. Zvizhinji zvezvirimwa zvembatatisi nemadomasi zvinorimwa muminda midiki yakazvimiririra; matanho ekudzivirira haatomboitwa pavari zvachose, kana mishonga yefungicidal inoitwa isina kukwana nhamba uye inotanga mushure mekuonekwa kwekunonoka kwetsvo pamusoro. Nekuda kweizvozvo, yega yega miriwo minda inoshanda sosi huru yehutachiona, kubva kune iyo zoosporangia inotakurwa nemhepo kuenda kune zvekudyara kudyara. Izvi zvinosimbiswa nekucherechedza kwedu kwakananga munzvimbo dzeMoscow, Bryansk, Kostroma, neRyazan: kukuvara kwezvirimwa mumapindu ega kunoonekwa kunyangwe kutanga kwekurapa kwefungicide kwekudyara kwekutengesa. Shure kwaizvozvo, denda muminda mikuru rinodziviswa nekushandisa fungicidal gadziriro, nepo mune yega minda kune kukurumidza kukura kwekunonoka kunaya.
Panyaya yekurapa kusiri iko kana kuti "bhajeti" kwekudyara kwekutengesa, foci yekupedzisira kwekutsvaira inoonekwa muminda; gare gare ivo vari kushingairira kusimukira, vachifukidza nzvimbo dzakakura nguva dzose (Elansky, 2015). Kutapukirwa muminda yega yega kunokanganisa madenda munzvimbo dzekutengesa. Munzvimbo dzese dzinorima mbatatisi muRussia, nzvimbo inogarwa nembatatisi mumapindu ega ega akakura zvakapetwa pane yakazara nzvimbo yeminda yevagadziri vakuru. Munzvimbo dzakadai, yakavanzika minda yemiriwo inogona kutaridzwa seyepasi rose inoculum sosi yekutengesa minda. Ngatiedzei kuziva izvo zvivakwa zvinozivikanwa neiyo genotypes yematambudziko muminda yega.
Kudyara kwembeu uye kuisa kwega kutonga kwewairi mbatatisi, mbeu yemadomasi yakawanikwa kubva kune vanogadzira vekunze vanogadzira, kurima kwenguva refu mbatatisi nematomatisi munzvimbo dzakafanana, kurapwa kusingaiti kwefungicide kana kusavapo kwavo zvachose kunotungamira kune epiphytoties dzakakomba munzvimbo dzakazvimirira, mhedzisiro kunova kuyambuka mahara, kusanganiswa uye kuumbwa kwemahospores mumapindu ega ega. Nekuda kweizvozvo, kwakanyanya kwazvo genotypic kusiyanisa kwepathogen kunoonekwa, apo chinenge chinetso chega chega chakasarudzika mune genotype yayo (Elansky et al., 2001, 2015). Kudyara mbeu mbatatisi yemhando dzakasiyana dzemageneti kunoita kuti zvisaite kuti mitsetse yeklonal inyanzvi kurwisa imwe mhando ichabuda. Iwo maratidziro akasarudzwa mune yakadai nyaya anosiyaniswa nekusiyana-siyana kwavo maererano nemhando dzakakanganiswa, mazhinji acho ane padhuze nenhamba yepamusoro yehutachiona hutachiona. Izvi zvakasiyana zvakanyanya nehurongwa hwe "clonal mitsara" inowanzoitika kuminda mikuru yemabhizimusi ezvekurima neyakaiswa mushe nzira yekudzivirira kubva kunonoka kunaya. "Clonal mitsetse" (apo mhando dzese dzekunonoka blight pathogen mumunda dzinomiririrwa neumwe kana akati wandei genotypes) dzinowanikwa kwese munyika umo kurimwa kwembatata kunoitwa chete nemapurazi makuru: USA, Netherlands, Denmark, nezvimwewo , Kune zvakare yakakwira genotypic dzakasiyana muminda yega. Pakupera kwezana ramakore rechimakumi maviri, "clonal lines" dzaive dzakapararira munzvimbo dzeAsia neFar Eastern zvikamu zveRussia (Elansky et al., 20), izvo, sezviri pachena, zvinokonzerwa nekushandiswa kwemhando imwe chete yekudyara chete mbatatisi kugadzirwa kwedu pachedu. Munguva pfupi yapfuura, mamiriro ezvinhu mumatunhu aya akatangawo kuchinja achienda mukuwedzera kweiyo genotypic kusiana kwevanhu.
Kushaikwa kwekurapwa kwakasimba ne fungicidal kugadzirira kune imwe, mhedzisiro yakanangana - hapana kuunganidzwa kweanodzivirira maronda muminda. Chokwadi, mhedzisiro yedu inoratidza kuti metalaxyl-inopesana nematraini anowanikwa zvakanyanya kushomeka zvakanyanya mumapindu ega ega pane ekudyara kwekutengesa.
Kuswedera padyo kwekudyarwa kwembatatisi uye madomasi, kwakajairika kune mapindu akazvimirira, kunobatsira kusimuka kwematambudziko pakati pezvirimwa izvi, semhedzisiro, mumakore gumi apfuura, pakati pematambudziko ari kure nemapatatisi, chiyero chematambudziko akatakura geni rekurwisa kune akasiyana Cherry tomato (T1), yaimbozivikanwa chete kune "matomati" maronda. Matinji ane iyo T1 geni mune dzakawanda kesi ane hukasha zvakanyanya kune ese mbatatisi nematomatisi.
Mumakore achangopfuura, kunonoka kunaya padomasi kwakatanga kuoneka mune dzakawanda zviitiko zvisati zvaitika pane mbatatisi. Mbeu dzematomatisi dzinogona kuzara nemaoospores muvhu kana oospores aripo mumbeu yemadomasi kana kuomerera pairi (Rubin et al., 2001). Mumakore gumi nemashanu apfuura, huwandu hwakawanda hwembeu dzakapakurwa dzakapetwa, dzinonyanya kutengwa kunze kwenyika, dzakaonekwa muzvitoro, kuti zvishandiswe nevakawanda vevagadziri vadiki. Mbeu dzinogona kuunza masisitimu ane genotypes akajairwa kumatunhu ekurima kwavo. Mune ramangwana, idzi genotypes dzinosanganisirwa muitiro wekuita bonde mune yega magadheni, izvo zvinotungamira mukubuda kweanyowani matsva genotypes.
Nekudaro, zvinogona kutaurwa kuti zvakavanzika magadheni "epasi rese arinyungudika", umo, semhedzisiro yekuchinjana kwemajini, genotypes dziripo dzinogadziriswa uye nyowani nyowani dzinoonekwa. Panguva imwecheteyo, kusarudzwa kwavo kunoitika pasi pemamiriro ezvinhu akasiyana zvakanyanya neakagadzirirwa mbatatisi mumapurazi mahombe: kusavapo kwemushonga wefungicidal, kusiyanisa kwekudyara, hukuru hwezvirimwa zvakakanganiswa nemhando dzakasiyana dzehutachiona uye hutachiona hutachiona, padyo kuna madomasi uye nightshades yemusango, kuyambuka kushingairira uye oospore kuumbwa, mukana wekuti oospores iite sechinhu chinotapukira hutachiona mugore rinotevera.
Zvese izvi zvinotungamira kune yakakwira kwazvo genotypic misiyano yevagari vemumba. Pasi pemamiriro epiphytotic, blight inononoka inopararira nekukurumidza muminda yemiriwo uye huwandu hwakawanda hwezvipembenene hunoburitswa, izvo zvinobhururuka kuenda kunzvimbo dzekutengesa padyo Nekudaro, vapinda muminda yekutengesa neiyo chaiyo sisitimu yekurima tekinoroji uye kuchengetedzwa kwemakemikari, ma spores akapinda mukati haakwanise kutanga epiphytotic mumunda, izvo zvinokonzerwa nekushayikwa kwemitsetse yeclonal inopesana nefungicides uye hunyanzvi kune dzakasiyana siyana dzakarimwa.
Imwe sosi yekutanga inoculum inogona kunge iri maronda machubhu akavharirwa mumidziyo yekutengesa. Aya ma tubers akakurwa, sekutonga, muminda ine hunyanzvi hwekurima uye yakasimba yekudzivirira makemikari. Iyo genotypes yeavo vakazvitsaura ayo akakanganisa ma tubers anochinjirwa mukusimudzira kwavo kwakasiyana siyana. Aya maronda ane ngozi zvakanyanya kukudyara kwekutengesa kupfuura inoculum inotangira kubva kune yega magadheni. Uku kufungidzira kunotsigirwawo nezvakabuda mukutsvaga kwedu. Vagari vakasarudzika kubva kuminda mikuru ine yakachengetedzwa zvakanaka kemikari chengetedzo uye yakanaka yekurima tekinoroji haina kusiyana mukukwirira kwemageneti. Kazhinji idzi dzinoverengeka clonal mitsara ine hukasha zvakanyanya.
Mitsetse kubva kuzvinhu zvinotengeswa zvembeu inogona kupinda muvanhu mumapindu emiriwo uye kubatanidzwa mune zvinoitika zvinoitika mavari. Nekudaro, mubindu remuriwo, kukwikwidza kwavo kunenge kwakadzikira zvakanyanya kupfuura mundima yekutengesa, uye munguva pfupi ivo vanozoguma kuvapo nenzira yechikwata checlonal, asi majini avo anogona kushandiswa mu "gadheni" revanhu.
Hutachiona hunowanikwa pane "vanozvipira" zvinomera uye pamirwi yemakonzerwa machubhu panguva yekukohwa haina kukodzera zvakanyanya kuRussia, nekuti Munzvimbo huru dzinokura mbatatisi dzeRussia, kwakadzika chando ivhu rinotonhora, uye zvinomera kubva mumatubhu izvo zvakadonhedza muvhu hazviwanzo kukura. Zvakare, sekuratidzira kwedu kwekuyedza, iyo yekupedzisira blight pathogen hairarame pakashata tembiricha kunyangwe pane machubhu akachengetedza kushanda kwawo. Munzvimbo isina mvura, panorimwa mbatatisi yekutanga, blight yekunonoka haiwanzoitika nekuda kwekukura uye kupisa kwekukura mwaka.
Nekudaro, izvozvi tiri kutarisa kupatsanurwa kweP infestans vanhu kupinda "mumunda" uye "gadheni" vanhu. Nekudaro, mumakore achangopfuura, maitiro akaonekwa achitungamira mukubatana uye kupindirana kwemagenotypes kubva kuvanhu ava.
Pakati pavo, mumwe anogona kucherechedza kuwedzera kwakawanda mukuverenga nekunyora kwevagadziri vadiki, kubuda kwemapakeji madiki embeu mbatatisi, kupararira kwekugadzirira fungicidal mumapakeji madiki, uye kurasikirwa kwekutya kwe "chemistry" nevanhu.
Mamiriro ezvinhu anomuka apo, nekuda kwekushanda nesimba kweumwe mutengesi, misha yese yakasimwa nembeu dzemidziyo yemhando imwechete uye inopihwa mapakeji madiki emushonga wezvipembenene. Zvinogona kufungidzirwa kuti mbatatisi dzemhando dzakasiyana dzichawanikwa pane kudyara kwekutengesa padyo.
Kune rimwe divi, mamwe makambani anotengesa mishonga inouraya zvipembenene ari kukurudzira zvirongwa zve "bhajeti" zvemakemikari. Muchiitiko ichi, huwandu hwokurapa hwakakurudzirwa hautarisirwe uye fungicides yakachipa inopihwa, uye kusimbisa hakusi kwekudzivirira kukura kwekupedzisira kwechirwere kusvika pakucheka misoro, asi pane kumwe kunonoka muepiphytoty kuitira kuti uwedzere goho. Zvirongwa zvakadaro zvinonzwisisika mune zvehupfumi kana uchirima mbatatisi kubva kune yakaderera-mbeu mbeu zvinhu, kana musimboti pasina mubvunzo wekuwana gohwo repamusoro. Nekudaro, mune iyi kesi, mukusiyana nevanhu vebindu, iyo yakakwenenzverwa magene emazamu embatatari inobatsira mukusarudzika kwemamwe marudzi ehupenyu, ayo ane njodzi kune aya akasiyana.
Kazhinji, maitiro ekuchinja kwe "gadheni" uye "munda" nzira dzekugadzira mbatatisi zvinoita sekunge isu zvine njodzi. Kudzivirira mhedzisiro yavo yakaipa, mune vese veimba uye nevezvokutengeserana, zvichave zvakafanira kudzora kuwanda kwembeu mbatatisi uye huwandu hwemanyowa anopihwa varidzi vega muzvirimwa zvidiki, uye kuteedzera zvirongwa zvekudzivirira mbatatisi uye mashandisirwo ye fungicidal kugadzirira mune yekutengesa chikamu.
Munzvimbo dzezvikamu zvakazvimiririra, kune kusimudzira kwakasimba kwete kwekupedzisira kunonoka, asiwo Alternaria. Vazhinji varidzi vezvikamu zvakavanzika havatore matanho akakosha ekudzivirira kurwisa Alternaria, vachikanganisa kuvandudzwa kweAltharia yekumomesa kwepamusoro kwemisoro kana kusimudzira kwekupedzisira kwechirwere. Naizvozvo, nekuvandudzwa kukuru kweExtaria pamhando dzinobatika, dzimba dzemumba dzinogona kushanda sosi yeinoculum yekudyara kwekutengesa.
Nzira dzekusiyana
Kuchinja maitiro
Sezvo kuitika kweshanduko kuri kweimwe nzira yekuenderera ine yakaderera frequency, iko kuitika kweshanduko kune chero locus kunoenderana nekuwanda kwekuchinja kweiyi locus uye saizi yehuwandu. Paunenge uchidzidza kuwanda kweshanduko dze P. infestans mafashama, huwandu hwemakoloni hwakakura pane inosarudzika nhepfenyuro midhiya mushure mekurapwa nemakemikari kana epanyama mutagen inowanzo gadziriswa. Sezvinoonekwa kubva pane data rakaburitswa muTebhenekeri 8, iyo shanduko frequency yeiyo imwechete strain pane dzakasiyana loci inogona kusiyana nematanho mazhinji ehukuru. Iyo yakakwira frequency yekuchinja mukushora metalaxyl inogona kunge iri imwe yezvikonzero zvekuunganidza kwehutachiona husingawirirani nahwo mune zvakasikwa.
Kuwanda kwekuchinja kwakangoerekana kwaitika kana kukonzereswa, kunoverengerwa pahwaro hwekuyedzwa kwerabhoritari, hakuwanzoenderana nemaitiro anoitika munharaunda dzevanhu, nekuda kwezvikonzero zvinotevera:
1. Mune inynchronous nyukireya fissions, hazvigoneke kufungidzira kuwanda kweshanduko pachizvarwa chimwe chenyukireya. Naizvozvo, kuyedza kwakawanda kunopa ruzivo chete zvakanangana nezve kuwanda kweshanduko, pasina kusiyanisa pakati pezviitiko zviviri zvekuchinja uye chiitiko chimwe chinotevera mitosis.
2. Nhanho-nhanho shanduko inowanzo deredza muyero weiyo genome, saka, pamwe nekutora chivakwa chitsva, hutano hwese hwechisimba hunoderera. Mazhinji ezviyero zvakawanikwa zvakaedzwa zvine hukasha hwakadzora uye hazvina kunyorwa muvanhu venyama. Nekudaro, iko kuwirirana kwakaringana pakati pechiyero chekuramba kweP. Infestans mutant kune phenylamide fungicides uye mwero wekukura pane yakasikwa nharaunda yaive paavhareji (-0,62), uye kuramba kwefungicides uye hasha pamashizha embatatisi (-0,65) et al., 1993), iyo inoratidza kusimba kweakachinja mutants. Shanduko dzekuramba kune dimethomorph dzakaperekedzwawo nekuderera kwakasimba mukubudirira (Bagirova et al., 2001).
3. Zvizhinji zvezviitiko zvenguva dzose uye zvakakonzera zvinochinja uye hazvizviratidze phenotypically mukuyedza, asi zvinomirira dura rakavanzika rekusiyana pakati pevanhu. Matant mutinosara akasarudzika muongororo yeRabhoritari inotakura shanduko dzakakura kana semi-hukuru (Kulish naDyakov, 1979). Sezvineiwo, diploidy yenyukireya inotsanangura kuyedza kusingabudirire kwekuwana mutants pasi pesimba reUV irradiation iyo ine huturu pamarimi akambomira (McKee, 1969). Zvinoenderana nekuverenga kwemunyori, shanduko dzakadai dzinogona kuitika nesvitsa isingasviki 1: 500000. Iko kushandurwa kwekuchinja kwakawanda kune imwe homozygous, phenotypically inoratidzwa mamiriro anogona kuitika nekuda kwekuita bonde kana asexual recombination (ona pazasi). Nekudaro, kunyangwe mune ino kesi, shanduko inogona kuvharwa neakakura alleles emusango-mhando nuclei mune cenotic (multinucleated) mycelium uye phenotypically yakagadziriswa chete panguva yekuumbwa kwemononuclear zoospores.
Tafura 8. Kakawanda kweP. Infestans shanduko kune kukura-kutadzisa zvinhu pasi pechiito che nitrosomethylurea (Dolgova, Dyakov, 1986; Bagirova et al., 2001)
Mukomboni | Kuchinja frequency |
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 |
Daaconil | <4 x 10-8 |
Dimethomorph | 6,3 x 10-7 |
Metalaxil | 6,9 x 10-6 |
Kukura kwevagari vevanhu zvakare kunoita chinhanho chakakosha mukumuka kweshanduko dzinongoitika dzega. Mune huwandu hwakakura kwazvo, mune huwandu hwemasero N> 1 / a, uko chiyero cheshanduko, shanduko inomira kuva chinhu chisina kujairika (Kvitko, 1974).
Masvomhu anoratidza kuti neavhareji infestation yemunda wembatatisi (makumi matatu nematatu pane chimwe chirimwa), 35x8 spores dzinoumbwa zuva nezuva pahekita imwe (Dyakov naSuprun, 1012). Sezvineiwo, huwandu hwakadai hune shanduko dzese dzinobvumidzwa nerudzi rwekuchinjana pane yega locus. Kunyangwe shanduko isingawanzoitika, inoitika nekuwanda kwe1984-10, ichawanikwa nevanhu chiuru kubva pamamirioni vanogara pahekita imwe yemunda wembatatisi. Nezve shanduko dzinoitika nehurefu hwepamusoro (semuenzaniso, 9-10), mune huwandu hwevanhu, shanduko dzakasiyana siyana dzakabatana dzinogona kuitika zuva nezuva (panguva imwe chete panzvimbo mbiri loci, i.e. maitiro ekuchinja anozotsiva zvekare.
Kutama
Kuna P. infestans, maviri akakosha marudzi ekufambisa anozivikanwa: kuvhara madaro (mukati memunda wembatatisi kana minda yakavakidzana) nekuparadzira zoosporangia nemhepo yemhepo kana mvura yekunaya, uye kumatunhu marefu - nekudyara tubers kana kutakurwa michero yemadomasi. Nzira yekutanga inovimbisa kuwedzera kwechikonzero chechirwere, chechipiri - kugadzirwa kwenzvimbo nyowani munzvimbo dziri kure kubva kupuraimari.
Kupararira kwehutachiona nemadomasi machubhu nemichero hazvisi izvo chete zvinobatsira mukubuda kwechirwere munzvimbo nyowani, asi zvakare ndiyo honzeri huru yekusiyana kwemajini muhuwandu hwevanhu. Munharaunda yeMoscow, mbatatisi dzinorimwa, dzinounzwa kubva kumatunhu akasiyana eRussia neWestern Europe. Michero yetamato inounzwa kubva kumatunhu ekumaodzanyemba eRussia (Astrakhan Dunhu, Krasnodar Territory, North Caucasus). Mbeu dzematomatisi, dzinogona zvakare kushanda senzvimbo dzehutachiona (Rubin et al., 2001), dzinotengwawo kubva kumatunhu ekumaodzanyemba eRussia, China, nyika dzeEurope nedzimwe nyika.
Zvinoenderana nekuverenga kwakaitwa naE.Mayr (1974), shanduko yemunhu munharaunda inokonzereswa neshanduko haidarike kupfuura gumi kusvika gumi neshanu pasisitimu, nepo mune vanhu vakavhurika, kuchinjana nekuda kwekuyerera kwemajeneti ingangoita 10-5 - 10 -3.
Kufamba muhutachiona hune hutachiona kunokonzera kupinda kweP infestans muEurope, ichipararira kumatunhu ese epasi panorimwa mbatatisi; zvakakonzera shanduko yakakomba kwazvo yehuwandu. Kunonoka kusvibiswa pamatatata kwakaonekwa munharaunda yeRussia Humambo panguva imwe chete nekuonekwa kwayo kuWestern Europe.
Sezvo chirwere ichi chakatanga kucherechedzwa muna 1846-1847 muBaltic States uye chete mumakore akatevera akapararira muBelarus neNorth-West matunhu eRussia, mavambo ayo ekuMadokero eEurope ari pachena. Iyo yekutanga sosi yekupedzisira kunonoka muNyika Yekare haina kunyatsooneka. Fungidziro yakagadzirwa naFry et al. (Fry et al., 1992; Fry, Goodwin, 1995, Goodwin et al., 1994) inoratidza kuti utachiona hwakatanga kupinda muNorth America kubva kuMexico, uko kwaipararira pamusoro pezvirimwa, ndokuzotakurwa. kuWestern Europe (muonde. 7).
Somugumisiro wekudzokororwa kwakadzokororwa (kaviri mhedzisiro ye "bhodhoro"), maonesheni mamwe akasvika kuEurope, ayo vana akakonzera denda munharaunda yese yeiyo Nyika Dzekare, uko kurimwa mbatatisi. Seumbowo hweiyi fungidziro, vanyori vanotaura, chekutanga, kuitika kwese kwese kwemhando imwechete yekukwirisa (A1) uye, chechipiri, homogeneity yeiyo genotypes yemhando dzakadzidzwa kubva kumatunhu akasiyana (ese ari maringe nemamorekera mamaki, kusanganisira 2 isozyme loci, DNA mafingapurindi, uye chimiro cheMitochondrial DNA chakafanana uye chinoenderana necone US-1 inotsanangurwa muUSA). Nekudaro, imwe dhata inosimudza kusahadzika pamusoro peanenge zvimwe zvezvipo zveiyo yakataurwa fungidziro. Kuongororwa kweP. Infestans mitochondrial DNA yakasarudzika kubva kune herbarium mbatatisi sampu yakatapukirwa panguva yekutanga epiphytotic yemakore ekuma40 yakaratidza kuti dzakasiyana muchimiro cheMitochondrial DNA kubva kudombo reUS-1, iro, nekudaro, raive risiri iro chete sosi yekutapukira. muEurope (Ristaino and said, 2001).
Mamiriro ezvinhu neanonoka blight akawedzera zvakare muma80s ezana ramakore XX. Shanduko dzinotevera dzaitika:
1) Avhareji hukasha hwehuwandu hwakawedzera, izvo zvakatungamira, kunyanya, mukupararira kwakapararira kwenzira inokuvadza zvakanyanya yekupedzisira kwekukuvara - kukuvara kune petioles uye madzinde.
2) Kwakave nekuchinja munguva yekuonekwa kwekupedzisira kwekukanganisa pamatatata - kubva pakupera kwaChikunguru kusvika kutanga kwaChikunguru uye kunyangwe kusvika pakupera kwaJune.
3) Iyo A2 yekukwirisa mhando, iyo yaimbove isipo muNyika Yekare, yave kuwanikwa kwese kwese.
Shanduko dzakatungamirwa nezviitiko zviviri: kushandiswa kukuru kweiyo fungicide metalaxyl nyowani (Schwinn uye Staub, 1980) uye kubuda kweMexico semutengesi wepasi rose wembatatisi (Niederhauser, 1993). Zvinoenderana neizvi, zvikonzero zviviri zvekushandurwa kwevanhu zvakaiswa kumberi - shanduko yerudzi rwekusangana pasi pesimba re metalaxyl (Ko, 1994) uye kuunzwa kukuru kwematambudziko matsva ane hutachiona hutachiona kubva kuMexico (Fry naGoodwin, 1995). Kunyangwe kuchinjika kwemhando dzekubata pasi pesimba re metalaxyl kwakawanikwa kwete naKo chete, asiwo mumabasa akaitwa murabhoritari yeMoscow State University (Savenkova, Chherepennicova-Anikina, 2002), yechipiri fungidziro iri nani. Pamwe pamwe nekuonekwa kwerudzi rwechipiri rwekusvirana, shanduko dzakakomba dzakaitika mumagenotypes eRussia P. infestans strains, kusanganisira mune asina kwazvakarerekera majini (isozyme neRFLP loci), pamwe nemamiriro eDNA yemitochondrial. Iyo yakaoma yeshanduko iyi haigone kutsanangurwa nekuita kwemetalaxyl; pachinzvimbo, kwaive nekutakura kukuru kwemarudzi matsva kubva kuMexico, ayo, achinyanya kuita hasha (Kato et al., 1997), yakasiya nzvimbo dzekare (US-1) , ichiva nesimba muhuwandu hwevanhu. Shanduko mukuumbwa kwevanhu veEuropean yakaitika munguva pfupi - kubva 1980 kusvika 1985 (Fry et al., 1992). Panharaunda yeyaimbova USSR, "matsva matsva" akawanikwa muunganidzwa kubva kuEstonia muna 1985, ndiko kuti, pakutanga kwePoland neGerman (Goodwin et al., 1994). Kekupedzisira "dambudziko rekare reUS-1" muRussia rakasarudzika kubva kumatamato ane hutachiona mudunhu reMoscow muna 1993 (Dolgova et al., 1997). Zvakare muFrance, maronda "ekare" akawanikwa mukudyara kwemadomasi kusvika pakutanga kwema90, ndiko kuti, mushure mekunge vanyangarika pamatatata kare (Leberton naAndrivon, 1998). Shanduko mu P. infestans matambudziko akakanganisa maitiro mazhinji, kusanganisira aya akakosha zvakanyanya, uye akawedzera kuomarara kwekunonoka kwekupedzisira.
Kuita zvepabonde
Kuti kurara kwepabonde kubatsire mukusiyana, zvakakosha, kutanga, kuvapo kwemhando mbiri dzekukwirana muhuwandu hwehuwandu hwepedyo ne1: 1, uye, chechipiri, kuvapo kwekutanga kwehuwandu hwevanhu.
Chiyero chemhando dzekubatana chinosiyana zvakanyanya muvanhu vakasiyana uye kunyangwe mumakore akasiyana muhuwandu hwakaenzana (Tafura 9,10, 90). Izvo zvikonzero zvekuchinja kwakakomba kwakadaro mukuwanda kwemhando dzekukwirana muhuwandu hwevanhu (se, semuenzaniso, muRussia kana muIsrael mukutanga kwema 2002s ezana ramakore rapfuura) hazvizivikanwe, asi zvinofungidzirwa kuti izvi zvinokonzerwa nekuunzwa kwezvimwe. makwikwi emakonesheni (Cohen, XNUMX).
Imwe isina kunangana data inoratidza iyo nzira yekuita bonde mune mamwe makore uye mune mamwe matunhu:
1) Zvidzidzo zvevanhu vanobva mudunhu reMoscow zvakaratidza kuti muvanhu gumi nematanhatu umo mugove weA13 mhando yekusangana yaive isingasviki gumi, huwandu hwese hwemajini, hwakaverengerwa matatu isozyme loci, yaive 2, uye mune gumi nemana vanhu mune mugove yeA10 yakapfuura 0,08%, kusiyana kwemajini kwakapetwa zvakapetwa kaviri (14) (Elansky et al., 2). Nekudaro, iyo inokwidziridza mukana wekusangana pabonde, ndiko kuwedzera kwakasiyana kwemajini evanhu.
2) Hukama pakati pechiyero chemhando dzekubatana muhuwandu hwevanhu uye kusimba kwe oospore kuumbwa kwakaonekwa muIsrael (Cohen et al., 1997) uye muNetherlands
(Flier et al., 2004). Zvidzidzo zvedu zvakaratidza kuti muhuwandu hwevanhu mune yakasarudzika nemhando yeA2 yekukwirisa yaive 62, 17, 9, uye 6%, oospores yakawanikwa mu78, 50, 30, uye 15% yemashizha akaongororwa embatatisi (ane maviri kana anopfuura mavara), zvichiteerana.
Samples ane maviri kana mamwe mavara anonyanya kuwanda ane oospores pane masampuli ane 2 nzvimbo (1 uye 32% yemasampuli, zvichiteerana) (Apryshko et al., 14).
Oospores yaive yakajairika mumashizha epakati nepazasi eiyo mbatatisi chirimwa (Mytsa et al., 2015; Elansky et al., 2016).
3) Mune mamwe matunhu, akasarudzika genotypes akawanikwa, iko kuitika kunosanganisirwa nekugadzirisa bonde. Nekudaro, muPoland muna 1989 uye muFrance muna1990, inokanganisa homozygous yeglucose-6-
phosphate isomerase (GPI 90/90). Sezvo zvakangowanikwa chete 10/90 heterozygotes akasangana kwemakore gumi, homozygosity inonzi inoratidzirwa pakuita bonde (Sujkowski et al., 100). MuColombia (USA), inoparadzanisa kusanganisa A1994 neGPI 2/100 uye A110 neGPI 1/100 zvakajairika, zvisinei, pakupera kwemwaka wa100 (Nyamavhuvhu 1994 naSeptember 16), maronda ane recombinant genotypes (A9 GPI 1 / 100 uye A110 GPI 2/100) (Miller et al., 100).
4) Mune vamwe vanhu vanobva kuPoland (Sujkowski et al., 1994) uye North Caucasus (Amatkhanova et al., 2004), kugoverwa kweminwe yeminwe DNA loci uye allozyme protein loci kunoenderana nekuparadzirwa kweHardy-Weinberg, iyo inoratidza
nezve yakakwenenzverwa chikamu chemupiro wekufunga zvebonde mukusiyana kwevanhu. Mune mamwe matunhu eRussia, hapana tsamba kune kugoverwa kweHardy-Weinberg muhuwandu hwevanhu yakawanikwa, asi kuvepo kwekubatana disequilibrium kwakaratidzwa, zvichiratidza hukuru hwekubereka kubereka (Elansky et al., 1999).
5) Kusiyana kweGenetic (GST) pakati pematambudziko ane akasiyana mating mhando (A1 neA2) yaive yakaderera pane pakati pevanhu vakasiyana (Sujkowski et al., 1994), iyo inoratidza zvisina kunangana michinjiko yepabonde.
Panguva imwecheteyo, mupiro wekufunga nezvebonde kune huwandu hwevanhu haugone kukwirira zvakanyanya. Uyu mupiro wakaverengerwa vanhu venzvimbo yeMoscow (Elansky et al., 1999). Zvinoenderana nekuverenga kwaLewontin (1979) "kugadzirisazve, uko kunogona kuburitsa misiyano mitsva kubva kune loci mbiri ine frequency isingapfuure chigadzirwa cheheterozygosities yavo, inova inoshanda chete kana hunhu hweheterozygosity kune ese alleles hwatove hwakakwirira."
Nechiyero chemhando mbiri dzekubatanidza, icho chinowanzoitika mudunhu reMoscow, chakaenzana na4: 1, kuwanda kwekudzokorora kuchave 0,25. Iko mukana wekuyambuka masisitimu uchave heterozygous kune vaviri vevatatu vakadzidza isozyme loci muvanhu vakadzidza yaive 0,01 (2 maronda kunze kwe177). Naizvozvo, mukana wekuitika kwemaviri heterozygotes semhedzisiro yekudzokorora haufanire kupfuudza chigadzirwa chavo chakawedzerwa nemukana wekuyambuka (0,25x0,02x0,02) = 10-4, i.e. zvepabonde zvinodzokororwa hazviwanzo kuwira mune zvakaenzanisirwa sampuro yematambudziko. Aya maverengera akaitirwa vanhu vanobva mudunhu reMoscow, ayo anozivikanwa nekusiyana kwakanyanya. Mune monomorphic vanhu vakaita seSiberia, iyo bonde, kunyangwe ikaitika muvanhu vamwe nevamwe, haigone kukanganisa kwavo kwakasiyana.
Uye zvakare, P. infestans inoratidzirwa neyakajairika chromosome kusarongeka mune meiosis, iyo inotungamira kune aneuploidy (Carter et al., 1999). Kukanganisa kwakadai kunoderedza kubereka kwemahybrids.
Parasexual recombination, mitotic gene kutendeuka
Mukuyedza kwekukamurwa kweP. Infestans mabhureki nekuchinja kwekurwisa kune akasiyana kukura inhibitors, kubuda kwemisolates inopesana neese inhibitors kwakawanikwa (Shattock naShaw, 1975; Dyakov, Kuzovnikova, 1974; Kulish, Dyakov,
1979). Mitsetse inopesana nemaviri ekukura inhibitors yakamuka nekuda kweheterokaryotization yeiyo mycelium, uye mune iyi kesi, yakanamatira panguva yekubereka ne mononuclear zoospores (Judelson, Ge Yang, 1998), kana vasina kunamatira mune monozoosporous vana, nekuti vaive netetraploid ( sezvo ekutanga akapatsanurwa ari diploid) nuclei (K, 1979). Heterozygous diploids yakaparadzaniswa pane yakaderera kwazvo frequency nekuda kwe haploidization, chromosome nondisjunction, uye mitotic kuyambuka pamusoro (Poedinok et al., 1982). Kuwanda kweaya maitiro kunogona kuwedzerwa nerubatsiro rweimwe mhedzisiro pane heterozygous diploids (semuenzaniso, UV irradiation yezvinomera zvinomera).
Kunyangwe kuumbwa kwemavara emahombekombe ane mbiri yekupokana kunoitika kwete chete mu vitro, asi zvakare muhutatari tubers inotapukirwa nemusanganiswa wezvinhu zvinoshanduka (Kulish et al., 1978), zviri nani kuongorora basa repasexual kudzokorora muchizvarwa che genotypes nyowani muhuwandu hwevanhu. Iko kuwanda kwema segregants kuumbwa nekuda kwe haploidization, nondisjunction yemakromosomes uye mitotic kuyambuka pamusoro pasina akakosha mhedzisiro haina basa (isingasviki 10-3).
Iko kubuda kwe homozygous segregants ye heterozygous strains inogona kuve yakavakirwa pane ese maviri mitotic kuyambuka pamusoro uye mitotic gene kutendeuka, iyo iri muP. Sojae inoitika ine frequency ye3 x 10-2 kusvika 5 x 10-5 pa locus, zvinoenderana neshungu ( Chamnanpunt et al., 2001).
Kunyangwe iko kuwanda kwezviitiko zveheterokaryons uye heterozygous diploids yakazove yakatarisira kumusoro (kusvika makumi ezana muzana), maitiro aya anoitika chete kana tsika dzakachinjika dzakawanikwa kubva padambudziko rimwe chete dzakapetwa. Paunenge uchishandisa akasiyana masisitimu akaparadzaniswa nemasikirwo, heterokaryotization haiitike (kana inoitika neakadzika kwazvo frequency) nekuda kwekuvapo kwekusaenderana kwezvirimwa (Poedinok naDyakov, 1981; Anikina et al., 1997b; Cherepennikova-Anikina et al., 2002 ). Nekudaro, iro basa repasexual recombination rinogona kungodzikiswa kuita intraclonal recombination mu heterozygous nuclei uye shanduko yemunhu majini kune imwe homozygous nyika isina kuita bonde. Maitiro aya anogona kuve eepidemiological kukosha mumatambudziko ane akawandisa kana epakati-akakosha fungicide kuramba kushanduka. Kuchinja kwaro kuenda kune imwe homozygous nyika nekuda kweiyo parasexual maitiro kunowedzera kushorwa kweiye anotakura shanduko (Dolgova, Dyakov, 1986).
Kujekesa kwemajini
Heterothallic marudzi Phytophthora anokwanisa kuberekana kuita hybrid oospores (ona Vorob'eva naGridnev, 1983; Sansome et al., 1991; Veld et al., 1998). Iyo yakasanganiswa yakasanganiswa yemhando mbiri dzePhytophthora yaive nehasha zvekuti yakauraya zviuru zvealders muUK (Brasier et al., 1999). P. infestans inogona kuitika nemamwe marudzi eiyo genus (P. erythroseptica, P. nicotianae, P. Cactorum, nezvimwewo) pane zvakajairika zvinomera zvinodyarwa uye muvhu, asi pane ruzivo rudiki mumabhuku nezve mukana weiyo interspecific hybrids . Pasi pemamiriro erabhoritari, mahybrids akawanikwa pakati paP infestans naP P. Mirabilis (Goodwin naFry, 1994).
Tafura 9. Chikamu che P. infestans chinosangana neA2 mhando yekusangana munyika dzakasiyana dzepasi kubva panguva ya1990 kusvika 2000 (zvinoenderana nedata rezvakavhurwa zvinyorwa nemasaiti www.euroblight.net, www.eucablight.org)
nyika | 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) | ||||||||||
England | 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) | ||||||||||
Mawodzanyemba. Ayarendi | 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) |
Tafura 10. Chiyero cheP. Infestans maficha ane A2 mating mhando mune dzakasiyana nyika dzepasi munguva kubva 2000 kusvika 2011.
nyika | 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) | |||||||
Germany | 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) | |||||
England | 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 | |||||||
Mawodzanyemba. Ayarendi | 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) | ||||||||
Vietnam | 0 (294, 2003-04) | ||||||||||
Uganda | 0 (8) |
Simba reiyo genotypic kuumbwa kwevanhu
Shanduko mune genotypic kuumbwa kwe P. infestans huwandu hwevanhu hunogona kuitika pasi pesimba rekutama kwema clones matsva kubva kune mamwe matunhu, maitiro ekurima (shanduko yemhando dzakasiyana, kushandiswa kwefungicides) uye mamiriro ekunze. Kunze kwekunze kunokanganisa matombo akasiyanasiyana pamatanho akasiyana ehupenyu kutenderera, nekudaro, vanhu pagore vanoona shanduko yedenderedzwa mumatanho emajeni anoenderana nesarudzo, nekuda kweshanduko mune yakanyanya kuita yejene kutsauka uye kusarudzwa.
Kufuridzira kwemhando dzakasiyana
Mitsva mitsva ine mageneti anoshanda ekumira akatwasuka (R-mageneti) isimba rakasarudzika rinosarudza macones ane ekuwedzera ehutachiona magene muP infestans vanhu. Mukushayikwa kwekusapokana kwakasimba mumhando dzembatatisi, iyo inotitadzisa kukura kwehutachiona hwevanhu, maitiro ekutsiva makona akakurisa muhuwandu anoitika nekukurumidza. Saka, mushure mekupararira mudunhu reMoscow yeDomodedovsky dzakasiyana, idzo dzine R3 yekudzivirira geni, kuwanda kwemakondoni ane huturu hweichi chakasiyana kuwedzera kubva pa0,2 kusvika ku0,82 mugore rimwe chete (Dyakov, Derevjagina, 2000).
Nekudaro, shanduko mune kuwanda kweiyo virulence genes (pathotypes) muhuwandu hwevanhu inoitika kwete kwete chete pasi pesimba remhando dzakarimwa dzembatatisi. Semuenzaniso, muBelarus kusvika 1977, maonesheni ane hutachiona hutachiona 1 ne4 hwakakunda, izvo zvakakonzerwa nekurimwa kwemarudzi embatatisi ane majekiseni ekurwisa R1 neR4 (Dorozhkin, Belskaya, 1979). Nekudaro, pakupera kwema 70s ezana ramakore reXXX, maonesheni akaonekwa aine hutachiona hwakasiyana uye kusanganiswa kwawo, uye mageneti ekuwedzera ekudzivirira haana kumbobvira ashandiswa mukuzadza mbatatisi (mamwe majini eropa) (Ivanyuk et al., 2002). Chikonzero chekuonekwa kwemakona akadaro, sezviri pachena, imhaka yekutamira kuEurope kwezvinhu zvinotapukira kubva kuMexico zvine mbatatisi tubers. Kumba, aya maonesheni akagadzirwa kwete chete pamatatata akarimwa, asiwo pamhando dzesango dzinotakura akasiyana majeni ekurwisa, saka kusanganiswa kwemajusi mazhinji ehutachiona mujenome kwaidiwa kuti urarame mumamiriro iwayo.
Kana zviri zvemhando dzakasiyana nonspecific kuramba, ivo, nekudzora mwero wekuberekana kwepathogen, vanononoka kushanduka kwehuwandu hwayo, iyo, sezvatotaurwa, ibasa rehuwandu. Sezvo hukasha huri polygenic, maonesheni ane huwandu hukuru hwemajini e "hasha" anounganidza nekukurumidza, inowedzera huwandu hwevanhu. Naizvozvo, nhangemutange ine hukasha haizi chigadzirwa chekushandurwa kumhando dzakarimwa nekushorwa kusingaite, asi zvinopesana, zvinowanzoonekwa mukudyara kwemhando dzinotapukira idzo dzinounganidza hutachiona hwehutachiona.
Nekudaro, muRussia, vanhu vane hukasha zvakanyanya veP.Pestestans vakawanikwa munzvimbo dzegore epiphytoties (vanhu vanobva kuSakhalin, Leningrad, neBryansk matunhu). Hasha dzevanhu ava dzakaratidza kuve dzakakwirira kupfuura dzevaMexico (Filippov et al., 2004).
Uye zvakare, mashoma oospores anoumbwa mumashizha emhando dzinodzivirira pane ayo anogona kubatwa (Hanson naShattock, 1998), ndiko kuti, kushorwa kusingaenzaniswi kwemhando iyi kunoderedza kugona kwehutachiona hwekudzokorora uye mukana weimwe nzira dzechando.
Kufuridzira fungicides
Fungicides haingoderedze chete nhamba yephytopathogenic fungi, i.e. inokanganisa hunhu hwehuwandu hwevanhu vavo, asi inogona zvakare kuchinja mafambisirwo eanogumbuka ega, i.e. inokonzeresa chimiro chehunhu hwevanhu. Pakati pezvakakosha zviratidzo zvehuwandu hwevanhu hunoshanduka pasi pesimba refungicides zvinotevera: shanduko mukushora fungicides, shanduko mune hukasha uye huturu, uye shanduko muhurongwa hwekubereka.
Kufuridzirwa kwema fungicides pakurwisa uye hukasha hwevanhu
Chiyero chekufurira kwakadai chakatemwa, kutanga pane zvese, nemhando yefungicide inoshandiswa, iyo inogona kuve yakakamurwa kuita polysite, oligosite uye monosite.
Boka rekutanga rinosanganisira akawanda ekubata fungicides. Kuramba kwavari (kana zvichibvira zvachose) kunodzorwa nenhamba huru yemagen'a asinganyatso kuratidza. Izvi zvivakwa zvinotarisisa kusavapo kweshanduko dzinooneka mukushomeka kwevagari mushure mekurapwa nema fungicides (kunyangwe mune kumwe kuyedza kumwe kuwedzera kwekushomeka kwakawanikwa). Iyo fungal huwandu hwakachengetedzwa mushure mekumwaya neyekubatana fungicides ine maviri mapoka ematambudziko:
1) Mitsetse inochengetwa munzvimbo dzekudyara dzisina kurapwa nemushonga. Sezvo pakanga pasina kusangana neiyo fungicide, hukasha uye kuramba kweaya maratidziro hakuchinje.
2) Mitsetse inosangana neiyo fungicide, iko kusunganidzwa kwaro panzvimbo dzekusangana kwaive pasi pekuuraya. Sezvambotaurwa pamusoro apa, kuramba kwechikamu chino chevanhu hakuchinji, zvisinei, nekuda kwekukanganisa kwakashata kwefungicide kunyangwe mukutetepa kwakadzika pametabolism yesero refungal, hutano hwakazara uye chinhu chayo chepasita, hasha, kuderera (Derevyagina naDyakov, 1990).
Nekudaro, kunyangwe chikamu chevanhu vasina kufa, chakafumurwa kusangana nefungicide, chine hukasha husina kusimba uye hachigone kuve sosi yeepiphytotic. Naizvozvo, kunyatsogadzirisa, uko kunoderedza kuwanda kwehuwandu hwehuwandu husingawirirane nefungicide, ndiwo mamiriro ekubudirira kwezviyero zvekudzivirira. Kuramba kune oligosite fungicides inodzorwa neakawanda ekuwedzera majini.
Kuchinja kwega kwega geni kunotungamira mukumwe kuramba mukushorwa, uye huwandu hwese hwekushomeka hwakakonzerwa nekuwedzerwa kweshanduko dzakadai. Naizvozvo, kuwedzera kwekushomeka kunoitika nhanho. Muenzaniso wekuwedzera kwekutsika kwekusagadzikana kushanduka mukushora fungicide dimethomorph, iyo inoshandiswa zvakanyanya kuchengetedza mbatatisi kubva kunonoka kunaya. Dimethomorph kuramba ndeye polygenic uye inowedzera. A-nhanho-shanduko inowedzera zvishoma kuramba.
Shanduko yega yega inotevera inoderedza saizi yakatarwa uye, nekudaro, kuwanda kweshanduko inotevera (Bagirova et al., 2001). Iko kuwedzera kwevhareji kuramba kwehuwandu hwevanhu mushure mekurapwa kwakawanda neiyo oligosite fungicide kunoitika zvishoma nezvishoma uye zvishoma nezvishoma. Chiyero cheichi chiitiko chinotemerwa nezvakaderera zvinhu zvitatu: kuwanda kwekuchinja kwemajini ekudzivirira, kuenzana kwekudzivisa majini pakusimba.
Kuwanda kwezviitiko zvega kwega shanduko inotevera kwakaderera pane yapfuura; nekudaro, maitiro ane damping hunhu (Bagirova et al., 2001). Nekudaro, kana maitiro ekudzokorora (ezvepabonde kana eparasexual) akaitika muhuwandu, zvino zvinokwanisika kusanganisa shanduko dzakasiyana dzevabereki mune yakasanganiswa strain uye nekukurumidza maitiro acho Naizvozvo, panmix vanhu vanowana kuramba nekukurumidza kupfuura maagamic, uye mune yekupedzisira, vanhu vasina zvinomera zvinopesana nekukasira kupfuura vanhu vakapatsanurwa nezvipingamupinyi zvakadaro. Panyaya iyi, kuvepo kwematambudziko muhuwandu hwevanhu hwakasiyana mumhando dzekusvitsa kunokasira maitiro ekuwana kushorwa nemafuta eoligosite.
Chechipiri uye chechitatu zvinhu hazvibatsire kukurumidza kuunganidzwa kwetimethomorph-inomiririra maficha muhuwandu hwevanhu. Kuchinja kwega kwega kunotevera kunopeta nekupokana, izvo zvisina basa, uye panguva imwechete kunoderedza zvese huwandu hwekukura munzvimbo yekunyepedzera uye hukasha (Bagirova et al., 2001; Danda, Kirk, 2004). Zvichida, saka, pakati pematambudziko echisikirwo eP. infestans, kunyangwe iwo akaunganidzwa kubva kumiti yekudyara mbatatisi inorapwa nedhimethomorph, pane angangoita asingagadzike.
Huwandu hwevanhu hunorapwa ne oligosite fungicide huchavewo nemapoka maviri emhando: iwo anga asati asangana neiyo fungicide, uye nekudaro haana kuchinja hunhu hwekutanga (kana zvinodzivirira zvikawanikwa pakati peboka iri, hazvizowanda nekuda kune hukasha hwepamusoro uye kukwikwidza kwematambudziko akaomarara), uye maratidziro ari kusangana neyakaipisisa kuwanda kweiyo fungicide. Icho chiri pakati peyekupedzisira kuti kuunganidzwa kweanodzivirira marara kunokwanisika, nekuti pano ivo vane zvakanakira kupfuura dzakaomarara.
Naizvozvo, kana uchishandisa oligosite fungicides, haina kunyatso chengetedza kugadzirisa kwakakosha sekukwirisa kwemushonga, kakati wandei kupfuura muyero unouraya, nekuti ne stepwise mutagenesis, iko kwekutanga kuramba kwehutachiona hwehutachiona hwakaderera.
Chekupedzisira, shanduko mukushora mononic fungicides dzinonyanya kuratidza, ndokuti, kumwe kusanduka kunogona kuzivisa yakakwira mwero wekudzivirira kusvika pakupera kurasikirwa kwekunzwa. Naizvozvo, kuwedzera kwekushomeka kwevanhu kunoitika nekukurumidza.
Muenzaniso weaya fungicides ndeye phenylamides, kusanganisira iyo yakajairika fungicide, metalaxyl. Shanduko dzekumirisana nayo dzinomuka neyakawandisa frequency, uye mwero wekurwisa mumutantiki wakanyanya kwazvo - inodarika kunetseka kwakadzvanywa nechinhu chiuru kana kudarika (Derevyagina et al., 1993). Kunyangwe hazvo huwandu hwekukura uye hukasha hwehutachiona husinga gadzikike zvichiderera pakatarisana nerufu rwehutachiona hunobatika kubva kune systemic fungicide, huwandu hwevanhu vanopikisa huri kukura nekukurumidza uye, mukufanana, hukasha hwayo huri kuwedzera. Naizvozvo, mushure memakore akati wandei ekushandisa fungicide, hukasha hwemishonga inodzivirira haugone chete kuenzana nehasha dzevanoziva, asi zvakare kuupfuura (Derevyagina naDyakov, 1992).
Mhedzisiro pakurangarira zvepabonde
Kubva zvaiitika kazhinji zveA2 mating mhando mu P. infestans huwandu hwevanhu hwakaenderana nekushandisa kwakasimba kwemetalaxyl kurwisa kunonoka kwekupedzisira, zvakafungidzirwa kuti metalaxyl inokonzeresa shanduko yemhando dzekukwirisa. MuP. Parasitica, shanduko yakadaro pasi pechiito chaChloroneb uye metalaxyl yakaratidza kuyedzwa (Ko, 1994). Ndima imwechete pane svikiro ine yakadzika mametalexyl yakatungamira mukubuda kwehemothallic isolates kubva padambudziko raP. Infestans inonzwisisa metalaxyl ine mhando yekukwikwidzana A1 (Savenkova naCherepnikova-Anikina, 2002). Munguva dzinotevera ndima pane midhiya ine yakanyanya kusangana metalaxyl, hapana kana imwechete yekuzvipatsanura yerudzi rweA2 pairing yakaonekwa; zvisinei, vazhinji vanozviparadzanisa, kana vayambuka neA2 vakazviparadzanisa, pachinzvimbo che oospores, vakaumba yakaipa mycelium kuunganidza uye vaive vasina kuchena. Zvikamu zvemhando isingamiririri ine iyo A2 yekukwirisa mhando pane midhiya ine yakanyanya kukosheswa kwemetalaxyl yakatibvumidza isu kuona matatu mafomu emhando yekukwirisa mhando shanduko: 1) kuzadza kusimba kana kuyambuka neA1 neA2 kuzviparadzanisa; 2) homotallism (kuumbwa kwe oospores mune monoculture); 3) kushandurwa kwemhando yeA2 yekukwikwidza kuita A1. Nekudaro, metalaxyl inogona kukonzera shanduko mumhando dzekusvirana muP infestans vanhu uye, nekudaro, iko kuitika kwekudzokorora kwepabonde mavari.
Kufuridzira pamusoro peyekudzokorora kwekudyara
Mamwe maantibayoti ekudzivirira majini akawedzera kuwanda kweiyo hyphal heterokaryotization uye yenyukireya diploidization (Poedinok naDyakov, 1981). Sezvambotaurwa, heterokaryotization ye hyphae panguva yekusanganiswa kwemhando dzakasiyana dzeP. Infestans inoitika zvisingawanzo nekuda kwechiitiko chekusapindirana kwezvirimwa mune iyi fungus. Nekudaro, magene ekurwisa mamwe maantibhakitiriya anogona kuve nemhedzisiro, inoratidzwa mukukunda kusapindirana kwezvirimwa. Iyi fuma yaive neiyo streptomycin yekudzivirira geni ye1S-1 mutant. Kuvapo kwezvinhu zvakadaro mumunda vanhu phytophthora kunogona kuwedzera kuyerera kwemajini pakati pematambudziko uye nekukurumidza kuchinjisa kwehuwandu hwevanhu kune mhando nyowani kana fungicides.
Mimwe fungicides nemishonga inorwisa mabhakitiriya inogona kukanganisa kuwanda kwemitotic kudzokororazve, iyo inogona zvakare kuchinja mafambisirwo ejenotype muhuwandu hwevanhu. Iyo inowanzoshandiswa fungicide benomyl inosunga kune beta-tubulin, puroteni inovakwa ma microtubules e cytoskeleton, uye nekudaro inokanganisa maitiro echromosome kupatsanurwa mu anaphase yemitosis, ichiwedzera kuwanda kwemitotic kudzokedzana (Hastie, 1970).
Iyo fungicide para-fluorophenylalanine, inoshandiswa kurapa chirwere cheDutch mumashure, ine chivakwa chimwe chete. Para-fluorophenylalanine yakawedzera kuwanda kwekudzokororazve mune heterozygous diploids P. infestans (Poedinok et al., 1982).
Kuchinja kwecyclic mune genotypic kuumbwa kwevanhu mune yehupenyu kutenderera kweP. Infestans
Iyo yechinyakare yekuvandudza kutenderera kweP. Infestans munzvimbo inotonhora ine zvikamu zvina.
1) Chikamu chekuwedzera kwekuwedzera kwehuwandu hwevanhu (polycyclic chikamu) nezvizvarwa zvipfupi. Ichi chikamu chinowanzo kutanga muna Chikunguru uye chinotora mwedzi 1,5-2.
2) Chikamu chekumisa kukura kwehuwandu hwevanhu nekuda kwekuderera kwakaperera muhuwandu hweasina kukanganiswa matishu kana kutanga kwemamiriro ekunze asina kunaka. Ichi chikamu chinowira kunze kwekutenderera kwegore mumapurazi anoita ekutanga-kukohwewa kwemashizha kubviswa.
3) Chikamu chekutonhora mumatubhi, chichiperekedzwa nekuderera kukuru kwehuwandu hwevanhu nekuda kwehutachiona hwehutachiona hwehutachiona, kukura kunononoka kwehutachiona mavari, kusavapo kwekutapukirwazve kwehutachiona, kuora uye kubaya kweanokanganisa tubers pasi peyakajairika mamiriro ekuchengetedza. .
4) Chikamu chekunonoka kukura muvhu uye pane mbesa (monocyclic phase), mune iyo nguva yechizvarwa inogona kusvika mwedzi kana kupfuura (kunopera kwaMay - kutanga kwaChikunguru). Kazhinji panguva ino, mashizha ane chirwere anonetsa kuona, kunyangwe nekucherechedza kwakakosha.
Chikamu chekuwedzera kwekuwedzera kwehuwandu hwevanhu (polycyclic chikamu)
Kucherechedzwa kwakawanda (Pshedetskaya, Kozubova, 1969; Borisenok, 1969; Osh, 1969; Dyakov, Suprun, 1984; Rybakova, Dyakov, 1990) yakaratidza kuti pakutanga kweepiphytoty, yakaderera ine hukasha uye zvishoma zvine hukasha zvimiro zvinotungamira, izvo zvinobva zvatsiviwa ne zvine mwero uye zvine hukasha.uye mwero wekukura kwehasha dzevagari ndiwo akakwirira, kushomeka kushomeka kwakasiyana kwemuti unogamuchira.
Sezvo huwandu hwevanhu hunowedzera, kuwanda kwemajini maviri akasarudzika akakosheswa akaunzwa mumhando dzemabhizimusi (R1-R4) uye nekusarura kwazvakarerekera (R5-R11) zvinowedzera. Saka, muhuwandu hwevanhu pedyo neMoscow muna 1993, avhareji yehutachiona kubva pakupera kwaChikunguru kusvika pakati paNyamavhuvhu yakawedzera kubva pa8,2 kusvika pa9,4, uye kuwedzera kukuru kwakacherechedzwa kune yakasarudzika hutachiona hwemajini R5 (kubva pa31 kusvika pa86% yematombo ane huturu) ( Smirnov, 1996).
Kuderera kwekukura kwehuwandu hwevanhu kunoperekedzwa nekuderera kweye parasiti chiitiko chevanhu. Naizvozvo, mumakore ekushungurudzika, ese ari maviri manhamba emarudzi uye huwandu hwemadzinza ane huturu hwakaderera pane epiphytotic iwo (Borisenok, 1969). Kana pakukwirira kweepiphytotic mamiriro ekunze achichinja kuva asingafadzi kunonoka kwekusviba uye mbatatari inodzikira, ipapo kuwanda kwematombo ane hukasha uye ane hukasha kunoderera (Rybakova et al., 1987).
Kuwedzera kwehuwandu hwemajini hunokanganisa huroyi uye hukasha hwevagari zvinogona kuve nekuda kwekusarudzwa kweakanyanya uye ane hukasha clones mune yakasanganiswa vanhu. Kuratidza kusarudzwa, nzira yekuongororwa kwekuchinja kwazvakarerekera yakagadziridzwa, iyo yakashandiswa zvinobudirira mune chemostat vanhu vembiriso (Adams et al., 1985) uye Fusarium graminearum (Wiebe et al., 1995).
Iko kuwanda kweiyo mutant inopesana ne blasticidin S mumunda wevanhu P. infestans yakaderera zvakafanana nekukura kwehukasha hwehuwandu hwevanhu, izvo zvinoratidza shanduko mumakona makuru panguva yekukura kwevanhu (Rybakova et al., 1987 ).
Nguva yechando mumatubu
Munguva yechando mumatopu embatatisi, humhutu uye hukasha hweP infestans mafashama anoderera, uye kuderera kwehutachiona kunoitika zvishoma nezvishoma pane hukasha (Rybakova naDyakov, 1990). Sezviri pachena, pasi pemamiriro ezvinhu anobatsira kukurumidza kukura kwehuwandu hwevanhu (r-kusarudzwa), "kuwedzerwa" hutachiona hutachiona uye hukasha hwakanyanya hunobatsira, saka kuvandudzwa kweepiphytotic kunoenderana nekusarudzwa kweakanyanya uye ane hukasha clones. Pasi pemamiriro ekuzadza kwezvakatipoteredza, kana isiri iyo mwero wekuberekesa, asi kushingirira kwekuvapo mumamiriro asina kunaka (K-kusarudzwa), inoita basa rakakosha, "mamwe" majeni ehutachiona uye hukasha zvinoderedza kusimba, uye maonesheni nemajeni aya. ndivo vekutanga kufa kunze, saka humhu hwevanhu huri kudonha
Chikamu chemiriwo muvhu
Ichi chikamu ndicho chinonyanya kushamisa muhupenyu hupenyu (Andrivon, 1995). Kuvapo kwaro kwakatemerwa zvakanyatsofungidzirwa - nekuda kwekushayikwa kweruzivo nezve zvinoitika kune pathogen kwenguva yakareba (dzimwe nguva inopfuura mwedzi) - kubva pakubuda kwembeu dzemazamu kusvika pakuonekwa kwenzvimbo dzekutanga dzechirwere pavari. Kubva pane zvakawonekwa uye kuyedzwa, hunhu hwefungus munguva ino yehupenyu hwakavakwazve (Hirst naStedman, 1960; Boguslavskaya, Filippov, 1976).
Kupararira kwefungus kunogona kuumba pane hutachiona hutachiona muvhu. Izvo zvinokonzeresa spores zvinomera ne hyphae, iyo inogona kumera kwenguva yakareba muvhu. Yepuraimari (yakaumbwa pamatubhi) uye yechipiri (pane mycelium muvhu) mhodzi dzinokwira pamusoro pevhu nema capillary mafungu, asi dzive nekwanisi yekukanganisa mbatatisi chete mushure mekunge mashizha ayo ezasi adonha osangana nepamusoro pevhu. Mashizha akadaro (kureva, mavara ekutanga echirwere anowanikwa paari) haaumbike nekukasira, asi mushure mekukura kwenguva refu uye kusimukira kwematata.
Nekudaro, iyo saprotrophic zvinomera chikamu inogona zvakare kuvapo muhupenyu hwekutenderera kweP infestans. Kana muchikamu cheparasiti chehupenyu hwekutenderera hukasha ndicho chakakosha chikamu chemuviri, saka mune saprotrophic chikamu kusarudzwa kunoitirwa kudzikisira hutachiona, sekuyedzwa kunoitwa kune mamwe phytopathogenic fungi (ona Carson, 1993). Naizvozvo, muchikamu chino cheiyo kutenderera, zvivakwa zvine hukasha zvinofanirwa kurasikirwa zvakanyanya zvakanyanya. Asi kusvika parizvino hapana kuyedza kwakananga kwakaitwa kusimbisa fungidziro dziri pamusoro.
Shanduko dzemwaka dzinokanganisa kwete chete pathogenic zvivakwa zveP infestans, asiwo kushora fungicides, iyo inokura muchikamu che polycyclic (panguva epiphytoties) uye inoderera munguva yekuchengetedza yechando (Derevyagina et al., 1991; Kadish naCohen, 1992). Kudonhedza kwakanyanya mukushora metalaxyl kwakacherechedzwa munguva iri pakati pekudyarwa kweanokanganisa tubers uye kuoneka kwenzvimbo dzekutanga dzechirwere mumunda.
Intraspecific hunyanzvi uye shanduko yayo
P. infestans iri kukonzera madenda muzvirimwa zviviri zvakakosha pakutengesa, mbatatisi nemadomasi. Epiphytoties pamatatata yakatanga nguva pfupi mushure mekunge fungus yapinda munzvimbo nyowani. Kukundwa kwetamato kwakacherechedzwa nguva pfupi mushure mekuonekwa kwehutachiona pamatatata, asi epiphytoties pane madomasi akaonekwa chete makore zana gare gare - pakati pezana ramakore rechiXNUMX. Izvi ndizvo zvinonyorwa naHallegli naNiederhauser nezve kukundwa kwemadomasi muUSA
(1962): "Kwemakore angangoita zana mushure mekurwiswa kwakanyanya kwa100, kwakaedzwa kushoma kana kusatomboitwa kuwana mhando dzedomasi risingamiri. Kunyangwe rakanonoka blight rakatanga kunyorwa pamadomasi kutanga kwa1845, haina kuve chinhu chekutarisirwa kwakanyanya kwevafudzi pamuti uyu kudzamara kwakasimba kwechirwere muna 1848. Panharaunda yeRussia, nguva yekupedzisira yetamato yakanyoreswa muzana ramakore rechi1946. “Kwenguva yakareba, vaongorori havana kucherechedza chirwere ichi, nekuti hachina kukonzera kukanganisa kwakanyanya kwehupfumi. Asi mumakore makumi matanhatu nemakumi manomwe. Epiphytoties yezana ramakore makumi maviri yekupedzisira kwekukuvara padomasi inoonekwawo muSoviet Union, kunyanya mudunhu reVolga, muUkraine, North Caucasus, kuMoldova ... ”(Balashova, 60).
Kubva ipapo, madomasi blight nekunonoka kusvibiswa ave pagore, akapararira mudunhu rese rekurimwa kwemaindasitiri nekumba uye anokonzeresa kukuvara kwehupfumi kune chibage ichi. Chii chakaitika? Nei kuoneka kwekutanga kwehutachiona pamatezi uye epiphytotic lesion yetsika iyi zvichiitika panguva imwe chete, nepo zvakatora zana ramakore kuti epiphytotic ioneke padomasi? Iyi misiyano inotsigira weMexico kwete weSouth America sosi yehutachiona. Kana iyo mhando Phytophthora infestans yakaumbwa sehutachiona hweMexicoan tuber-inotakura marudzi eiyo genus Solanum, saka zvinonzwisisika nei mbatatisi dzakarimwa dziri muchikamu chimwe chete chejenasi semhando dzeMexico dzakakanganiswa zvakanyanya, asi nekuda kwekushayikwa co-shanduko nehutachiona, iyo isina kuvandudza nzira dzekupokana chaiko uye nekusagadzikana.
Iyo tomato ndeyechikamu chakasiyana chejenasi, mhando yekuchinjana kwayo ine misiyano yakakura kubva kumarudzi ane tuberous, saka, kunyangwe hazvo iyo nyanya isiri kunze kwehunyanzvi hwekudya hweP infestans, kusimba kwekukundwa kwayo kwanga kusingakwani yekurasikirwa kwakakomba kwehupfumi.
Iko kubuda kweepiphytoties pane nyanya kunokonzerwa nekuchinja kwakakomba kwemajini muhutachiona, izvo zvakawedzera kuchinjika kwayo (pathogenicity) panguva yeparasitism. Isu tinotenda kuti iyo nyowani, yakasarudzika fomu yekuparadza madomasi ndiyo T1 mujaho wakatsanangurwa naM. Gallegly, uchikanganisa mhando dzekrisita matomatisi (Red Cherry, Ottawa), inopesana nemujaho weT0 wakapararira pamatata (Gallegly, 1952). Sezvineiwo, shanduko (kana nhevedzano yeshanduko) iyo yakashandura mujaho weT0 mumujaho weT1 uye yakatungamira mukubuda kwemakona akagadziriswa zvakanyanya kukundwa kwetamato. Sezvazvinowanzoitika, kuwedzera kwehutachiona kune mumwe mugari kwakaperekedzwa nekuderera kwaro kune imwe, ndiko kuti, kutanga, kusati kwazadzikiswa kwehunyanzvi kwakamuka - kune mbatatisi (mujaho T0) uye kuna nyanya (nhangemutange T1).
Ndeupi humbowo hunotsigira iyi fungidziro?
- Kuitika pamatato nematomatisi. Pamashizha emadomasi, mujaho weT1 unotungamira, nepo pamashizha embatatisi usingawanzoitika. Sekureva kwaSF Bagirova naT.A. Oreshonkova (isina kuburitswa) mudunhu reMoscow muna 1991-1992, kuitika kwemujaho weT1 mukudyara mbatatisi kwaive 0%, uye mumadomasi kudyara - 100%; muna 1993-1995 - 33% uye 90%, zvichiteerana; muna 2001 - 0% uye 67%. Dhata yakafanana yakawanikwa muIsrael (Cohen, 2002). Ongororo nehutachiona hwembatatari tubers ine yakasarudzika yemujaho weT1 uye musanganiswa wekuzviparadzanisa T0 neT1 zvakaratidza kuti voga vemujaho weT1 havana kuchengetedzwa zvakanaka mumatubhu uye vanotsiviwa nevasarudzika veiyo T0 mujaho (Dyakov et al., 1975; Rybakova, 1988).
2) Dynamics yemujaho weT1 mumadomasi kudyarwa. Huru hutachiona hwemashizha emadomasi anoitwa neakasarudzika eiyo T0 mujaho, uyo unotungamira mukuongororwa kwehutachiona munzvimbo dzekutanga dzakagadzirwa pamashizha. Izvi zvinosimbisa chirongwa chinowanzo gamuchirwa chekufamba kwehutachiona: Huru hukuru hwehutachiona kubva kumatatata hunoumbwa nemujaho weT0, zvisinei, huwandu hushoma hweT1 clones inochengetwa mumatatata, kamwe padomasi, bvisa mujaho weT0 uye uzviunganidze kuguma epiphytotic nguva. Izvo zvakare zvinokwanisika kuti kune imwe nzira sosi yematomatisi hutachiona hutachiona neiyo T1 mujaho, isina kusimba sembatatari tubers nemashizha, asi inogara iripo. Naizvozvo, sosi iyi haina zvainoita pamaumbirwo ehuwandu hwevanhu vari kutapira madomasi, asi zvichizo tarisa kuunganidzwa kwerudzi rweT1 (Rybakova, 1988; Dyakov et al., 1994).
3) Kushatirwa kumatato nematomatisi. Hutachiona hwehutamatisi nemashizha embatatisi ane magaera emarudzi T0 neT1 zvakaratidza kuti zvekare zvine hukasha hwembatatisi pane zvemadomasi, uye iyo yekupedzisira ine hukasha hwetamato kupfuura mbatatisi. Iyi misiyano inoratidzwa mukushandurwa kwevanoparadzaniswa nerudzi rusiri "rwoga" kubva kune vanhu vakavhengana panguva dzendima dzemashizha mugreenhouse (Dyakov et al., 1975) uye muminda yeminda (Leberton et al., 1999); misiyano muiyo shoma inotapukira inotakura, latency nguva, saizi yenzvimbo dzinotapukira uye kugadzirwa kwepore (Rybakova, 1988; Dyakov et al., 1994; Legard et al., 1995; Forbes et al., 1997; Oyarzun et al., 1998; Leberton et al., 1999; Vega-Sanchez et al., 2000; Knapova, Gisi, 2002; Sussuna et al., 2004).
Hasha dzekuzvipatsanura dzerudzi rweT1 kumhando dzemadomasi dzinoshaya majekiseni ekurwisa dzakakwirira zvekuti izvi zvinoparadzanisa spore pamashizha senge pane svutugadzike pasina kuumbiridza hutachiona (Dyakov et al., 1975; Vega-Sanchez et al., 2000) .
4) Virulence yetapiri nematomatisi. Mujaho weT1 unokanganisa mhando dzemadomasi Cherry nePh1 yekudzivirira geni, nepo T0 mujaho usingakwanise kutapukisa marudzi aya, i.e. ine hutachiona hwakadzikira. Zvichienderana nevanosiyanisa
Iwo maR-genes embatatisi akapindirana zvine hukama Mhando dzakasarudzika kubva kumadomasi mashizha hadzina huturu kupfuura "mbatatisi" marata (Tafura 11).
5) Kwazvakarerekera mamaki. Kuongororwa kwevasina kwavakarerekera mamaki muhuwandu hwevanhu P. infestans parasitizing pamatezi nematomatisi zvakare inopupurira kune multidirectional intraspecific sarudzo. Mune vanhu veBrazil ve P. infestans, matomatisi mashizha akapatsanurwa aive emutsetse weclonal US-1, uye iwo anobva kumashizha embatatari aive eiyo BR-1 mutsetse (Suassuna et al., 2004). MuFlorida (USA), kubvira 1994, dombo reUS-90 rakatanga kutonga matapiri (iine chiitiko chinopfuura makumi mapfumbamwe muzana), uye inonamira US-8 neUS-11 padomasi, uye vekupedzisira voga vanoshusha nyanya. kupfuura mbatatisi (Weingartner, Tombolato, 17). Misiyano yakakosha mumagenotype frequency (DNA zvigunwe) mumapatatisi uye matomatisi akapatsanurwa akagadzirirwa 2004 P. infestans strains akaunganidzwa muUnited States kubva muna 1200 kusvika 1989 (Deahl et al., 1995).
Uchishandisa nzira yeAFLP yakaita kuti zvikwanisike kupatsanura masaini 74 akaunganidzwa kubva kumazamu embatatisi nemadomasi muna 1996-1997. muFrance neSwitzerland, mumapoka manomwe. Mhando dzembatatisi nedomasi hadzina kunyatso kusiyaniswa, asi iyo "mbatatisi" michina yakazove yemageneti akasiyanasiyana kupfuura "madomasi". Iwo ekutanga akawanikwa mune ese masumbu manomwe, uye ekupedzisira, mana chete, izvo zvinoratidza imwe yakasarudzika genome yeiyo yekupedzisira (Knapova naGisi, 7).
6) Maitiro ekuzvitsaura. Kana huwandu hwehutachiona hwemhando mbiri dzezvirimwa hwakashanduka huchienda mukudzikisira hunyanzvi kuenda kune yavo "yega", ipapo nzira dzakasiyana-siyana dzepamberi uye postmeiotic dzinomira dzinodzivirira kupindirana kwemajini (Dyakov naLekomtseva, 1984).
Ongororo dzinoverengeka dzakatsvaga mhedzisiro yesosi yematambudziko evabereki pane kugona kwehybrisization. Pakuyambuka mitsara yakasarudzika kubva kumarudzi akasiyana ejenor Solanum muEcuador (Oliva et al., 2002), zvakaonekwa kuti maronda ane A2 mating mhando kubva kusango Solanaceae (clonal mutsetse EC-2) yakayambuka yakaipisisa nematra kubva kudomasi ( mutsara EC -3), uye zvakanyatso kuyambuka neiyo mbatatisi strain (EC-1).
Ese mahybrids akawanikwa asiri-pathogenic. Vanyori vanotenda kuti iyo yakaderera muzana yekusanganiswa uye kudzikiswa kwepathogenicity mune mahybrids zvinokonzerwa neepostmeiotic nzira dzekuberekana dzega dzevanhu.
Mukuyedza kwaBagirova et al. (1998), huwandu hwakawanda hwetatata nematomatisi akayambuka pamwe nemidziyo yeiyo T0 neT1 nhangemutange. Iyo yakaorera kwazvo yaive michinjikwa yematombo eT1xT1 akaparadzaniswa nemadomasi (36 oospores mumicroscope field yekutarisa, 44% ye oospore kumera), yakanyanya kushanda yaive michinjikwa yemadzinza eT0xT1 akasiyiwa kubva kune akasiyana mauto (huwandu hwakaderera hweanosimudzira uye akamera oospores , yakakwira chikamu cheabortive uye isina kusimukira oospores) ... Kugona kwemuchinjiko pakati pevamwe vakasarudzika mujaho weT0 wakaparadzaniswa nemapatata wakazove wepakati. Sezvo mutumbi mukuru wemujaho weT0 uchikanganisa mbatatisi, ine sosi yekuvimbika yechando - mbatatari tubers, semhedzisiro yekukosha kwemahombekombe seanotonhora zvikamu zvinotapukira zvevanhu kubva kumatatata zvakaderera. "Tomato fomu" rakachinjika rinokwanisa kuchando padomasi muchimiro che oospores (ona pazasi) uye nekudaro rinoramba riine hupfumi hwepamusoro pemaitiro epabonde. Nekuda kwekubereka kwaro kukuru, T1 inowana mukana wakazvimiririra wehutachiona hwekutanga mudomasi. Mhedzisiro yakawanikwa naKnapova et al. (Knapova et al., 2002) inogona kududzirwa nenzira imwecheteyo. Michero yemhando dzakasarudzika kubva kumatatata nematra kubva kudomasi yakapa yakanyanya kuwanda maoospores - 13,8 pa sq. Mm. yepakati (ine kupararira kwe5-19) uye yepakati muzana yekumera kwemaoospores (6,3 nekupararira kwe0-24). Kuyambuka kwenzira dzakaparadzaniswa nemadomasi kwakapa yakaderera muzana muzana yeoospores (7,6 ine kupararira kwe4-12) ine yakakwira kwazvo chikamu chekumera kwavo (10,8). Iyo michinjikwa pakati pematambudziko akasiyiwa kubva kumatatata yakapa iyo yepakati nhamba ye oospores (8,6 ine yakawanda yekuparadzira dhata - 0-30) uye yakadzika kwazvo muzana yekumera kwe oospores (2,7). Nekudaro, maronda kubva kumatatata haana kubereka pane aya anobva kudomasi, asi michinjikwa yekudyidzana haina kupa mhedzisiro yakaipa kupfuura yekudyara. Izvo zvinokwanisika kuti mutsauko nedata riri pamusoro naBagirova et al. zvinotsanangurwa nenyaya yekuti vaongorori veRussia vakashanda nematrains akasarudzika mukutanga kwema 90 ezana ramakore rechimakumi maviri, uye vaongorori veSwitzerland - nematambudziko akasarudzika kuma 90s apfuura.
Kubereka kushoma kunogona kuve kukonzerwa neheteroploidy yematambudziko. Kana mune vanhu veMexico, uko kusangana pabonde uye hutachiona hwekutanga nevana vane oosporous kuri kwakajairika, mazhinji ematambudziko akaongororwa eP.Infestans idiploid, saka munyika dzeOld World kupindira kwepolymorphism yeploidy kunoonekwa (di-, tri- nematetraploid strains, pamwe neheterokaryotic strains ane heteroploid nuclei), nematambudziko ane mhando dzakasiyana dzekusvirana, i. yakaenzana inobereka, yakasiyana mu nucleus ploidy (Therrien et al., 1989, 1990; Whittaker et al., 1992; Ritch, Daggett, 1995). Misiyano yenuclei muantheridia uye oogonia inogona kukonzeresa kubereka.
Kana zviri zvekuchinjana kwenyukireya pakati pehyphae panguva yeanastomoses, izvi zvinodzivirirwa nekusawirirana kwezvirimwa, izvo zvinopatsanura huwandu hweasexual mumakombo mazhinji akasarudzika (Poedinok naDyakov, 1987; Gorbunova et al., 1989; Anikina et al., 1997b).
7) Kusangana kwevanhu. Idzi data riri pamusoro rinoratidza kuti kusanganiswa pakati pe "mbatatisi" uye "madomasi" maronda eP. Infestans zvinogoneka. Kudzokororazve hutachiona hwevanhu vakatsaukana kwakasiyana zvakare kunogoneka, kunyangwe nekudzora hukasha.
Kuongororwa kwevanhu vanocherekedza vanhu vari vega kubva kuminda iri pedyo nembatatisi nemadomasi muna1993 kwakaratidza kuti chingangoita chikamu chimwe muzvina chevanhu vakazviparadzanisa nemashizha emadomasi vakatamiswa kubva kumunda wembatatisi uri pedyo (Dolgova et al., 1997). Pfungwa, zvinogona kufungidzirwa kuti kusiyana kwevanhu munzvimbo mbiri kunowedzera uye kunotungamira mukubuda kwemhando dzakasiyana-siyana (f.sp. mbatatisi uye f.sp. tomato), kunyanya sezvo oospores inogona kuramba iri mumarara emiti (Drenth et al., 1995; Bagirova, Dyakov, 1998) uye madomasi mbeu (Rubin et al., 2001). Nekudaro, madomasi parizvino ane sosi yekumutsiridzwa patsva yakazvimiririra nembatatari tubers.
Zvisinei, zvose zvakaitika zvakasiyana. Kukundwa ne oospores kwakabvumira utachiona kudzivirira nhanho yakatetepa muhupenyu hwayo - monocyclic nhanho yezvinomera muvhu, panguva iyo zvipembenene zvinoderera, izvo zvishoma nezvishoma zvinodzoserwa muzhizha muchikamu che polycyclic.
Tafura 11. Kakawanda kwehutachiona hwehutachiona kumhando dzakasiyana dzembatatisi muP infestans strains
nyika | Gore | Avhareji nhamba yehutachiona hutachiona mumatambudziko | Munyori | |
kubva kumatatata | kubva kudomasi | |||
France | 1995 | 4.4 | 3.3 | Leberton et al., 1999 |
1996 | 4.8 | 3.6 | Leberton, Andrivon, 1998 | |
France, Switzerland | 1996-97 | 6.8 | 2.9 | Knapova, Gisi, 2002 |
United States | 1989-94 | 5 | 4.8 | Goodwin et al., 1995 |
USA, Zap. Washington | 1996 | 4.6 | 5 | Dorrance et al., 1999 |
1997 | 6.3 | 3.5 | " | |
Ecuador | 1993-95 | 7.1 | 1.3 | Oyarzun et al., 1998 |
Israel | 1998 | 7 | 4.8 | Cohen, 2002 |
1999 | 6 | 5.7 | " | |
2000 | 6.7 | 6.1 | " | |
Russia, Mosk. nharaunda | 1993 | 8.9 | 6.7 | Smirnov, 1996 |
Russia, matunhu akasiyana | 1995 | 9.4 | 8 | Kozlovskaya nevamwe. |
1997 | 9.2 | 9.2 | " | |
2000 | 8.7 | 4.8 | " |
Chekutanga zoosporangia uye zoospores, iyo inokura oospores, ine huwandu hwakanyanya hwehutachiona, kunyanya kana oospores yakaumbwa parthenogenetically pasi pesimba remapheromone erudzi nerumwe rudzi rwekusangana. Naizvozvo, izvo zvinotapukira pamamato emadomasi akakura kubva pambeu dzakatapukirwa ne oospores ine hutachiona hwakanyanya kune ese madomasi nembatatisi.
Idzi shanduko dzakatungamira kune kumwe kugadziridzwa kwevagari, kwakaratidzwa mune zvinotevera zvakakosha shanduko kubva padenda rekuona.
- Mbeu dzematomatisi dzakabatwa dzave sosi yakakosha yehutachiona hwekutanga hwembatatisi (Filippov, Ivanyuk, meseji yako).
- Epiphytoties pamatato yakatanga kucherechedzwa kutanga kwaJune, inenge mwedzi yapfuura kupfuura zvakajairwa.
- Mukudyarwa kwembatatisi, chikamu chemujaho weT1 chakawedzera, icho chakambowanikwa imomo mune chisina kukosha (Ulanova et al., 2003).
- Mitsetse yakasarudzika kubva kumadomasi mashizha haisisina kusiyana nemhando yematatisi muhutachiona pamasiyano embatatisi ehutachiona hutachiona uye akatanga kudarika "mbatatisi" maratidziro mune hukasha kwete chete pane madomasi, asi nembatatisi (Lavrova et al., 2003; Ulanova et al. , 2003).
Nekudaro, panzvimbo pekusiyana, paive nekubatana kwevanhu, kubuda kwehuwandu hwevanhu pazvirimwa zviviri zvine huturu hwakanyanya uye hukasha kune ese marudzi.
mhedziso
Nekudaro, kunyangwe makore anodarika zana nemakumi mashanu ekudzidza kwakadzama kwaP. Infestans, mubhayoloji, kusanganisira huwandu hwevanhu hweiyo biusative agent yezvirwere zvakakosha zvezvirimwa zvakasimwa solanaceous, zvakawanda zvinoramba zvisingazivikanwe. Hazvisi pachena kuti kupindirana kwezvikamu zvega zvega zvehupenyu zvinokanganisa sei chimiro chevanhu, ndeapi majekiseni ekuchinja kwakasiyana kwehukasha uye huturu, chii chiyero chehurongwa hwekubereka uye hwekubereka muvanhu vakasikwa, sei kusapindirana kwezvirimwa kunogarwa nhaka, basa rembatatisi nematomatisi mune hutachiona hwekutanga hwezvirimwa izvi uye mune chii chinogadzirisa chimiro chevanhu vehutachiona. Kusvika parizvino, nyaya dzakakosha dzakadai sedzemagariro ekuchinja hukasha hwehutachiona kana kukukurwa kwekupokana kweembatatisi hakuna kugadziriswa. Nekudzika uye kuwedzerwa kwekutsvaga pamusoro pembatata kunonoka kusvibiswa, hutachiona hunounza matambudziko matsva kuvatsvagiri. Nekudaro, kuvandudzwa kwesimba rekuyedza, kumuka kwenzira nyowani dzekusanganisa nemajini nemapuroteni kunotibvumidza kutarisira mhinduro inobudirira yemibvunzo yakabvunzwa.
Chinyorwa chakaburitswa mumagazini "Kudzivirirwa kweMbatatisi" (Nha. 3, 2017)