S.N. Elansky, L.Yu. Kokaeva, N.V. Statsyuk, Yu.T. Dyakov
Taw qhia
Oomycete Phytophthora infestans (Mont.) De Bary, tus neeg sawv cev ntawm lub ntsej muag blight, cov kab mob tseem ceeb tshaj plaws ntawm cov qos yaj ywm thiab txiv lws suav, tau ua tib zoo mloog los ntawm cov kws tshawb fawb los ntawm txawv teb chaws rau ntau tshaj ib puas xyoo thiab ib nrab. Kev tshwm sim tam sim ntawd tshwm sim nyob rau hauv Europe li ntawm nruab nrab ntawm xyoo pua puv XNUMX, nws ua rau muaj kev kis mob qos yaj ywm uas tseem nyob hauv lub cim xeeb ntawm ntau tiam neeg.
Txog rau tam sim no, nws feem ntau hu ua "nceb ntawm Irish kev tshaib plab". Yuav luag ib puas xyoo tom qab thawj phaum mob, cov qos ntoo dub Mev hom tiv taus cov kab blight lig tau pom, cov hau kev hla lawv nrog cog qos yaj ywm tau tsim (Muller, 1935), thiab cov thawj lig blight-resistant ntau yam tau txais (Pushkarev, 1937). Txawm li cas los xij, tsis ntev tom qab pib ntawm lawv txoj kev ua lag luam sau qoob, kev sib tw ntawm cov kab mob blight lig uas tau ua rau cov neeg tsis taus khov kom tiv taus ntau ntau yam. thiab kev qhia txog cov tshuaj tiv thaiv tshiab los ntawm cov qos yaj ywm Mev qus rau hauv ntau hom tau pib poob sai.
Ua tsis tiav nrog kev siv cov tshuaj monogenic (ntsug) ua rau lub cev tsis sib haum xeeb saib xyuas rau ntau txoj hauv kev ntawm kev siv cov nonspecific polygenic (kab rov tav) tsis kam. Xyoo tsis ntev los no, cov haiv neeg muaj kev sib ntau tau pib ntau zuj zus nyob hauv ib tus neeg ntawm cov cab, ua rau yaig ntawm txawm nonspecific tsis kam. Lub hnub tuaj ntawm cov tshuaj tua kab-tua tau ua rau muaj teeb meem hauv kev siv cov tshuaj tiv thaiv qos yaj ywm.
Vim tias qhov sib txawv tseem ceeb ntawm oomycetes thiab cov kab mob hu ua fungi hauv tshuaj lom neeg, ultrastructure, thiab cov metabolism, fungicides, tshwj xeeb tshaj yog cov kab ke uas siv los tiv thaiv cov nroj tsuag los ntawm ntau cov kab mob fungal, yog qhov tsis muaj txiaj ntsig tiv thaiv oomycetes.
Yog li, hauv kev siv tshuaj tiv thaiv tiv thaiv blight lig, ntau yam (txog 12 zaug hauv ib lub caij lossis ntau dua) kev txau nrog kev sib cuag nrog kev npaj ntawm cov kev pom dav dav tau siv. Lub kauj ruam ntawm kev tawm tsam yog kev siv phenylamides, uas yog lom rau oomycetes thiab sib kis kab ke hauv cov nroj tsuag. Txawm li cas los xij, lawv txoj kev siv dav siv sai sai ua rau kev tsub zuj zuj ntawm cov kab mob tsis haum hauv cov neeg muaj peev xwm tiv thaiv kab mob (Davidse et al., 1981), uas cuam tshuam cov nroj tsuag tsis zoo. P. Cov kab mob no tsuas yog qhov cab ntawm qhov tsis zoo, qhov kev puas tsuaj rau hauv kev ua liaj ua teb hauv organic tsis tuaj yeem cuam tshuam tsis muaj kev siv tshuaj los ntawm kev tiv thaiv (Van Bruggen, 1995).
Qhov saum toj no piav qhia txog kev saib xyuas ntau heev them nyiaj los ntawm cov kws tshawb fawb los ntawm txawv teb chaws los kawm txog P. infestans neeg, kev hloov pauv ntawm lawv ntau thiab caj ces muaj pes tsawg leeg, nrog rau kev hloov caj ceg ntawm kev hloov pauv.
Lub neej voj voog ntawm R. INFESTANS
Oomycete Phytophthora infestans nthuav dav mycelium nrog haustoria hauv cov nplooj qos. Pub rau ntawm cov nplooj qhwv cov tawv nqaij, nws ua rau tsim ntawm tsaus me ntsis, uas tig mus rau dub thiab rot hauv huab cua ntub. Nrog txoj kev swb muaj zog, tag nrho cov nplooj tuag. Tom qab lub sijhawm pub mis, outgrowths yog tsim nyob rau ntawm mycelium - sporangiophores - uas loj hlob sab nraud dhau ntawm lub plab. Nyob rau hauv huab cua ntub, lawv tsim ib cov paj dawb ib ncig ntawm me ntsis ntawm qhov tsis tau sab hauv ntawm nplooj. Ntawm qhov kawg ntawm sporangiophores, txiv qaub-puab zoosporangia raug tsim, uas tawg tawm thiab nqa los ntawm tshuaj tsuag los nag (Daim duab 1). Ntog mus rau hauv cov tee dej rau saum npoo ntawm cov nplooj qos, sporangia germinate nrog 6-8 zoospores, uas, tom qab ib ntus ntawm kev txav, muaj puag ncig, them nrog lub plhaub thiab tua tawm nrog lub raj kab mob. Cov ntoo tawm tuaj dhau ntawm lub plab me me mus rau cov nplooj ntoo ua ke. Hauv qee qhov xwm txheej, sporangia tuaj yeem cog rau hauv txoj kev loj hlob raj ncaj qha mus rau cov ntaub so ntswg nplooj. Hauv cov xwm txheej zoo, lub sijhawm los ntawm kev kis tus kab mob mus rau kev tsim kom muaj cov sporulation tshiab tsuas yog 3-4 hnub.
Ib zaug ntawm hauv av thiab lim los ntawm cov av, sporangia tuaj yeem kis tau cov hlab. Kev cuam tshuam loj heev rot rot thaum lub sij hawm cia; nyob rau hauv cov cuam tshuam tsis muaj zog, tus kab mob yuav nyob ntev mus txog lub caij tom ntej. Tsis tas li ntawd, tus neeg sawv cev ntawm lub cev blight tuaj yeem ua rau lub caij ntuj no nyob rau hauv daim ntawv ntawm oospores (tuab-walled so kev sib deev spores) hauv cov av ntawm cov nroj tsuag khib nyiab thiab ntawm cov noob txiv lws suav. Oospores yog tsim los ntawm cov nroj tsuag muaj sia nyob chaw thaum muaj kev sib txawv ntawm cov khub sib txawv ntsib nrog ntau noo noo. Nyob rau lub caij nplooj ntoo hlav, cov qhov tso hnoos qeev yog tsim rau cov kab mob cog thiab cov cog ntoo nrog cov oospores; Qee qhov xwm txheej, mycelium tuaj yeem loj hlob ntawm cov tuber uas kis rau mus rau ntsuab ntsuab ntawm cov nroj tsuag thiab feem ntau tshwm sim nyob rau sab qaum kev qia.
Ib qhov sib txawv tseem ceeb ntawm oomycetes thiab feem ntau ntawm cov neeg hu ua fungi nyob hauv qhov tseem ceeb ntawm daim ntawv pov thawj hauv lawv lub neej nrog gametic meiosis thiab kev tawg ntawm zygotes (oospores) tsis muaj lub zog ntawm fim nuclear. Cov yam ntxwv no, ntxiv rau dipolar heterotallism hloov bisexuality, yuav zoo li ua rau nws siv tau rau oomycetes cov txheej txheem tsim los rau kev kawm cov neeg ntawm eukaryotes ntau dua (kev soj ntsuam ntawm panmixia thiab subdivision ntawm cov neeg, intra- thiab interpopulation gene flow, thiab lwm yam). Txawm li cas los xij, peb lub ntsiab lus tsis tso cai hloov tag nrho cov kev kawm no rau kev kawm P. infestans.
1. Nrog rau kev ua ke nrog cov xov paj, nws tus kheej-tsim muaj thiab parthenogenetic oospores yog tsim nyob rau hauv cov neeg (Fife thiab Shaw, 1992; Anikina et al., 1997a; Savenkova, Cherepnikoba-Anirina, 2002; Smirnov, 2003), thiab qhov ntau zaus ntawm lawv tsim tau txaus los cuam tshuam ntawm cov ntawv xeem.
2. Cov txheej txheem kev sib daj sib deev hauv P. infestans ua qhov tsis tseem ceeb rau kev ua haujlwm ntawm cov pejxeem loj, vim tias cov pwm tuaj yeem ua kom muaj feem ntau los ntawm kev cog qoob loo, tsim rau ntau dua 90% ntawm cov txiaj ntsig ntawm qhov kev soj ntsuam ntawm hom kev tawm tsam los ntawm cov txheej txheem ib txwm ntawm cov khoom noj nruab nrab ... lub caij cog qoob loo yog ob peb tiam ntawm asexual sporulation (kev txhim kho kab mob polycyclic). Oospores ua si lub luag haujlwm tseem ceeb hauv kev tiv thaiv kab mob ntawm lub sijhawm thaum tsis muaj nroj tsuag ntsuab (thaum caij ntuj no) thiab nyob rau thawj qhov muaj tus kab mob. Tom qab ntawd, thaum lub caij ntuj sov, clonal yug me nyuam tshwm sim thiab ib qho kev nce ntxiv los yog, hloov pauv, qhov kev txo qis ntawm tus neeg ntawm cov pob zeb uas tshwm sim los ntawm kev sib deev, uas yog qhov tseem ceeb tau txiav txim siab los ntawm kev xaiv ntawm qhov ntau yoog. Yog li no, qhov sib piv ntawm ib leeg pob rau ib tus pej xeem thaum pib thiab thaum xaus ntawm epiphytotics tuaj yeem sib txawv kiag li.
3. Lub voj voog tau piav qhia yog tus xeeb ceem ntawm cov haiv neeg ib txwm muaj P. infestans hauv lawv lub tebchaws, Central America. Lwm thaj chaw hauv ntiaj teb, kev sib daj sib deev tsis tau paub txog ntau tshaj 100 xyoo; Lub neej ntawm lub neej muaj kev sib cav ntau, thiab kev kis tau yog focal hauv qhov: kev kis kab mob los ntawm ib leeg cog tau hla mus rau nplooj, ua rau thawj foci ntawm tus kab mob, uas tuaj yeem sib koom ua ke thaum loj hlob ntawm tus kab mob.
Yog li, hauv qee thaj tsam yuav muaj kev hloov pauv ntawm kev sib deev thiab asexual kev voj voog, thaum nyob hauv lwm tus - tsuas yog lub voj voog asexual.
Keeb kwm ntawm P. INFESTANS
P. infestans tau tshwm sim hauv Tebchaws Europe thaum kawg ntawm thawj ib nrab ntawm xyoo 1991th. Txij li thaum cov qos yaj ywm ib txwm nyob rau qaum teb Qaum Teb Asmeskas, nws tau kwv yees tias tus cab no tau coj los ntawm muaj rau Tebchaws Europe thaum lub sijhawm tawg ntawm Chilean saltpeter. Txawm li cas los xij, cov kev tshawb fawb nqa tawm ntawm Rockefeller Center qos chaw nres tsheb hauv Toluca Valley, Mexico yuam kom qhov kev xav no tau hloov kho dua tshiab (Niederhauser 1993, XNUMX).
1. Hauv lub Toluca Valley, lub nroog tuberous hom qos yaj ywm (Solanum demissum, S. bulbocastanum, thiab lwm yam) muaj cov noob sib txawv rau cov ntsug ua ke nrog cov qib siab tsis tseem ceeb, uas qhia tau ntev txog kev hloov pauv nrog tus cab. Miskas qab teb, nrog rau cov qoob loo qos yaj ywm, tsis muaj cov noob tsis kam tiv taus.
2. Hauv Toluca Valley, muaj qhov sib cais nrog mating hom A1 thiab A2, qhov txiaj ntsig ntawm qhov uas cov neeg sib cuam tshuam ntawm P. infestans tau dav; hos nyob hauv lub teb chaw ntawm cov qos yaj ywm, Qab Teb Asmeskas, cov kab mob cab sib kis ncaj qha.
3. Nyob hauv Toluca Valley, muaj txhua xyoo kev mob loj heev ntawm qhov blight lig. Yog li no, ntawm cov kws tshawb fawb Asmeskas North (Cornell University), lub tswv yim hais txog Mesoamerica (Central America) raws li qhov chaw yug ntawm qos phytophthora yog tsim los (Goodwin li al., 1994).
Cov kws tshawb fawb Asmeskas teb chaws tsis qhia txog cov tswv yim no. Lawv ntseeg tias cov qos xyaw thiab nws cov tsiaj P. infestans muaj ib qho chaw nyob sib luag - South American Andes. Lawv tau txhawb lawv txoj kev xav los ntawm kev tshawb fawb txog molecular ntawm kev tsom xam ntawm DNA polymorphisms ntawm mitochondrial genome (mtDNA) thiab nuclear genes RAS thiab tub-tubulin (Gomez-Alpizar li al., 2007). Lawv pom tias cov kab sau los ntawm ntau qhov chaw ntawm lub ntiaj teb nqis los ntawm peb txoj kab sib txawv ntawm cov caj ces uas (tag nrho peb) muaj nyob hauv South American Andes. Andean haplotypes yog xeeb leej xeeb ntxwv ntawm ob kab: cais tawm ntawm qhov qub tshaj plaws ntawm mtDNA kab yog pom ntawm Solanaceae tsiaj qus los ntawm ntu Anarrhicomenum hauv Ecuador, thaum cais tawm ntawm kab thib ob yog ib txwm muaj ntawm qos yaj ywm, txiv lws suav, thiab tsaus ntuj dub. Hauv Toluca, txawm tias qhov mob haplotypes tsawg tsawg yog nqis los ntawm tsuas yog ib caj ces, nrog kev hloov caj ces ntawm Toluca lim (tsawg allelic zaus ntawm qee qhov chaw kuj tsis paub meej) qhia txog cov muaj zog muaj txiaj ntsig vim muaj kev tsis txaus siab tsis ntev los no.
Tsis tas li ntawd, ib hom tsiaj tshiab P. andina tau pom nyob hauv Andes, morphologically thiab cov caj ces zoo ib yam li P. infestans, uas, raws li cov kws sau ntawv, taw rau lub Andes ua qhov kub kub ntawm kev hais tawm hauv cov phytophthora. Thaum kawg, hauv Tebchaws Europe thiab Tebchaws Asmeskas, P. Cov neeg muaj kab mob coob tshaj plaws suav tag nrho ob tog Andean, thaum nyob hauv Toluca tsuas yog ib tus.
Cov ntawv tshaj tawm no tau ua rau cov lus teb los ntawm cov pab pawg neeg tshawb fawb los ntawm txawv teb chaws, uas tau ua ntau txoj hauv kev sim los kho qhov kev tshawb fawb dhau los (Goss et al., 2014). Hauv txoj haujlwm no, thawj zaug, ntau cov xov xwm me me ntawm cov xov tooj DNA tau siv los kawm DNA polymorphisms; qhov thib ob, rau kev tsom xam ntawm pawg, kev tsiv teb tsaws chaw, lub sijhawm ntawm kev sib cais ntawm cov neeg, thiab lwm yam. ntau tshaj cov qauv tau raug siv (F-txheeb cais, Bayesian kev kwv yees, thiab lwm yam) thiab, thib peb, kev sib piv tau siv tsis yog nrog Andean hom P. andina, uas ib qhov xwm keeb tsim muaj (P. infestans x Phytophthora sp.) , tab sis kuj nrog hom Mev teb tau P. mirabilis, P. Ipomoeae, thiab Phytophthora theemoli - ua kom kaw cov P. infestans koom nrog rau tib pawg (Kroon et al., 2012). Raws li kev soj ntsuam no, nws tau qhia txog tias lub hauv paus ntawm tsob ntoo phylogenetic ntawm txhua hom ntawm cov genus Phytophthora coj mus rau hauv qhov kev tshawb fawb, tshwj tsis yog lub keeb P. andina, yog los ntawm Mev hom, thiab cov khiav dej num ntws muaj qhov kev coj Mexico - Andes, thiab tsis yog lwm tus rov qab, thiab nws pib sib haum nrog European. txoj kev paub tswj hwm ntiaj teb Tshiab (300-600 xyoo dhau los). Yog li, qhov tshwm sim ntawm P. infestans hom, tshwj xeeb rau qhov yeej ntawm cov qos yaj ywm, tau tshwm sim hauv theem nrab caj ces nruab nrab ntawm kev tsim cov tuberous nightshades, i.e. hauv Central America.
Genome ntawm P. INFESTANS
Hauv xyoo 2009, ib pab pawg kws tshaj lij thoob ntiaj teb tau ua tiav P infestans genome (Haas li al, 2009), qhov loj me yog 240 MB. Qhov no yog ob peb zaug ntau dua li nyob hauv txheeb ze hom tsiaj P. sojae (95 Mb), ua rau cov hauv paus hniav ntawm cov taum pauv, thiab P. Ramorum (65 Mb), cuam tshuam zoo li cov ntoo uas muaj txiaj ntsig xws li ntoo qhib, beech, thiab lwm tus. Cov ntaub ntawv tau pom tias cov genome muaj ntau ntawm cov ntawv luam ntawm cov qib sib lawv liag - 74%. Lub genome muaj 17797 protein-coding genes, feem ntau ntawm cov caj ces koom nrog cov txheej txheem ntawm tes, suav nrog DNA replication, hloov ntaub ntawv thiab txhais cov protein.
Kev sib piv ntawm cov genus ntawm cov genus Phytophthora qhia tawm lub koom haum txawv txav ntawm cov genome, suav nrog cov ntu ntawm cov txheej txheem ntawm cov neeg tsis pom zoo, uas cov noob ceev muaj feem tsawg thiab cov ntsiab lus ntawm cov ntu rov qab yog qhov tsawg, thiab ib cheeb tsam uas tsis muaj kev tso cai ntawm cov noob, nrog lub noob tsawg thiab cov ntsiab lus ntawm rov qab thaj tsam. Cov kev thaiv tau suav txog 70% (12440) ntawm tag nrho cov P. infestans muaj protein-coding noob. Tsis pub dhau kev txhaws cia, cov noob feem ntau sib luag nrog qhov nruab nrab intergenic nrug ntawm 604 bp. Hauv thaj chaw nruab nrab ntawm kev thaiv thaj chaw, qhov kev ncua deb intergenic ntau dua (3700 bp) vim tias qhov nce ntawm qhov ntom ntawm cov khoom rov ua dua tshiab. Kev hloov kho sai sai cov txiaj ntsig cov ntaub ntawv kab ntxwv muaj nyob hauv cov cheeb tsam cov noob tsis zoo.
Sequence tsom xam ntawm P. Infestans genome pom tau tias kwv yees li ib feem peb ntawm cov genome yog cov khoom lag luam hloov pauv. Lub genome ntawm P. infestans muaj ntau tsev neeg sib txawv ntawm transposons dua li lwm cov neeg paub txog genomes. Feem ntau ntawm P. infestans transposons feem rau Gypsy tsev neeg.
Coob tus ntawm cov tsev neeg tshwj xeeb muab kev koom tes hauv pathogenesis tau txheeb pom hauv P. infestans genome. Ib feem tseem ceeb ntawm lawv encode effector protein uas hloov cov physiology ntawm tus tswv tsev cog qoob loo thiab pab txhawb kom nws kis tau. Lawv poob rau hauv ob pawg dav: apoplastic effectors, uas ua nyob rau hauv qhov chaw sib cais (apoplasts), thiab cytoplasmic effectors, uas nkag rau hauv hlwb ntawm haustoria. Apoplastic effectors muaj zais zais cov hydrolytic enzymes xws li proteases, lipases thiab glycosylases uas rhuav tshem cov nroj tsuag hlwb; cov tshuaj tiv thaiv kev tiv thaiv cov nroj tsuag tiv thaiv enzymes; thiab necrotizing co toxins xws li Nep1-zoo li cov protein (NPLs) thiab Pcf-zoo li me me cysteine-nplua nuj protein (SCRs).
P. infestans effector noob yeej muaj ntau thiab feem ntau loj dua li cov noob tsis-muaj. Cov paub zoo tshaj plaws yog cov cytoplasmic effectors RXLR thiab Crinkler (CNR). Cov khaub ncaws cytoplasmic zoo tshaj plaws ntawm oomycetes yog RXLR cov protein. Txhua RXLR effector cov noob tau tshawb pom kom deb li deb muaj pawg amino-terminal pab pawg Arg-XLeu-Arg, qhov twg X yog cov amino acid. Raws li qhov kev tshawb nrhiav, nws tau hais tias muaj 563 RXLR caj ces hauv P. infestans genome, uas yog 60% ntau dua hauv P. sojae thiab P. ramorum. Kwv yees li ib nrab ntawm RXLR caj ces hauv P. infestans genome yog hom tsiaj tshwj xeeb. RXLR effectors muaj ntau yam kev sib txawv ntawm ntu. Ntawm lawv, ib qho loj thiab 150 tsev neeg me tau pom. Tsis zoo li cov khoom tiv thaiv tseem ceeb, RXLR effector cov noob feem ntau nyob hauv cov gene-tsis zoo thiab rov ua cov nplua nuj nyob hauv cov genome. Lub xov tooj ntawm tes uas txiav txim siab qhov kev hloov pauv ntawm cov cheeb tsam no txhawb txoj kev rov ua dua nyob hauv cov noob.
Cytoplasmic CRN txiaj ntsig tau pib ua lub npe nyob hauv P. infestans transcripts encode cog cov ntaub so ntswg necrosis peptides. Txij li thaum lawv nrhiav pom, me ntsis tau paub txog tsev neeg ntawm cov effectors no. Kev tsom xam ntawm P. Infestans genome qhia ib tsev neeg loj heev ntawm 196 CRN caj, uas yog loj dua li ntawm P. sojae (100 CRN) thiab P. ramorum (19 CRN). Zoo li RXLRs, CRNs yog cov protein ntau thiab muaj kev tiv thaiv zoo N-davhlau LFLAK (50 amino acids) thiab ib qho DWL sau uas nyob ib sab muaj cov noob sib txawv. Feem ntau CRNs (60%) muaj lub teeb liab peptide.
Muaj peev xwm ntawm ntau CRNs los cuam tshuam cov txheej txheem ntawm tes ntawm lub tswv tsev cog tseg tau kawm. Thaum txheeb xyuas cov nroj tsuag necrosis, qhov kev tshem tawm ntawm CRN2 cov kab mob ua rau nws muaj peev xwm txheeb xyuas thaj chaw C-cov cheeb tsam uas muaj 234 amino acids (txoj haujlwm 173-407, DXG sau npe) thiab ua rau lub cell tuag. Kev tsom xam ntawm P. infestans CRN noob qhia txog plaub thaj chaw sib txawv ntawm C-davhlau ya nyob twg, uas tseem ua rau lub cell tuag nyob rau hauv tsob ntoo. Cov no suav nrog cov DC tshiab tau txheeb (P. Infestans muaj 18 noob thiab 49 pseudogenes), nrog rau D2 (14 thiab 43) thiab DBF (2 thiab 1) cov npe uas zoo ib yam li cov kinases protein. Proteins ntawm CRN domains qhia nyob rau hauv ib tsob nroj yog khaws cia (thaum tsis muaj lub teeb liab peptides) hauv cov xovtooj ntawm cov nroj tsuag thiab txhawb kev tuag ntawm tes los ntawm kev ua haujlwm ntawm intracellular mechanism. Lwm 255 ntu uas muaj CRN domains feem ntau yuav tsis ua haujlwm raws caj ces.
Qhov kev nce ntawm tus naj npawb thiab qhov loj ntawm RXLR thiab CRN effector cov tsev neeg tau txais vim yog qhov tsis muaj vim tias tsis muaj lub zog tsis sib xws thiab rov ua dua tshiab. Dua li ntawm qhov tseeb hais tias cov genome muaj ntau ntawm cov xov tooj txawb nquag, tseem tsis muaj cov pov thawj ncaj qha ntawm kev hloov chaw ntawm cov muaj txiaj ntsig cov noob.
Cov hau kev siv hauv kev kawm txog cov qauv pej xeem
Txoj kev kawm txog keeb kwm caj ces ntawm cov pej xeem tam sim no nyob ntawm kev txheeb xyuas cov ntshiab kab lis kev cai ntawm nws cov kab mob sib kis. Kev ntsuam xyuas ntawm cov neeg tsis muaj cais cov kev coj noj coj ua tseem ua rau lub hom phiaj tshwj xeeb, xws li, piv txwv, kawm txog kev ua nruj ua tsiv ntawm cov pej xeem los yog muaj cov kab mob tsis haum rau cov fungicides hauv nws (Filippov et al., 2004; Derevyagina li al., 1999). Hom kev tshawb fawb no koom nrog kev siv cov hau kev tshwj xeeb, qhov kev piav qhia uas dhau dhau ntawm qhov ntsuas ntawm kev tshuaj xyuas no. Rau qhov sib piv ntawm cov kab mob sib txawv, ntau txoj kev siv, raws ob qho tib si ntawm kev tsom xam ntawm cov qauv DNA thiab ntawm kev kawm txog phenotypic manifestations. Kev sib piv ntawm cov pejxeem tau hais txog ntau qhov sib cais, uas yuav tsum tau ua raws li qee qhov kev xav tau ntawm cov txheej txheem kev siv. Qhov tsim nyog, lawv yuav tsum ua raws li cov cai hauv qab (Cooke, Lees, 2004, Mueller, Wolfenbarger, 1999):
- pheej yig, yooj yim rau kev siv, tsis tas yuav siv sijhawm tseem ceeb, ua raws li cov thev naus laus zis feem ntau muaj (piv txwv li PCR);
- yuav tsum tsim kom muaj cov neeg nyob sab nraud ntawm codominant cov cim ywj pheej;
- muaj kev rov ua dua siab;
- siv cov ntaub so ntswg tsawg kawg los kuaj;
- qhia meej rau cov substrate (qhov sib kis hauv kab lis kev cai yuav tsum tsis txhob cuam tshuam rau cov txiaj ntsig);
- tsis tas yuav siv cov txheej txheem tsim kev kub ntxhov thiab tshuaj lom ntau.
Hmoov tsis zoo, tsis muaj txoj hau kev txuam nrog rau tag nrho cov ntsuas dhau los. Rau qhov sib piv kawm txog cov kab mob hauv peb lub sijhawm, cov txheej txheem raug siv los ntawm kev soj ntsuam ntawm phenotypic cov cwj pwm: virulence rau cov qos yaj ywm thiab hom txiv lws suav (cov qos yaj ywm thiab soob), hom mating, spectra ntawm peptidase isoenzymes thiab glucose-6-phosphate isomerase, thiab ntawm kev tsom xam ntawm DNA qauv: ntev polymorphism cov kev txwv tsis pub tshaj (RFLP), uas feem ntau ntxiv nrog kev ua tiav ntawm kev sojntsuam RG 57, kev sojntsuam ntawm microsatellite rov ua dua (SSR thiab InterSSR), amplification nrog random primers (RAPD), amplification ntawm cov kev txwv txiav (AFLP), amplification nrog primers homologous mus rau ntu ntawm cov khoom txawb (piv txwv li, Inter SINE PCR), kev txiav txim siab ntawm mitochondrial DNA haplotypes.
Cov lus qhia luv luv ntawm cov hau kev ntawm kev sib piv kawm ntawm cov leeg siv hauv kev ua haujlwm nrog P. Infestans
Phenotypic cim ntaus cim rau
"Qos yaj" kev sib tw
Cov qos yaj ywm "cov qos yaj ywm" feem ntau yog cov kev tshawb nrhiav thiab siv cim. Cov qos yaj ywm uas yooj yim muaj ib lub noob rau cov qos, "cov nyom" ntau - tsawg kawg yog ob qhov. Dub li al. (1953), suav sau tag nrho cov ntaub ntawv muaj rau lawv, pom tias cov haiv neeg phytophthora muaj peev xwm kis cov nroj tsuag nrog cov thev naus laus zis / cov noob sib xyaw nrog cov P. infestans virulence gene / noob, thiab pom cov haiv neeg 1, 2, 3, thiab 4 uas kis rau cov nroj tsuag nrog cov noob R1, R2, R3 thiab R4, feem, i.e. kev sib cuam tshuam ntawm cov cab thiab tus tswv tsev tshwm sim raws li cov gene rau lub hauv paus ntsiab lus. Ntxiv mus, Dub, nrog kev koom tes ntawm Gallegly thiab Malcolmson, nrhiav cov tsis kam cov caj ces R5, R6, R7, R8, R9, R10 thiab R11, nrog rau haiv neeg sib xws (Dub, 1954; Dub & Gallegly, 1957; Malcolmson & Dub, 1966; Malcolmson, 1970).
Muaj ntau lub cev ntawm cov ntaub ntawv ntawm cov tawv nqaij muaj pes tsawg leeg ntawm cov pathogen los ntawm thaj chaw sib txawv. Yog tias tsis txheeb xyuas cov ntaub ntawv no kom paub meej, peb yuav qhia tsuas yog qhov dav dav: qhov twg ntau yam nrog cov tshiab tiv thaiv cov noob los yog lawv cov kev sib txuas tau siv, thaum xub thawj muaj qee qhov tsis muaj zog ntawm lig blight, tab sis tom qab ntawd kev sib tw nrog cov sib deev sib deev tau tshwm sim thiab raug xaiv thiab tawm ntawm cov blight rov pib dua. Qhov tshwj xeeb virulence tawm tsam thawj 4 hom kev tawm tsam (R1-R4) tsis tshua pom muaj nyob hauv kev sib sau ua ntej kev qhia txog kev cog qoob loo ntau yam nrog cov noob, tab sis tus naj npawb ntawm cov virulent lim ntau ntxiv thaum cov kab mob parasitized ntawm ntau yam uas nqa cov noob. Koob 5-11, ntawm qhov tod tes, yog qhov muaj ntau nyob rau hauv kev sib sau (Shaw, 1991).
Kev tshawb fawb ntawm qhov sib piv ntawm cov sib txawv ntawm cov haiv neeg sib txawv thaum lub caij cog qoob loo, ua nyob rau lub sijhawm xyoo 1980s, qhia tau tias thaum pib ntawm kev txhim kho tus kabmob, cov pob txha tsis muaj kev txhoj puab thiab 1-2 virulence noob predominate hauv cov pejxeem.
Ntxiv mus, nrog rau kev txhim kho ntawm cov blight lig, qhov kev tsub zuj zus ntawm cov thawj cov pob zeb tsawg zuj zus thiab tus naj npawb ntawm cov "kev sib tw" nrog cov kev siab phem nce ntxiv. Qhov tshwm sim tom qab kawg los ntawm qhov kawg ntawm lub caij nce mus txog 100%. Thaum khaws cov qe, muaj qhov txo qis ntawm kev txhoj puab heev thiab poob ntawm tus kheej cov virulence noob. Lub zog ntawm clone hloov tuaj yeem tshwm sim hauv ntau hom sib txawv (Rybakova & Dyakov, 1990). Txawm li cas los xij, peb cov kev tshawb fawb nyob rau xyoo 2000-2010 tau qhia tias cov kev sib tw nyuaj tau pom los ntawm thaum pib ntawm epiphytotics ntawm cov kab mob sib cais los ntawm ob qho tib si qos yaj ywm thiab txiv lws suav. Qhov no tej zaum vim muaj kev hloov pauv ntawm cov neeg ntawm P. Infestans hauv tebchaws Russia.
Txog xyoo 1988-1995, qhov ntau zaus tshwm sim ntawm "superraces" nrog txhua lossis tag nrho cov virulence genes hauv thaj tsam sib txawv tau mus txog 70-100%. Cov xwm txheej zoo li no tau sau tseg, piv txwv li, hauv Belarus, hauv thaj chaw Leningrad, Moscow, hauv North Ossetia thiab hauv Tebchaws Yelemees (Ivanyuk et al., 2002a, 2002b; Politiko, 1994; Schober-Butin et al., 1995).
"Soob" kev sib tw
Hauv cov noob txiv lws suav, tsuas yog 2 lub noob ntawm kev tsis kam mus txog tom qab blight tau pom - Ph1 (Gallegly & Marvell, 1955) thiab Ph2 (Al-Kherb, 1988). Xws li cov qos yaj ywm sib tw, kev sib cuam tshuam ntawm cov txiv lws suav thiab P. infestans tshwm sim ntawm cov noob-noob-noob. Kev sib tw T0 kis tau ntau yam uas tsis muaj cov tshuaj tiv thaiv cov noob (feem ntau ntawm cov lag luam siv ntau yam), cov haiv neeg T1 kis tau ntau yam nrog cov Ph1 gene (Ottawa), thiab T2 haiv neeg - ntau yam nrog cov gene Ph2.
Hauv tebchaws Russia, yuav luag tshwjxeeb T0 pom ntawm cov qos yaj ywm; T0 tau kwv yees rau cov txiv lws suav thaum pib ntawm lub caij, tab sis tom qab ntawd nws hloov los ntawm kev sib tw T1 (Dyakov li al., 1975, 1994). Tom qab 2000, T1 rau cov qos yaj ywm hauv ntau tus neeg tau pib tshwm sim thaum pib ntawm lub sijhawm epiphytotic. Hauv Tebchaws Asmeskas, cov noob qoob loo tsis yog-pathogenic rau lws suav, nrog rau kev sib tw T0, T1, thiab T2, thaum T1 thiab T2 tau yeej rau cov txiv lws suav (Vartanian & Endo, 1985; Goodwin li al., 1995).
Mating type
Los ua qhov kev tshawb fawb, tester (kev siv) cov kab mob uas paub hom kev ua niam txiv - A1 thiab A2 - yog qhov yuav tsum tau ua. Qhov ntsuas cais tawm yog inoculated nrog lawv hauv khub hauv Petri cov tais diav nrog oat agar nruab nrab. Tom qab tsim kom loj hlob nyob rau 10 hnub, daim hlau tau soj ntsuam seb puas muaj qhov q lossis oospores hauv nruab nrab hauv thaj chaw sib chwv ntawm cov leeg. Muaj 4 txoj kev xaiv: txoj haujlwm yog hom A1 mating hom, yog tias nws tsim oospores nrog cov A2 tester, rau A2, yog tias nws tsim oospores nrog cov A1 tester, rau A1A2, yog tias nws tsim oospores nrog ob tus kuaj, lossis yog huv (00), yog tias nws tsis ua oospores nrog tsis muaj tus simter (ob pab pawg kawg yog tsawg).
Txhawm rau txiav txim siab ntau dua ntawm hom kev ua niam txiv, kev sim tau txheeb xyuas thaj tsam ntawm cov genome cuam tshuam nrog hom mating, nrog lub hom phiaj ntawm lawv qhov kev siv txuas ntxiv los txiav txim siab hom kev sib raug zoo los ntawm PCR. Ib qho ntawm cov kev sim ua tiav thawj zaug los txheeb xyuas qhov chaw zoo li no tau ua los ntawm cov kws tshawb fawb Asmeskas (Judelson et al., 1995). Siv tus qauv RAPD, lawv muaj peev xwm txheeb xyuas lub cheeb tsam W16 txuam nrog hom mating hauv cov xeeb ntxwv ntawm ob txoj kev hla dhau, thiab tsim ib khub ntawm 24-bp primers kom loj dua nws (W16-1 (5'-AACACGCACAAGGCATATAAATGTA-3 ') thiab W16-2 (5' -GCGTAATGTAGCGTAACAGCTCTC-3 ') Tom qab txwv PCR cov khoom lag luam nrog cov kev txwv enzyme HaeIII, nws muaj peev xwm cais kev sib cais nrog kev sib piv hom A1 thiab A2.
Lwm qhov kev sim ua kom tau txais PCR cov cim los txiav txim siab txog hom kev sib raug zoo tau ua los ntawm cov kws tshawb nrhiav Kauslim (Kim, Lee, 2002). Lawv txheeb xyuas qee yam khoom siv cov qauv AFLP. Raws li qhov tshwm sim, ib khub ntawm cov ntawv PHYB-1 (rau pem hauv ntej) (5'-GATCGGATTAGTCAGACGAG-3 ') thiab PHYB-2 (5'-GCGTCTGCAAGGCGCATTTT-3') tau tsim, tso cai xaiv kom muaj suab nrov ntawm cov cheeb tsam genome cuam tshuam nrog hom A2 mating. Tom qab ntawd, lawv txuas ntxiv cov haujlwm no thiab tsim cov qauv siv 5 'AAGCTATACTGGGACAGGGT-3' (INF-1, rau pem hauv ntej) thiab 5'-GCGTTCTTTCGTATTACCAC-3 '(INF-2), tso cai xaiv kom nrov ntawm thaj av Mat-A1 cov yam ntxwv ntawm cov leeg nrog hom mating. A1. Kev siv PCR kuaj mob ntawm kev ua niam txiv tau pom tias muaj txiaj ntsig zoo thaum kawm txog cov neeg ntawm P. infestans hauv Czech koom pheej (Mazakova et al., 2006), Tunisia (Jmour, Hamada, 2006), thiab lwm thaj chaw. Hauv peb lub chaw kuaj (Mytsa, Elansky, tsis tshaj tawm), 34 P. cov kab mob kis los ntawm cov kab mob qos thiab lws suav cov kabmob ntawm thaj chaw sib txawv ntawm Russia (Kostroma, Ryazan, Astrakhan, Moscow oblasts) tau tshawb xyuas. Cov txiaj ntsig ntawm kev ntsuas PCR siv cov kev ntsuas tshwj xeeb ntau dua 90% zoo dua nrog cov txiaj ntsig ntawm kev tsom xam ntawm hom kev ua niam txiv los ntawm cov txheej txheem ib txwm nyob ntawm nruab nrab cov khoom noj khoom haus.
Cov lus 1. Qhov txawv ntawm tsis kam tiv taus nyob hauv Tus Sib Koom 1 clone (Elansky et al., 2001)
Qauv sau qhov chaw | Tus naj npawb ntawm cais cais | Tus naj npawb ntawm rhiab (S), tsis muaj zog tiv taus (SR) thiab tsis kam ua haujlwm (R) lim, pcs (%) | ||
S | SR | R | ||
G. Vladivostok | 10 | 1 (10) | 4 (40) | 5 (50) |
G. Chita | 5 | 0 | 0 | 5 (100) |
Irkutsk | 9 | 9 (100) | 0 | 0 |
G. Krasnoyarsk | 13 | 12 (92) | 1 (8) | 0 |
Yekaterinburg lub nroog | 15 | 8 (53) | 1 (7) | 6 (40) |
O. Sakhalin | 66 | 0 | 0 | 66 (100) |
Thaj av Omsk | 18 | 0 | 0 | 18 (100) |
Metalaxyl tsis kam ua raws li cov pej xeem lub cim
Thaum xyoo 1980s, kev tawm suab nrov ntawm cov blight lig tshwm sim los ntawm metalaxyl-resistant P. cov kab mob tau pom nyob hauv ntau thaj tsam. Cov qoob loo qos yaj ywm hauv ntau lub teb chaws tau poob ntau heev (Dowley & O'Sullivan, 1981; Davidse et al., 1983; Derevyagina, 1991). Txij thaum ntawd los, hauv ntau lub teb chaws ntawm lub ntiaj teb, kev tshuaj xyuas tas li ntawm qhov tshwm sim ntawm phenylamide hom kab mob hauv P. cov kab mob tsawg kawg tau ua. Ntxiv nrog rau kev soj ntsuam kev muaj tiag ntawm kev cia siab rau kev siv cov tshuaj phenylamide uas muaj cov tshuaj, tsim kom muaj lub zog ntawm kev tiv thaiv kev tiv thaiv thiab kev kwv yees txog mob epiphytoties, kev tawm tsam cov tshuaj no tau dhau los ua ib qho cim cim dav siv rau kev sib piv ntawm cov neeg ntawm tus kab mob no. Txawm li cas los xij, kev siv ntawm kev tawm tsam mus rau metalaxyl nyob rau hauv kev sib piv cov pej xeem kev tshawb fawb yuav tsum tau nqa tawm mus rau hauv tus account qhov tseeb tias: 1 - lub hauv paus caj ces ntawm kev tiv thaiv tseem tsis tau raug txiav txim siab meej meej, 2 - kev tawm tsam rau metalaxyl yog kev xaiv qhov sib txawv uas tuaj yeem hloov pauv raws li kev siv phenylamides, 3 - txawv cov theem ntawm rhiab heev rau metalaxyl lim tsis pub dhau ib qho clonal kab (rooj. 1).
Spectra ntawm isozymes
Isozyme cov cim cim yog feem ntau ywj siab ntawm cov xwm txheej sab nraud, qhia Mendelian cov qub txeeg qub teg thiab yog codominant, cia kom paub qhov txawv ntawm homo- thiab heterozygotes. Kev siv cov protein ua cov cim hluav taws xob ua rau nws muaj peev xwm txheeb xyuas ob qho tib si rov txhim kho caj ces ntawm caj ces, suav nrog chromosomal thiab kev hloov pauv hauv noob caj noob ces, thiab tib cov amino acid hloov chaw.
Electrophoretic cov kev tshawb fawb ntawm cov protein tau qhia tias cov enzymes feem ntau muaj nyob hauv cov tsiaj hauv daim ntawv ntawm ob peb feem sib txawv hauv electrophoretic kev txav mus los. Cov zauv feem yog qhov tshwm sim ntawm kev sau ntau hom ntawm cov enzymes los ntawm cov loci sib txawv (isozymes lossis isozymes) lossis los ntawm cov sib txawv ntawm txhua tus neeg hauv thaj chaw (allozymes lossis alloenzymes). Ntawd yog, isozymes yog cov sib txawv ntawm ib qho enzyme. Cov foos sib txawv muaj cov kev ua kom zoo tib yam, tab sis txawv me ntsis hauv cov amino acid hloov pauv hauv peptide thiab them. Xws li cov sib txawv tau pom thaum lub sijhawm electrophoresis.
Thaum kawm P. infestans lim ntshav, tsom iav ntawm isoenzymes ntawm ob cov protein, peptidase thiab piam thaj-6-phosphate isomerase, tau siv (cov enzyme no yog monomorphic hauv cov neeg Lavxias Txhawm rau cais lawv mus rau isozymes hauv thaj chaw hluav taws xob, cov khoom noj protein npaj cais los ntawm cov kab mob muaj nyob hauv kev tshawb fawb tau thov rau lub phaj gel tso rau hauv thaj chaw hluav taws xob. Tus nqi ntawm kev sib txawv ntawm cov nyias muaj nyias protein hauv cov gel nyob ntawm tus nqi thiab qhov hnyav phaus; yog li ntawd, hauv thaj chaw hluav taws xob, kev sib xyaw ntawm cov protein sib cais tau cais ua nyias feem, uas tuaj yeem pom tau siv cov dyes tshwj xeeb.
Txoj kev kawm txog peptidase isoenzymes yog nqa tawm ntawm cellulose acetate, hmoov txhuv nplej siab lossis polyacrylamide gels. Qhov yooj yim tshaj plaws yog cov qauv raws li kev siv cov cellulose acetate gels tsim los ntawm Helena Lub Chaw Ntsuas Hluav Taws Xob. Nws tsis tas yuav muaj ntau qhov ntsuas cov khoom siv, nws tso cai rau ib qho kom tau txais cov kab sib luag ntawm cov gel tom qab electrophoresis rau ob qho tag nrho cov enzyme loci, nws qhov kev siv tsis tas siv sijhawm loj thiab cov khoom siv (Daim duab 2).
Ib qho me me ntawm mycelium tau pauv mus rau hauv 1,5 ml microtube, 1-2 tee ntawm cov dej distilled ntxiv rau nws. Tom qab ntawd, tus qauv yog homogenized (piv txwv li, siv hluav taws xob xyaum nrog cov ntawv yas kom haum rau lub microtube) thiab sedimented rau 25 vib nas this ntawm lub centrifuge ntawm 13000 rpm. 8 μl los ntawm txhua lub microtube. lub supernatant pauv mus rau tus thov phaj.
Cov cellulose acetate gel tshem tawm ntawm lub thawv ntas dej, blotted ntawm ob daim ntawv lim ntawv lim thiab muab tso nrog txheej txheej ua haujlwm nyob rau sab saum toj yas txheej ntawm tus neeg thov kev pabcuam. Qhov kev daws teeb meem los ntawm cov phaj yog pauv los ntawm tus neeg thov rau lub zaws 2-4 zaug. Lub pob zeb tau pauv mus rau chav electrophoresis,
Cov lus 2. Cov sib xyaw ua ke ntawm cov tshuaj siv rau staining cellulose acetate gel hauv kev tsom xam ntawm peptidase isoenzymes, ib cov xim poob (bromophenol xiav) tau muab tso rau ntawm ntug ntawm cov gel.
TRIS HCl, 0,05M, Ph 8,0 2 ml
Peroxidase, 1000 U / ml 5 ncos
o-dianisidine, 4 mg / ml 8 tee
MgCl2, 20 mg / ml 2 tee
Gly-Leu, 15 mg / ml 10 tee
L-amino-acid oxidase, 20 u / ml 2 tee
Electrophoresis yog ua tiav rau 20 feeb. ntawm 200 V. Tom qab electrophoresis, cov gel hloov pauv mus rau lub rooj pleev xim thiab tha xim nrog cov tshuaj pleev xim tshwj xeeb (Cov Lus 2). 10 ml ntawm 1,6% DIFCO agar yog preliminarily melted nyob rau hauv lub qhov cub microwave, txias rau 60 ° C, tom qab ntawd 2 ml ntawm agar sib xyaw nrog cov xim sib xyaw thiab nchuav mus rau lub gel. Cov kab txaij tshwm hauv 15-20 feeb. L-amino-acid oxidase tshuaj ntxiv rau tsuas yog ua ntej tov tov nrog molten agar.
Hauv cov haiv neeg Lavxias, Pep 1 thaj chaw yog sawv cev los ntawm genotypes 100/100 thiab 92/100. Homozygote 92/92 tsis tshua muaj neeg los (kwv yees li 0,1%). Locus Rehr 2 yog sawv cev los ntawm peb lub genotypes 100/100, 100/112, thiab 112/112, thiab tag nrho 3 qhov txawv yog qhov muaj ntau (Elanky thiab Smirnov, 2003, Daim Duab 2).
Kev tshawb fawb genome
Kev txwv qhov ntev ntawm polymorphism nrog kev sib txuas ntxiv (RFLP-RG 57)
Tag nrho cov DNA tau kho nrog Eco R1 enzyme txwv, DNA tawg yog sib cais los ntawm electrophoresis hauv agarose gel. Nuclear DNA yog qhov loj heev thiab muaj ntau qhov kev rov ua dua, uas ua rau nws nyuaj rau ncaj qha soj ntsuam ntau tawg tsam tau los ntawm kev ua ntawm kev txwv cov enzymes. Yog li ntawd, cov DNA tawg sib cais hauv cov gel tau hloov mus rau ib daim nyias nyias tshwj xeeb thiab siv rau kev sib xyaw nrog RG 57 sojntsuam, uas suav nrog nucleotides daim ntawv lo nrog cov pa hluav taws xob lossis cov ntawv fluorescent. Qhov kev sojntsuam no sib txuas nrog cov kev rov ua kom zoo ibyam zuj zus (Goodwin li al., 1992, Forbes li al., 1998). Tom qab ua kom pom tseeb ntawm cov txiaj ntsig ntawm kev sib xyaw ntawm lub teeb ci- lossis cov khoom siv hluav taws xob, ntau qhov chaw ntawm cov keeb kwm kev sib txuas ntawm tus kheej (ntiv tes ntiv tes) tau txais, sawv cev los ntawm 25-29 tawg tsam (Forbes li al., 1998). Asexual (clonal) cov xeeb ntxwv yuav muaj tib lub profiles. Los ntawm kev ua raws li cov kab ua ke nyob rau ntawm electrophoretogram, qhov zoo sib xws thiab qhov sib txawv ntawm cov kab mob sib piv tau txiav txim.
Mitochondrial DNA haplotypes
Hauv cov hlwb eukaryotic feem ntau, mtDNA tau nthuav tawm nyob rau hauv daim ntawv ntawm ob lub cev sib npaug ntawm cov xov tooj cua DNA, uas, tsis zoo li lub nuclear chromosomes ntawm eukaryotic hlwb, replicates semi-conservatively thiab tsis cuam tshuam nrog protein molecules.
Cov mitochondrial genome ntawm P. infestans tau ua tiav, thiab ib tug xov tooj ntawm cov haujlwm tau mob siab rau kev tsom xam ntawm kev txwv kev cais txog qhov ntev (Carter et al, 1990, Goodwin, 1991, Gavino, Fry, 2002). Tom qab Griffith and Shaw (1998) tau tsim cov qauv yooj yim thiab nrawm rau kev txiav txim siab mtDNA haplotypes, qhov cim no tau los ua ib qho ntawm cov nyiam tshaj plaws hauv P. Cov kev tshawb fawb tsawg tshaj plaws.Qhov qhov tseem ceeb ntawm cov qauv muaj nyob rau hauv qhov txuas ntxiv ntawm ob mitochondrial DNA tawg (los ntawm cov genome) nrog primers F2-R2 thiab F4-R4 (Cov Lus 3) thiab lawv cov kev txwv tom ntej nrog kev txwv nrog enzymes MspI (1st ntu) thiab EcoR1 (thib 2 puas). Cov txheej txheem tso cai rau koj txheeb xyuas 4 haplotypes: Ia, IIa, Ib, IIb. Hom II txawv ntawm hom I los ntawm qhov pom ntawm 1881 bp ntxig thiab qhov chaw sib txawv ntawm cov chaw txwv nyob hauv thaj tsam P2 thiab P4 (Daim duab 3).
Txij li thaum xyoo 1996, ntawm cov kev txhawm coj los sau rau thaj chaw ntawm teb chaws Russia, tsuas yog haplotypes Ia thiab IIa tau sau tseg (Elansky et al., 2001, 2015). Lawv tuaj yeem raug txheeb xyuas tom qab kev sib cais cov khoom txwv nrog primer F2-R2 hauv thaj chaw hluav taws xob (Daim duab 4, 5). Cov hom ntawm mtDNA tau muab siv rau hauv kev sib piv ntawm cov kab mob thiab ntau yam. Hauv ntau cov kev tshawb fawb, hom mitochondrial DNA tau siv los cais cov kab clonal thiab passportize P. infestans cais (Botez et al., 2007; Shein li al., 2009). Siv tus txheej txheem PCR-RFLP, nws tau xaus tias mtDNA yog cov kabmob sib npaug hauv tib hom P. infestans strain (Elansky and Milyutina, 2007). Cov kev ua kom txawv: 1x (500 sec. 94 ° C), 40x (30 sec. 90 ° C, 30 sec. 52 ° C, 90 sec. 72 ° C); 1x (5 min. 72 ° C). Cov tshuaj tiv thaiv kev sib xyaw: (20 μl): 0,2 U Taq DNA polymerase, 1x 2,5 mM MgCl2-Taq tsis, 0,2 mM txhua dNTP, 30 pM primer thiab 5 ng ntawm cov tshuaj ntsuam DNA, dej deionized - txog 20 μl.
Kev Txwv cov PCR cov khoom yog nqa nrog 4-6 teev ntawm qhov kub txog 37 ° C. Kev txwv kev sib xyaw (20 μl): 10x MspI (2 μl), 10x txwv tsis pub (2 μl), dej deionized (6 μl), PCR khoom (10 μl).
Cov Lus 3. Cov Qauv Siv tau los ua kom pom tseeb ntawm mtDNA polymorphic cheeb tsam
Tsom | Primer | Primer ntev thiab kev tso kawm | PCR khoom ntev | Txwv Tsis Pub Siv |
---|---|---|---|---|
P2 | F2: 5'- TTCCCTTTGTCCTCTACCGAT | 21; 13619-13639 Ib | 1070 | MspI |
R2: 5'- TTACGGCGGTTTAGCACATACA | 22; 14688-14667 Ib | |||
P4 | F4: 5'- TGGTCATCCAGGTTTATGTT | 22; 9329-9350 Ib | 964 | EcoRI |
R4:5 - CCGATACCAGATACCAGCACCAA | 22; 10292-10271 Ib |
Kev tso suab kom nrov (RAPD)
Thaum nqa tawm RAPD, ib qho tshuaj primer yog siv (qee zaus qee qhov pib ib txhij) nrog rau cov xaim hluav taws xob tsis xws luag, feem ntau 10 nucleotides nyob rau hauv ntev, nrog cov ntsiab lus siab (los ntawm 50%) ntawm GC nucleotides thiab tsis kub annealing kub (txog 35 ° C). Xws hom av "thaj av" rau ntau qhov chaw sib ntxiv hauv lub genome. Tom qab ua kom nrov dua, ntau ntau ntawm amplicons tau txais. Lawv tus lej yog nyob ntawm tus primer (s) siv thiab cov kev mob tshwm sim (MgCl2 concentration thiab annealing kub).
Kev xam pom ntawm amplicons yog nqa los ntawm distillation hauv polyacrylamide lossis agarose gel. Thaum ua kev soj ntsuam RAPD, nws yog ib qho tsim nyog yuav tsum tau ua tib zoo saib xyuas kom huv huv ntawm cov ntaub ntawv txheeb xyuas, txij li thaum kev sib kis nrog lwm cov khoom muaj sia nyob tuaj yeem ua rau muaj ntau qhov ntxiv ntawm cov khoom cuav, uas muaj ntau heev nyob rau hauv kev txheeb xyuas ntawm cov khoom ntshiab (Perez et al, 1998). Kev siv tus qauv no hauv kev kawm txog P. infestans genome yog pom hauv ntau yam haujlwm (Judelson, Roberts, 1999, Ghimire et al., 2002, Carlisle et al., 2001). Cov kev xaiv ntawm cov tshuaj tiv thaiv cov teeb meem thiab cov primers (51 10-nucleotide primers tau kawm) tau muab rau hauv tsab xov xwm los ntawm Abu-El Samen et al., (2003).
Kev Ntsuam Xyuas Rov Ntsuas Dua (SSR)
Cov me me rov qab ua dua tshiab (qhov tsis yooj yim ua kom rov ua dua, SSR) yog qhov ua kom luv dua ntawm theem 1-3 ntawm qee zaum (qee zaum txog 6) nucleotides tam sim no hauv cov hluav taws xob nuclear ntawm txhua qhov eukaryotes. Tus naj npawb ntawm cov pheej rov ua dua tshiab tuaj yeem hloov ntawm 10 txog 100. Lub zos Microsat satellite tshwm sim nrog qhov ntsuas siab thiab muaj ntau dua los sis tsawg dua li faib thoob plaws hauv lub genome (Lagercrantz li al., 1993). Polymorphism ntawm microsat satellite kab ke yog cuam tshuam nrog qhov sib txawv ntawm cov xov tooj ntawm qhov rov ua dua ntawm qhov pib motif. Cov cim cim me me yog codominant, uas ua rau nws siv tau rau kev txheeb xyuas cov pej xeem cov qauv, txiav txim siab kev sib koom tes, cov qauv txav ntawm lub cev, thiab lwm yam. Ntawm lwm qhov zoo ntawm cov cim no, ib qho yuav tsum nco ntsoov lawv cov polymorphism siab, zoo reproducibility, nruab nrab, thiab muaj peev xwm ua tsis siv neeg tsom xam thiab ntsuas .Analysis ntawm polymorphism ntawm microsatellite rov ua dua yog siv los ntawm PCR amplification siv primers ntxiv rau cov kab sib txawv ntawm lub microsatellite loci. Thaum pib, kev tsom xam tau ua nrog kev sib cais ntawm cov tshuaj tiv thaiv cov khoom ntawm polyacrylamide gel. Tom qab ntawd, cov neeg ua haujlwm ntawm Applied Biosystems tau npaj siab siv cov tshuaj fluorescently sau npe nrog rau kev tshawb nrhiav cov khoom tshuaj tiv thaiv uas siv lub tshuab hluav taws xob tsis siv neeg (Diehl li al., 1990), thiab tom qab ntawd tus txheej txheem automatic DNA sequencers (Ziegle et al., 1992). Sau npe ntawm cov ntawv pleev xim nrog ntau cov roj fluorescent ua rau nws muaj peev xwm txheeb xyuas ntau lub cim ntawm ib zaug ntawm ib txoj kab thiab, raws li, nce qhov khoom lag luam ntawm tus qauv thiab nce qhov tseeb ntawm qhov kev tsom xam.
Thawj cov ntawv tshaj tawm tau hais txog kev siv SSR kev soj ntsuam rau qhov kev kawm P. infestans tau tshwm sim thaum 2000s. (Knapova, Gisi, 2002). Tsis yog txhua qhov cim sau tseg los ntawm cov sau phau ntawv qhia tau pom tias muaj qib siab ntawm polymorphism, txawm li cas los xij, ob ntawm lawv (4B thiab G11) tau muaj nyob rau hauv cov teeb tsa ntawm 12 SSR cov cim sau los ntawm Lees et al. (2006) thiab tom qab tau saws hauv Eucablight tshawb fawb network (www.eucablight .org) raws li tus qauv rau P. infestans. Ob peb xyoos tom qab, kev tshawb fawb tau tshaj tawm txog kev tsim lub system rau kev soj ntsuam cov kab mob sib txuam ntawm P. infestans DNA raws li yim lub cim SSR (Li et al., 2010). Thaum kawg, tom qab ntsuam xyuas tag nrho cov ntawv sau tseg ua ntej lawm thiab xaiv cov tshaj plaws ntawm lawv, nrog rau ua kom zoo dua qub, ntawv cim fluorescent, thiab cov kev ua kom nrov zog, tib pawg kws sau ntawv tau tshaj tawm cov txheej txheem ntawm ib-kauj ruam multiplex tsom xam, suav nrog 12 tus cim (Cov Lus 4; Li li al. , 2013a). Cov primers siv nyob rau hauv cov kab ke no tau xaiv thiab cim npe nrog ib ntawm plaub qhov cim xim fluorescent (FAM, VIC, NED, PET) kom cov thaj tsam ntawm cov allele ntau thiab tsawg ntawm cov ntawv luam tawm qub uas muaj cov ntawv sau sib tshooj tsis sib tshooj.
Cov kws sau ntawv ua qhov kev soj ntsuam ntawm PTC200 amplifier (Kev Tshawb Fawb MJ, Tebchaws Asmeskas) siv QIAGEN multiplex PCR cov khoom siv lossis QIAGEN Typeit Microsatellite PCR cov khoom siv. Qhov ntim ntawm qhov sib xyaw cov tshuaj tiv thaiv yog 12.5 μL. Cov kev ua kom nrov zog muaj raws li nram no: rau QIAGEN multiplex PCR: 95 ° C (15 min), 30x (95 ° C (20 s), 58 ° C (90 s), 72 ° C (60 s), 72 ° C (20 min); rau QIAGEN Hom-nws Microsatellite PCR: 95 ° C (5 min), 28x (95 ° C (30 sec), 58 ° C (90 sec), 72 ° C (20 sec), 60 ° C (30 feeb).
Cais cais thiab kev pom tseeb ntawm PCR cov khoom tau ua tiav siv ABI3730 tsis siv neeg capillary DNA tsom xam (Siv Biosystems).
Cov Lus 4. Tus yam ntxwv ntawm 12 tus cim SSR cim siv rau kev tua neeg ntawm P. Infestans (Li et al., 2013a)
Lub npe | Tus naj npawb ntawm alleles | Qhov loj me alleles (bp) | Ntxhoo |
PiG 11 | 13 | 130-180 | F: NED-TGCTATTTATCAAGCGGGGG R: GTTTCAATCTGCAGCCGTAAGA |
Pib 02 | 4 | 255-275 | F: NED-ACTTGCAGAACTACCGCCCC R: GTTTGACCACTTTCCTTGTTTC |
PinfSSR11 | 4 | 325-360 | F: NED-TTAAGCCACGACATGAGCTG R: GTTTAGACAATTGTTTGTGGTCGC |
D13 | 16 | 100-185 | F: FAM-TGCCCCCTGCTCACTC R: GCTCGAATTCATTTACAGACTTG |
PinfSSR8 | 4 | 250-275 | F: FAM-AATCTGATCGCAACTGAGGG R: GTTTACAAGATACACACTCTCGCTCC |
PinfSSR4 | 7 | 280-305 | F: FAM-TCTTGTTCGAGTATGCGACG R: GTTTCACTTCGGAGAAAAGGCTTC |
Pib 04 | 4 | 160-175 | F: VIC-AGCGGCTACCGATGG R: GTTTCAGCGGCTTTCGAC |
Pib 70 | 3 | 185-205 | F: VIC-ATGAAAATCGTTCAATGCTCG R: CGTTGGAATATTTTTTTTTTCG |
PinfSSR6 | 3 | 230-250 | F: GTTTTGGTGGGGCTGAAGTTTT R: VIC-TCGCCACAAGATTTATTCCG |
Pib 63 | 3 | 265-280 | F: VIC-ATGACGAAGATGAAAGTGAGG R: CGTATTTTCCTGTTATCTAACACC |
PinfSSR2 | 3 | 165-180 | F: PET-CGACTTCTTACATCAACCGGC R: GTTTGTTTGGACTGCGTCTTTAGC |
Pi4B | 5 | 200-295 | F: PET-AAAAAAAGCCTTTTGGTTCA R: GCAAGCGAGGTTTGTAGATTT |
Ib qho piv txwv ntawm kev pom kev tsom xam cov txiaj ntsig tau tshwm sim hauv Daim Duab. 6. Cov txiaj ntsig tau soj ntsuam tau siv GeneMapper 3.7 software los ntawm kev sib piv cov ntaub ntawv tau nrog cov ntaub ntawv ntawm cov kev paub cais. Txhawm rau pab txhawb kev txhais lus ntawm cov ntsiab lus ntsuam xyuas, nws yog ib qho tsim nyog yuav tsum muaj xws li 1-2 kev siv cais nrog ib tus paub hauv keeb kwm hauv txhua qhov kev kawm.
Cov qauv kev tshawb fawb tau sim rau ntau qhov ntawm cov qauv ntsuas tom qab, tom qab ntawd tus sau phau ntawv tus qauv kev cai ntawm chav chaw kuaj mob ntawm ob lub koomhaum, Lub Koom Haum James Hutton (UK) thiab Wageningen University & Tshawb Xyuas (Netherlands), uas, nrog rau kev siv cov ntawv FTA txheem rau kev yooj yim kev sau thiab kev xa khoom ntawm P. infestans DNA cov qauv coj los ua rau nws muaj peev xwm los tham txog qhov ua tau ntawm kev ua lag luam siv ntawm qhov kev txhim kho no. Ib qho ntxiv, txoj kev ceev thiab raug ntawm genotyping P. infestans cais siv cov kev soj ntsuam multiplex SSR ua rau nws muaj peev xwm los ua cov kev tshawb fawb txog cov neeg ntawm cov kab mob no rau lub ntiaj teb, thiab kev tsim lub ntiaj teb cov ntaub ntawv ntawm lig blight nyob rau hauv lub moj khaum ntawm qhov project Eucablight (www.eucablight.org), suav nrog , suav nrog cov txiaj ntsig ntawm kev tsom xam microsatellite, ua rau nws muaj peev xwm taug qab qhov tshwm sim thiab kis ntawm cov genotypes tshiab thoob ntiaj teb.
Amplified kev txwv kev ncua ntev polymorphism (AFLP). AFLP (amplified fragment ntev polymorphism) yog thev naus laus zis rau kev tsim cov pa roj carbon monoxide hom cim uas siv qee yam tshwj xeeb. Hauv AFLP, DNA raug kho nrog ob txoj kev txwv nrog cov ntshav siab. Cov hloov kho tshwj xeeb yog lig rau cov nplaum tas ntawm cov kev txwv.
Cov thooj ntawv no tom qab ntawd nthuav dav siv cov primers ntxiv rau qhov hloov chaw txuas ntxiv thiab txwv chaw, thiab ntxiv ib qho nqa ib lossis ntau ntau lub hauv paus ntawm lawv 3 'xaus. Cov kev tawg tsam tau txais yog nyob ntawm kev txwv cov enzymes thiab xaiv xaiv cov nucleotides ntawm 3'-xaus ntawm cov primers (Vos li al., 1995). AFLP - kev tso tawm genotyping yog siv los kawm sai sai ntawm kev hloov caj ces ntawm ntau yam muaj sia.
Cov lus piav qhia ntxaws txog cov qauv yog muab nyob rau hauv kev ua haujlwm ntawm Mueller, Wolfenbarger, 1999, Savelkoul li al., 1999. Ntau ntau cov haujlwm ua haujlwm piv rau kev daws teebmeem ntawm AFLP thiab SSR cov hau kev tau ua tiav los ntawm cov kws tshawb nrhiav hauv Suav teb. Cov phenotypic thiab cov cim ntawm genotypic ntawm 48 P. infestans cais sau nyob rau tsib cheeb tsam ntawm North China tau kawm. Raws li AFLP spectra, yim qhov sib txawv ntawm DNA genotypes tau txheeb xyuas, hauv kev sib piv ntawm SSR genotypes, uas tsis muaj ntau haiv neeg tau tshwm sim (Guo et al., 2008).
Ua kom haum nrog primers homologous mus rau kab txuas ntawm cov khoom txawb
Cov cim cim tau los ntawm cov txheej txheem ntawm cov txheej txheem retrotransposons yog qhov yooj yim rau kev qhia txog caj ces, kev kawm txog kev tshuaj ntsuam caj ces thiab cov kev hloov pauv (Schulman, 2006). Yog tias qhov pib tau tsim los ua kom tiav cov qib ruaj khov ntawm qee lub xov tooj txawb, nws muaj peev xwm ua kom nrov nrov ntawm cov cheeb tsam genome nyob ntawm lawv. Nyob rau hauv kev tshawb fawb ntawm tus neeg sawv cev causative ntawm lig blight, tus qauv ntawm kev ua kom nrov nrov ntawm cov cheeb tsam genome siv cov txheej txheem sib txuas rau cov txheej txheem tseem ceeb ntawm SINE (Luv Interspersed Nuclear Elements) retroazone tau siv tiav (Lavrova thiab Elansky, 2003). Siv cov txheej txheem no, kev sib txawv tau qhia tawm txawm nyob rau hauv cov xeeb ntxwv asexual ntawm ib tus isolate. Hauv qhov no, nws tau xaus lus tias qhov inter - SINE - PCR txheej txheem yog qhov tshwj xeeb heev thiab tus nqi ntawm kev txav ntawm SINE cov ntsiab lus hauv Phytophthora genome yog tus siab.
Hauv cov genome ntawm P. infestans, 12 tsev neeg ntawm luv luv retrotransposons (SINEs) tau pom tias; cov tsiaj faib ntawm luv luv retrotransposons tau tshawb xyuas, cov khoom (SINEs) tau pom tias pom nyob hauv cov genome tsuas yog P. infestans (Lavrova, 2004).
Cov yam ntxwv ntawm kev siv ntawm cov hau kev ntawm txoj kev sib piv ntawm cov leeg hauv cov kev tshawb fawb pej xeem
Thaum npaj kev kawm, nws yog ib qho tsim nyog yuav tsum nkag siab meej txog cov hom phiaj uas nws ua tiav thiab siv cov hauv kev tsim nyog. Yog li, qee txoj kev cia ua kom muaj ntau ntawm qhov ua cim ywj pheej, tab sis tib lub sijhawm tsis tshua muaj kev rov ua dua thiab muaj kev cia siab rau cov khoom siv reagents, cov tshuaj tiv thaiv kev cuam tshuam, thiab cov khoom sib kis raws li kev kawm. Yog li, hauv txhua qhov kev tshawb fawb ntawm ib pawg ntawm cov neeg sib txawv, nws yog qhov yuav tsum tau siv ntau cov qauv (siv) kev cais, tab sis txawm tias qhov no, cov txiaj ntsig ntawm ntau qhov kev sim tau nyuaj rau kev sib txuas ua ke.
Pawg hom no suav nrog RAPD, AFLP, InterSSR, InterSINE PCR. Tom qab ua kom nrov dua, ntau ntawm DNA tawg ntawm cov ntau qhov sib txawv tau txais. Nws raug nquahu kom siv cov tswv yim zoo li no thaum tsim nyog los tsim cov kev sib txawv ntawm cov kab mob sib txawv (niam txiv-xeeb leej xeeb ntxwv, hom tsiaj qus, thiab lwm yam), lossis hauv rooj plaub thaum kev txheeb xyuas cov ncauj lus kom ntxaws ntawm cov qauv me me. Yog li, AFLP txheej txheem tau siv dav hauv kev teeb tsa caj ces ntawm P. infestans (van der Lee li al., 1997) thiab hauv kev tshawb fawb intrapopulation (Knapova, Gisi, 2002, Cooke li al, 2003, Flier et al, 2003). Cov hau kev no yog qhov tsis txaus ntseeg los siv thaum tsim cov ntaub ntawv ntawm cov kab mob puas lawm, txij li ntawd los nws yog qhov tsis yooj yim sua kom los ua ke cov kev txheeb xyuas cov txiaj ntsig thaum ua qhov kev tshuaj ntsuam hauv lub chaw ntsuas sib txawv.
Txawm hais tias zoo li qhov yooj yim thiab ceev ntawm kev tua (DNA cais tsis muaj qhov huv si, ua kom pom tseeb, pom lub txiaj ntsig), cov pab pawg ntawm cov hau kev no xav kom siv cov txheej txheem tshwj xeeb rau cov ntawv teev cov txiaj ntsig: distillation hauv polyacrylamide gel nrog cov ntawv sau (hluav taws xob los yog luminescent) primers thiab tom qab raug lub teeb lossis hluav taws xob. Kev ua kom zoo nkauj ethidium bromide agarose gel kev ntsuas tsis zoo rau cov qauv no vim tias coob ntawm DNA tawg ntawm ntau qhov sib txawv tuaj yeem fuse.
Lwm txoj hauv kev, ntawm qhov tsis sib xws, tso cai rau koj los ua kom muaj qee qhov me me ntawm cov nta nrog siab heev reproducibility. Pab pawg no suav nrog kev kawm txog mitochondrial DNA haplotypes (tsuas yog ob lub haplotypes Ia thiab IIa tau sau tseg hauv tebchaws Lavxias), hom mating (feem cais feem ntau tau faib ua 2 hom: A1 thiab A2, SF tus kheej tsis muaj tsawg) thiab peptidase isozyme spectra (ob lub zos Pep1 thiab Pep2 , suav nrog ob isozymes txhua tus) thiab piam thaj-6-phosphate isomerase (hauv tebchaws Russia tsis muaj qhov txawv txav rau tus cwj pwm no, txawm hais tias tseem ceeb polymorphism tau sau tseg hauv lwm lub tebchaws hauv ntiaj teb). Nws raug nquahu kom siv cov nta no thaum tshuaj ntsuam xyuas kev suav sau, sau cov ntaub ntawv nyob hauv ib cheeb tsam thiab thoob ntiaj teb. Kev soj ntsuam ntawm isozymes thiab haplotypes ntawm mitochondrial DNA, nws yog qhov ua tau yam tsis muaj qauv phuam txhua, thaum nyob hauv kev tsom xam ntawm hom mating, ob qhov ntsuas cais nrog paub hom mating yog qhov yuav tsum tau ua.
Cov kev mob tshwm sim thiab cov tshuaj reagents tuaj yeem tsuas yog cuam tshuam ntawm qhov sib txawv ntawm cov khoom ntawm lub electrophoretogram;
Tam sim no, feem ntau ntawm cov pejxeem nyob sab Europe feem ntawm Russia yog sawv cev los ntawm hom ntawm ob hom mating (Cov Lus 6), ntawm lawv muaj cais nrog hom Ia thiab IIa ntawm mitochondrial DNA (lwm hom mtDNA pom nyob hauv lub ntiaj teb tsis tau pom muaj nyob hauv Lavxias tom qab 1993). Cov yeeb yaj kiab ntawm peptidase isozymes yog sawv cev los ntawm ob lub genotypes ntawm Pep1 locus (100/100, 92/92 thiab heterozygote 92/100, thiab 92/92 caj ces tsis tshua muaj siab (<0,3%)) thiab los ntawm ob qhov genotypes ntawm Pep 2 locus (100/100 , 112/112 thiab heterozygote 100/112, nrog cov qauv ntawm 112/112 tshwm sim tsawg tshaj li 100/100, tab sis kuj muaj ntau zaus).
Nws tsis muaj qhov txawv txav ntawm cov teeb meem ntawm isoenzymes ntawm cov piam thaj-6-phosphate isomerase tom qab xyoo 1993 (kev ploj ntawm txoj kab clonal US-1); txhua qhov kev tshawb nrhiav cais tau muaj 100/100 caj ces (Elansky thiab Smirnov, 2002).
Cov pab pawg thib peb ntawm cov txheej txheem tso cai tau txais ib pab pawg txaus ntawm qhov ua cim ywj pheej nrog qhov siab rov ua dua. Niaj hnub no, cov pab pawg no suav nrog RFLP-RG57 kev sojntsuam, uas ua rau 25-29 DNA tawg ntawm ntau qhov sib txawv. RFLP-RG57 tuaj yeem siv tau ob qho tib si thaum ntsuas cov qauv thiab sau cov ntaub ntawv pov thawj. Txawm li cas los xij, hom no nws kim heev dua li cov dhau los, nws yog lub sijhawm siv sijhawm, thiab yuav tsum muaj qhov ntau txaus kom tiav cov DNA uas tau lim kom txaus. Yog li no, cov kws tshawb nrhiav txoj haujlwm yuam kom txwv cov ntim ntawm cov khoom siv kuaj xyuas.
Txoj kev loj hlob ntawm RFLP-RG57 nyob rau thaum ntxov 90s ntawm lub xyoo pua xeem ho pom cov kev tshawb fawb ntawm cov pej xeem ntawm cov neeg ua txhaum kev tuaj lig. Nws tau dhau los ua cov txheej txheem ntawm cov txheej txheem raws li xaiv thiab tsom xam ntawm "Clonal kab" (saib hauv qab). Nrog rau RFLP-RG57, mating hom, hom ntiv tes DNA (RFLP-RG57 txujci), tsom kwm ntawm peptidase thiab piam thaj-6-phosphate isomerase isoenzymes, thiab mitochondrial DNA hom yog siv los txheeb xyuas cov kab mob ntawm cov kab. Ua tsaug rau nws, nws tau qhia al., 1994), kev hloov pauv ntawm cov neeg laus los ntawm cov neeg tshiab (Drenth et al, 1993, Sujkowski et al, 1994, Goodwin et al, 1995a), thiab clonal lineages uas yeej nyob hauv ntau lub teb chaws ntawm lub ntiaj teb tau pom. Kev tshawb fawb ntawm Lavxias hom siv cov qauv no qhia tau lub siab genotypic polymorphism ntawm hom kab ke ntawm European feem thiab monomorphism ntawm cov neeg ntawm cov neeg Asian thiab Far Eastern ntawm Russia (Elansky et al, 2001). Thiab tam sim no hom no tseem yog qhov tseem ceeb hauv kev tshawb fawb pej xeem ntawm P. infestans. Txawm li cas los xij, nws txoj kev faib khoom dav yog qhov cuam tshuam los ntawm nws txoj kev nce nqi thiab kev siv zog hauv kev tua.
Lwm qhov kev cog lus zoo uas tsis tshua muaj siv hauv kev tshawb fawb P. infestans yog tsom tsom microsatellite (SSR). Tam sim no, txoj kev no yog dav siv los cais tawm cov kab clonal. Txog kev txheeb xyuas ntawm cov kab mob, xws li cov cim phenotypic cov cim vim muaj cov virulence noob rau cov qos yaj ywm (Avdey, 1995, Ivanyuk li al., 2002, Ulanova li al., 2003) thiab txiv lws suav tau siv dav (thiab txuas ntxiv siv). Los ntawm tam sim no, noob ntawm virulence rau qos ntau yam tau poob lawv cov nqi raws li cim cim rau cov pej xeem cov kev tshawb fawb vim muaj qhov tshwm sim ntawm qhov siab tshaj plaws (lossis ze rau nws) tus naj npawb ntawm virulence noob hauv feem ntau ntawm cais. Tib lub sijhawm, T1 virulence noob rau cov noob txiv lws suav coj tus nqi sib thooj Ph1 yog tseem muaj kev vam meej ua qhov cim kev coj (Lavrova li al., 2003; Ulanova li al., 2003).
Hauv ntau qhov kev tshawb fawb, fungicide tsis kam yog siv los ua cim. Qhov kev coj ua no yog qhov tsis xav siv nyob rau hauv cov kev tshawb fawb los ntawm cov pej xeem vim yog qhov yooj yim pom ntawm kev hloov pauv nyob hauv cov kab mob caj dab tom qab daim ntawv thov ntawm metalaxyl- (lossis mefenoxam-) muaj cov fungicides hauv thaj teb. Piv txwv li, qhov sib txawv tseem ceeb hauv theem ntawm kev tawm tsam tau qhia nyob rau hauv Sib1 clonal kab (Elansky et al., 2001).
Yog li, mating hom, peptidase isozyme spectrum, mitochondrial DNA hom, RFLP-RG57, SSR yog cov cim ntaus cim rau tsim cov txhab nyiaj cov ntaub ntawv thiab kev ntaus cim cov cim rau hauv kev suav sau. Txhawm rau piv txwv cov qauv txwv, yog tias nws yuav tsum tau siv ntau tshaj plaws ntawm cov cim cim, koj tuaj yeem siv AFLP, RAPD, InterSSR, Inter-SINE PCR (Cov Lus 5). Txawm li cas los xij, nws yuav tsum nco ntsoov tias cov hau kev no tsis tshua zoo rov ua dua qub, thiab hauv txhua tus neeg sim (ua kom lub zog hluav taws xob electrophoresis phaum), ntau qhov siv cais yuav tsum siv.
Rooj 5. Sib piv ntawm cov kev sib txawv ntawm kev tshawb fawb ntawm cov leeg P. Cov Kab Mob Txav
kauj | TS | Isofer tub ceev xwm | MtDNA | RFLP-RG57 | HLUA | ISSR | SSR | AFLP | Rev |
---|---|---|---|---|---|---|---|---|---|
Cov ntaub ntawv ntau | Н | Н | Н | С | В | В | С | В | В |
Luam tau rov qab | В | В | В | В | Н | Н | С | С | С |
Qhov ua tau ntawm cov khoom cuav | Н | Н | Н | Н | В | С | Н | С | В |
nqi ntawm | Н | С | Н | В | Н | Н | Н | С | Н |
Kev tawm dag zog | Н | Н | Н | В | NS * | NS * | Н | С | NS * |
Kev tsom xam ceev ** | В | Н | Н | С | Н | Н | Н | Н | Н |
Lus Cim: H - qis, C - nruab nrab, B - siab; НС * - kev siv ua haujlwm yog tsawg thaum siv agarose gel lossis automatic
genotyper, nruab nrab - los ntawm distillation nyob rau hauv polyacrylamide gel nrog sau cov ntawv primers,
** - tsis suav lub sijhawm siv rau kev loj hlob mycelium rau kev rho tawm DNA.
Cov pej xeem qauv
Clonal kab
Thaum tsis muaj recombination lossis nws tsis tseem ceeb los pab txhawb rau cov qauv pejxeem, cov pejxeem muaj qee tus lej ntawm cov pob zeb, kev hloov caj ces ntawm cov uas tsis tshua muaj tshwm sim.
Nyob rau hauv cov neeg zoo li no, nws yog qhov qhia ntau dua los kawm tsis yog qhov nquag ntawm cov neeg ib leeg, tab sis ntau zaus ntawm cov keeb kwm genotypes uas muaj cov keeb kwm ua ntej (cov kab mob clonal lossis clonal lineages) thiab txawv tsuas yog hauv kev hloov pauv. Cov kev tshawb fawb pej xeem ntawm lig blight pathogen thiab kev soj ntsuam ntawm cov kab clonal tau nce nrawm dua vim muaj qhov tshwm sim ntawm RFLP-RG57 tus qauv hauv 90s thaum ntxov ntawm lub xyoo pua xeem. Nrog rau RFLP-RG57, mating hom, spectra ntawm peptidase thiab piam thaj-6-phosphate isomerase isoenzymes, thiab mitochondrial DNA hom yog siv los txheeb xyuas cov kab clonal. Cov yam ntxwv ntawm cov kab clonal feem ntau tshwm sim hauv Daim Ntawv 6.
Clone US-1 muaj cov neeg nyob thoob plaws txhua qhov chaw txog rau thaum xyoo 80s, tom qab ntawd nws tau pib hloov los ntawm lwm cov clones thiab ploj ntawm Tebchaws Europe thiab North America. Tam sim no nws pom nyob rau hauv Far East (Philippines, Taiwan, Tuam Tshoj, Nyiv, Kauslim Teb, Kauslim Teb, Koh li al., 1994, Mosa et al, 1993), hauv Africa (Uganda, Kenya, Rwanda, Goodwin li al, 1994, Vega-Sanchez et al., 2000; Ochwo li al., 2002) thiab nyob rau South America (Ecuador, Brazil, Peru, Forbes et al., 1997, Goodwin li al., 1994). Tsis muaj ib hom kab raug rau US-1 kab tau muaj qhia nyob hauv Australia ib leeg. Pom tau tias, P. infestans cais tau tuaj rau tebchaws Australia nrog lwm nthwv dej ntawm lub tebchaws (Goodwin, 1997).
Clone US-6 tau tsiv teb chaws los ntawm sab qaum teb Mexico mus rau California thaum xyoo 70 thiab tau muaj kev kis mob thoob plaws hauv cov qos yaj ywm thiab txiv lws suav tom qab 32 xyoo tsis muaj mob. Vim tias nws qhov kev kub ntxhov siab, nws tau xa nyiaj rau Asmeskas-1 tus kabmob thiab pib ua tus ntug dej hiav txwv sab hnub poob ntawm Tebchaws Meskas (Goodwin et al., 1995a).
Cov genotypes US-7 thiab US-8 tau tshawb pom hauv Tebchaws Asmeskas xyoo 1992 thiab twb tau muaj rau xyoo 1994 tau muab faib dav dav hauv Tebchaws Asmeskas thiab Canada. Nyob rau ib lub caij ua liaj teb, clone US-8 muaj peev xwm yuav luag tag nrho clone US-1 hauv cov qos yaj ywm hauv thaj av pib kis tau ob qho tag nrho cov clones hauv kev sib npaug sib npaug (Miller thiab Johnson, 2000).
Clones BC-1 txog BC-4 tau txheeb xyuas nyob rau hauv British Columbia hauv qee qhov sib cais me me ntawm Goodwin li al., 1995b). Clone US-11 tau dav dav hauv Tebchaws Meskas thiab pabcuam hauv Asmeskas-1 hauv Taiwan. Clones JP-1 thiab EC-1, nrog rau clone US-1, muaj ntau yam nyob rau Nyiajpoom Teb thiab Ecuador, feem (Koh li al., 1994; Forbes et al., 1997).
SIB-1 yog tus tub luam uas kov yeej hauv tebchaws Russia hla ib cheeb tsam loj heev los ntawm thaj av Moscow mus rau Sakhalin. Hauv cheeb tsam Moscow, nws tau tshawb pom hauv xyoo 1993, thiab qee cov neeg ua liaj ua teb feem ntau yog los ntawm cov kab ntawm cov kab no, tiv taus zoo rau metalaxyl. Tom qab xyoo 1993, qhov xwm txheej ceev ntawm no clone txo qis. Sab nraum Urals xyoo 1997-1998, SIB-1 tau pom nyob txhua qhov chaw, tsuas yog Khabarovsk Thaj Chaw (cov clone SIB-2 tau dav nyob ntawd). Qhov sib thooj ntawm qhov sib txawv ntawm cov pob zeb nrog ntau hom kev sib tham tsis suav nrog cov txheej txheem kev sib deev hauv Siberia thiab Far East. Hauv cheeb tsam Moscow, hauv kev sib piv rau Siberia, cov pejxeem sawv cev los ntawm ntau lub pob zeb; yuav luag txhua txhua kem tawm nyias muaj nyias qhov cim uas tsis muaj qhov cim ntau (Elansky li al., 2001, 2015). Qhov kev sib txawv no tsis tuaj yeem piav qhia ib leeg los ntawm kev pom zoo ntawm cov kab mob fungal los ntawm ntau qhov chaw ntawm lub ntiaj teb nrog cov khoom noob txawv teb. Txij li ob hom kev sib raug zoo tshwm sim hauv cov neeg, nws yog qhov ua tau tias nws muaj ntau haiv neeg kuj yog vim xav rov ua dua. Yog li, hauv British Columbia, qhov tshwm sim ntawm cov genotypes BC-2, BC-3, thiab BC-4 tau xav tias vim yog kev sib txuas ntawm cov pob zeb BC-1 thiab US-6 (Goodwin li al., 1995b). Nws yog tau hais tias hom leeg tau pom nyob hauv nroog Moscow. Piv txwv li hom khaub thuas MO-4, MO-8 thiab MO-11 heterozygous rau PEP qhov chaw tuaj yeem yog cov menyuam nruab nrab ntawm cov hlab ntshav MO-12, MO-21, MO-22, muaj hom A2 ua ke thiab homozygous rau ib allele ntawm PEP chaw thiab hom MO-8, muaj hom A1 mating thiab homozygous rau lwm allele ntawm cov thaj chaw. Thiab yog tias qhov no yog qhov tseeb, thiab hauv cov neeg tshiab ntawm P. infestans muaj ib qho kev nyiam ntawm kev ua lub luag haujlwm ntawm kev sib deev, tom qab ntawv cov ntaub ntawv muaj nuj nqi ntawm kev tsom xam ntawm multilocus clones yuav txo qis (Elansky et al., 2001, 2015).
Kev sib txawv hauv cov kab clonal
Txog rau thaum 90s ntawm xyoo pua 20, cov kab ntawv clonal US-1 tau ua thoob plaws hauv lub ntiaj teb. Feem ntau ntawm cov teb thiab cheeb tsam cov neeg muaj feem tsis suav nrog kev sib tw nrog US-1 tus qauv. Txawm li cas los xij, kev sib txawv ntawm kev sib cais kuj raug saib xyuas, feem ntau yuav tshwm sim los ntawm ib qho kev hloov pauv. Kev sib hloov tau tshwm sim nyob rau hauv ob qho tib si nuclear thiab mitochondrial DNA thiab cuam tshuam, ntawm lwm yam, cov theem ntawm cov tshuaj tiv thaiv phenylamide thiab tus lej ntawm cov virulence noob. Cov kab uas txawv ntawm cov keeb kwm dhau los ntawm kev hloov pauv tau qhia los ntawm cov lej ntxiv tom qab teev teev tom qab lub npe ntawm cov cim keeb kwm qub (piv txwv li US-1.1 mutant kab ntawm US-1 clonal kab). Cov ntiv tes ntiv tes rau DNA kab US-1.5 thiab Asmeskas-1.6 muaj cov kab ntxiv ntawm ntau qhov sib txawv (Goodwin et al., 1995a, 1995b); cov kab xoo US-6.3 kuj txawv ntawm US-6 hauv ib kab lus txuas ntxiv (Goodwin, 1997, Rooj 7).
Hauv kev kawm txog mitochondrial DNA, nws tau pom tias tsuas yog hom 1b mitochondrial DNA tau pom nyob rau hauv txoj kab clonal US-1 (Carter et al., 1990). Txawm li cas los xij, hauv kev kawm txog cov kab ntawm cov kab mob no los ntawm Peru thiab Philippines, cais tawm tau pom tias nws cov kabmob mitochondrial DNA tau sib txawv ntawm 1b nyob rau ntawm qhov chaw ntxig thiab tshem tawm (Goodwin, 1991, Koh et al., 1994).
Rooj 6. Multilocus genotypes ntawm qee tus P. infestans clonal kab
Lub npe | Mating type | Isozymes | Cov ntiv tes DNA | MtDNA hom | |
GPI | PEP | ||||
US-1 | A1 | 86/100 | 92/100 | 1.0111010110011E + 24 | Ib |
US-2 | A1 | 86/100 | 92/100 | 1.0111010010011E + 24 | - |
US-3 | A1 | 86/100 | 92/100 | 1.0111000000011E + 24 | - |
US-4 | A1 | 100/100 | 92/92 | 1.0111010010011E + 24 | - |
US-5 | A1 | 100/100 | 92/100 | 1.0111010010011E + 24 | - |
US-6 | A1 | 100/100 | 92/100 | 1.0111110010011E + 24 | II ib |
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 | II ib |
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 |
NPE-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 |
Nco tseg: * - tsis muaj ntaub ntawv.
Rooj 7. Ntau hom genotypes thiab lawv cov kab mutant
Lub npe | Mating type | | Cov ntiv tes DNA (RG57) | Sau ntawv | |
GPI | PEB-1 | ||||
US-1 | A1 | 86/100 | 92/100 | 1011101011001101000110011 | Keeb Thawj Xeeb 1 |
US-1.1 | A1 | 86/100 | 100/100 | 1011101011001101000110011 | Kev Sib Hloov Hauv PEP |
US-1.2 | A1 | 86/100 | 92/100 | 1011101010001101000110011 | Kev Sib Hloov RG57 |
US-1.3 | A1 | 86/100 | 92/100 | 1011101001001101000110011 | Kev Sib Hloov RG57 |
US-1.4 | A1 | 86/100 | 100/100 | 1011101010001101000110011 | Kev Sib Hloov RG57 thiab PEP |
US-1.5 | A1 | 86/100 | 92/100 | 1011101011001101010110011 | Kev Sib Hloov RG57 |
US-6 | A1 | 100/100 | 92/100 | 1011111001001100010110011 | Keeb Thawj Xeeb 2 |
US-6.1 | A1 | 100/100 | 92 /92 | 1011111001001100010110011 | Kev Sib Hloov Hauv PEP |
US-6.2 | A1 | 100/100 | 92/100 | 1011101001001100010110011 | Kev Sib Hloov RG57 |
US-6.3 | A1 | 100/100 | 92/100 | 1011111001011100010110011 | Kev Sib Hloov RG57 |
US-6.4 | A1 | 100/100 | 100/100 | 1011011001001100010110011 | Kev Sib Hloov RG57 thiab PEP |
US-6.5 | A1 | 100/100 | 92/100 | 1011111001001100010010011 | Kev Sib Hloov RG57 |
BR-1 | A2 | 100/100 | 100/100 | 1011101000001100001111011 | Keeb Thawj Xeeb 3 |
BR-1.1 | A2 | 100/100 | 100/100 | 1010101000001100001110011 | Kev Sib Hloov RG57 |
Kuj tseem muaj kev hloov pauv hauv lub tsom iav ntawm isozymes. Raws li txoj cai, lawv tshwm sim los ntawm kev rhuav tshem ntawm ib tus kab mob thaum pib ua mob heterozygous rau qhov enzyme mus rau hauv homozygous. Xyoo 1993, ntawm cov txiv lws suav txiv hmab txiv ntoo, peb tau txheeb pom muaj cov yam ntxwv ntawm cov yam ntxwv ntawm US-1: RG57 ntiv tes ntiv tes, mitochondrial DNA hom, thiab 86/100 cov cim rau cov piam thaj-6-phosphati-isomerase, tab sis nws yog homozygous (100/100) rau thawj peptidase locus hloov ib 92/100 heterozygote ntse ntawm txoj kab no. Peb tau muab cov cim ntawm cov qauv no coj los ua ke MO-17 (Cov Lus 6). Txoj kab kev hloov ua ke ntawm US-1.1 thiab US-1.4 kuj txawv ntawm US-1 los ntawm kev hloov pauv ntawm thawj peptidase locus (Cov Lus 7).
Kev hloov pauv ua rau muaj kev hloov pauv ntawm cov virulence noob rau cov qos thiab lws suav hom yog qhov muaj ntau heev. Lawv tau sau tseg ntawm kev nyob ib leeg ntawm cov kab clonal US-1 hauv cov neeg tuaj ntawm Netherlands (Drenth et al., 1994), Peru (Goodwin et al., 1995a), Poland (Sujkowski li al., 1991), North North America (Goodwin et al., ., 1995b). Cov kev sib txawv ntawm cov qos yaj ywm virulence cov noob kuj tseem raug sau tseg ntawm kev sib cais ntawm cov kab mob caj dab US-7 thiab US-8 hauv Canada thiab Tebchaws Asmeskas (Goodwin et al., 1995a), ntawm cov cais tawm ntawm SIB-1 kab hauv Asian feem ntawm Russia (Elansky et al, 2001) )).
Isolates nrog cov sib txawv muaj zog hauv theem ntawm kev tawm tsam phenylamide tshuaj tau pom nyob rau hauv monoclonal teb cov neeg, txhua tus tau koom nrog txoj kab clonal Sib-1 (Elansky li al, 2001, Rooj 1). Yuav luag txhua qhov kev cuam tshuam ntawm cov kab clonal US-1 yog qhov muaj kev cuam tshuam zoo rau metalaxyl; txawm li cas los xij, muaj peev xwm tiv taus cov kab ntawm cov kab no tau raug cais nyob hauv Philippines (Koh et al., 1994) thiab hauv Ireland (Goodwin li al., 1996).
Cov neeg niaj hnub P. infestans
Central America (Mexico)
P. cov neeg nyob hauv Mexico muaj qhov sib txawv ntawm lwm cov neeg hauv ntiaj teb, uas feem ntau vim yog nws txoj haujlwm keeb kwm. Ntau cov kev tshawb fawb ntawm cov neeg no thiab cuam tshuam txog P. infestans hom tsiaj ntawm lub pob zeb Phytophthora, nrog rau hom kab ke hauv zos ntawm Solanum, ua rau qhov kev txiav txim siab tias qhov kev hloov kho ntawm cov kab mob nyob hauv nruab nrab ntawm Mexico tau tshwm sim ua ke nrog kev hloov ntawm cov tswv tsev thiab muaj kev sib deev nrog kev sib deev (Grünwald, Flier , 2005). Ob hom kev ua niam txiv yog sawv cev hauv cov pejxeem, thiab cov sib npaug sib npaug, thiab muaj oospores hauv av, ntawm cov nroj tsuag thiab cov tub ntawm cov qos yaj ywm thiab cov tsiaj qus ntsig txog Solanum tau lees tias muaj cov txheej txheem kev sib deev hauv cov pejxeem (Fernández-Pavía li al., 2002). Cov kev tshawb fawb tsis ntev los no ntawm Toluca Valley thiab nws cov chaw (thaj chaw muaj keeb kwm ntawm cov kab mob) paub tseeb hais tias muaj ntau haiv neeg ntawm cov pej xeem hauv zos ntawm P. infestans (134 multilocus genotypes hauv ib qho piv txwv ntawm 176 cov qauv) thiab muaj ntau qhov sib txawv ntawm qhov pom hauv thaj av (Wang li al., 2017). Qhov tseem ceeb ua rau muaj qhov sib txawv no yog qhov sib faib ntawm qhov sib txawv ntawm cov yam ntxwv ntawm cov toj siab hauv nruab nrab Mexico, qhov sib txawv ntawm kev cog qoob loo thiab cov qos yaj ywm siv nyob rau hauv kwj hav thiab roob, thiab muaj cov tsiaj cov tub hom Solanum uas tuaj yeem ua lwm yam kev hloov chaw (Fry et al ., 2009).
Txawm li cas los xij, nws yuav tsum raug sau tseg tias cov neeg ntawm P. infestans nyob rau sab qaum teb Mexico yog cuaj luaj thiab zoo ib yam li cov neeg Asmeskas Qaum Teb, uas qhia tau tias cov no yog cov neeg tshiab (Fry et al., 2009).
North America
Cov neeg Asmeskas North ntawm P. infestans yeej ib txwm muaj cov qauv yooj yim heev thiab lawv tus cwj pwm clonal tau tsim ntev ua ntej siv cov tsom tsom microsatellite. Mus txog xyoo 1987, cov kab tseem ceeb claus US-1 ua tus thawj hauv Tebchaws Meskas thiab Canada (Goodwin et al., 1995). Hauv lub xyoo nruab nrab xyoo 70, thaum cov kab mob ironaxyl uas muaj cov tshuaj fungicides tau tshwm sim, cov clone no tau pib hloov los ntawm lwm cov, tiv thaiv ntau cov genotypes uas tau tsiv los ntawm Mexico (Goodwin li al., 1998). Thaum xaus ntawm 90s. lub cim US-8 hloov tag nrho US-1 lub genotype hauv tebchaws Asmeskas thiab tau dhau los ua cov kab tshaj plaws ntawm cov qos yaj ywm (Fry et al., 2009; Fry et al., 2015). Qhov xwm txheej tau sib txawv nrog cov txiv lws suav, uas tas li muaj ntau cov kab clonal, thiab lawv cov kev hloov pauv ntawm xyoo mus rau xyoo (Fry et al., 2009).
Nyob rau xyoo 2009, kev muaj mob loj ntawm cov txiv lws suav liab tsoo hauv tebchaws Asmeskas. Ib qhov tshwj xeeb ntawm tus kabmob kis tau zoo tshaj plaws yog nws yuav luag thooj txhij nyob rau ntau qhov chaw hauv tebchaws Northwest, thiab nws tau ua rau muaj feem cuam tshuam nrog kev muag ntau yam ntawm cov kab mob cov txiv lws suav hauv cov vaj loj loj (Fry et al., 2013). Cov qoob loo poob muaj ntau heev. Kev tsom xam ntawm lub tshuab hlwb hlau ntawm qhov cuam tshuam tau nthuav tawm tias tus kabmob kis thoob plaws lub ntiaj teb US-22 A2-mating. Hauv xyoo 2009, cov kev faib tawm ntawm cov keeb kwm no hauv Asmeskas cov pej xeem ntawm P. infestans tau nce mus txog 80% (Fry et al., 2013). Hauv cov xyoo tom ntej, qhov kev faib ua feem ntawm kev txhoj puab heev ntawm US-23 (feem ntau yog rau cov txiv lws suav) thiab Asmeskas-24 (ntawm cov qos yaj ywm) tau nce zuj zus hauv cov pejxeem, txawm li cas los xij, tom qab 2011, kev tshawb pom ntawm US-24 poob qis, thiab txog rau hnub tim, kwv yees li 90% ntawm cov neeg muaj kabmob pathogen hauv Tebchaws Asmeskas yog sawv cev los ntawm Asmeskas-23 cov caj ces (Fry et al., 2015).
Hauv tebchaws Canada, ib yam li nyob rau tebchaws Asmeskas, thaum kawg ntawm 90s. qhov tseem ceeb tshaj plaws US-1 tau los ntawm Asmeskas-8, txoj haujlwm tseem ceeb ntawm uas tseem tsis tau hloov pauv mus txog 2008. Xyoo 2009-2010. Hauv tebchaws Canada, muaj cov kab mob tshwm sim rau cov neeg muag cov txiv lws suav, tab sis lawv raug mob los ntawm genotypes US-23 thiab US-8 (Kalischuk li al., 2012). Qhov pom tseeb ntawm thaj chaw txawv ntawm cov genotypes tau zoo kawg li: US-23 yeej nyob rau cov xeev sab hnub poob ntawm Canada (68%), thaum Asmeskas-8 tau kav ntau lub xeev nyob sab hnub tuaj (83%). Xyoo tom ntej, Asmeskas-23 kis mus rau cov cheeb tsam sab hnub tuaj; txawm li cas los xij, qhov dav dav, nws cov feem hauv cov pej xeem poob qis me ntsis tiv thaiv keeb kwm ntawm qhov tshwm sim ntawm genotypes US-22 thiab US-24 hauv lub tebchaws (Peters et al., 2014). Txog niaj hnub no, Asmeskas-23 tswj tuav txoj haujlwm tseem ceeb thoob tebchaws Canada; US-8 tam sim no nyob rau hauv British Columbia, thaum US-23 thiab US-24 tam sim no nyob hauv Ontario (Peters, 2017).
Yog li, North American cov haiv neeg ntawm P. infestans feem ntau yog cov kab clonal. Tau 40 xyoo dhau los, tus naj npawb ntawm cov pom ntawm clonal genotypes tau mus txog 24. Txawm hais tias qhov tseeb ntawm ob hom kev sib deev muaj nyob hauv cov pejxeem, qhov tshwm sim ntawm qhov tshwm sim ntawm cov genotypes tshiab yog vim qhov kev sib deev tso tawm tseem tsawg heev. Txawm li cas los xij, hauv 20 lub xyoo dhau los, ntau qhov xwm txheej ntawm qhov tshwm sim ntawm cov neeg xav tau ntawm qhov tsis txaus ntseeg tau sau tseg (Gavino et al., 2000; Danies li al., 2014; Peters li al., 2014), thiab hauv ib kis, qhov tshwm sim ntawm kev hla yog qhov keeb kwm US-11 , uas tau cog rau hauv North America tau ntau xyoo (Gavino li al., 2000). Txog rau xyoo 2009, kev hloov pauv hauv cov qauv ntawm cov pejxeem tau cuam tshuam nrog kev tshwm sim tshiab, kev tawm tsam ntau dua nrog lawv cov neeg tsiv teb tsaws chaw thiab tshem tawm ntawm cov neeg ua ntej dhau los. Dab tsi tshwm sim thaum xyoo 2009-2010 nyob hauv Asmeskas thiab Canada, epiphytotics thawj zaug qhia tau hais tias nyob rau lub caij nyoog ntawm kev hloov kho lub ntiaj teb, kev kis tus kab mob tuaj yeem cuam tshuam nrog kev sib kis ntawm cov kab mob genotypes tshiab thaum muag cov khoom cog cog.
South America
Txog thaum nyuam qhuav, cov kev tshawb fawb ntawm South American cov neeg ntawm P. infestans kuj tsis yog ib txwm muaj los sis tsis muaj neeg coob. Nws tau paub tias cov qauv ntawm cov neeg no yog qhov yooj yim heev thiab suav nrog 1-5 caj qaum ntawm ib lub tebchaws (Forbes et al., 1998). Yog li, xyoo 1998, cov genotypes US-1 (Brazil, Chile) BR-1 (Brazil, Bolivia, Uruguay, Paraguay), EC-1 (Ecuador, Colombia, Peru thiab Venezuela), AR-1, AR tau pom ntawm cov qos yaj ywm -2, AR-3, AR-4 thiab AR-5 (Argentina), PE-3 thiab PE-7 (sab qab teb Peru). Mating hom A2 yog tam sim no hauv Brazil, Bolivia thiab Argentina thiab tsis pom dhau lub Bolivian-Peruvian ciam teb nyob rau thaj tsam ntawm Pas Dej Titicaca, tom qab uas EC-1 A1 caj ces loj hauv Andes. Ntawm cov txiv lws suav, Asmeskas-1 tseem yog cov caj ces uas nrov thoob plaws South America.
Qhov xwm txheej ntau dua los sis tsawg dua qub hauv 2000s. Lub ntsiab lus tseem ceeb yog qhov kev tshawb pom nyob rau sab qaum teb Andes ntawm kev noj cov qos yaj ywm loj hlob (S. brevifolium thiab S. tetrapetalum) ntawm cov kab tshiab clonal EC-2 ntawm A2 hom (Oliva et al., 2010). Cov kev tshawb fawb phylogenetic tau qhia tias cov kab no tsis zoo ib yam rau P. infestans, txawm hais tias nws tau txheeb ze rau nws, hauv kev sib txuas nrog qhov nws tau xav kom txiav txim siab nws, nrog rau lwm txoj kab, EC-3, cais los ntawm tsob ntoo txiv lws suav S. betaceum loj hlob hauv Andes, ib hom tshiab hu ua P. andina; txawm li cas los xij, cov xwm txheej ntawm cov hom no (ib hom tsiaj ywj siab lossis kev sib xyaw ntawm P. infestans nrog qee tus tseem tsis paub kab) tseem tsis meej (Delgado li al., 2013).
Tam sim no, txhua tus neeg Asmeskas Qab Teb cov neeg ntawm P. infestans yog clonal. Txawm hais tias tseem muaj ob qho kev sib raug zoo, tsis muaj kev nrhiav tus neeg muaj peev xwm paub tau. Ntawm cov txiv lws suav, tus qauv US-1 yog qhov muaj qhov txawv, pom thaj chaw los ntawm cov qos yaj ywm los ntawm cov neeg hauv zos, keeb kwm tseeb uas tseem tsis tau paub. Hauv Brazil, Bolivia thiab Uruguay, BR-1 tus qauv tam sim no; hauv Peru, nrog rau Asmeskas-1 thiab EC-1, muaj ob peb lwm yam hauv zos cov genotypes. Hauv Andes, txoj haujlwm tseem ceeb yog tau tuav los ntawm txoj kab xoo EC-1, kev sib raug zoo ntawm cov uas muaj qhov tsis ntev los no P. andina tseem tsis tau paub. Lub tsuas "tsis khov" qhov chaw rau lub sijhawm 2003-2013. muaj cov kev hloov tseem ceeb hauv cov pej xeem, dhau los ua Chile (Acuña et al., 2012), qhov twg hauv 2004-2005. cov pathogen cov pej xeem los ua cov yeeb yam los ntawm kev tawm tsam metalaxyl thiab ib qho tshiab mitochondrial DNA haplotype (Ia hloov ntawm Ib tam sim no Ib). 2006 txog 2011 Hauv cov pejxeem, cov cim 21 (raws li SSR) tau muaj, qhov sib faib uas tau nce mus txog 90%, tom qab ntawd lub xibtes dhau mus rau genotype 20, qhov nquag ntawm qhov tshwm sim uas nyob hauv ob xyoos tom ntej tau khaws cia txog 67% (Acuña, 2015).
Teb chaws Europe
Hauv keeb kwm ntawm Tebchaws Europe, muaj tsawg kawg ob nthwv dej ntawm kev tsiv teb tsaws ntawm P. infestans los ntawm North America: nyob rau xyoo 1th. (HERB-1) thiab pib xyoo pua 70th (US-1). Qhov ua kom sib faib cov khoom ntawm metalaxyl-muaj cov fungicides hauv XNUMXs. coj mus rau kev tsiv tawm ntawm cov tawm tswv yim tseem ceeb US-XNUMX thiab nws hloov nrog cov genotypes tshiab. Vim li ntawd, nyob rau feem ntau cov tebchaws nyob sab Europe, cov neeg ntawm cov kab mob yog sawv cev tsuas yog los ntawm ntau cov kab clonal.
Kev siv cov microsatellite tsom rau kev soj ntsuam ntawm cov neeg muaj peev xwm ua rau nws muaj peev xwm txheeb xyuas qhov kev hloov pauv hnyav uas tau tshwm sim nyob rau Western Europe xyoo 2005-2008. Xyoo 2005, kab tshiab tau tshawb pom nyob rau hauv tebchaws Askiv, hu ua 13_A2 (lossis "Xiav 13") thiab ua cim los ntawm hom A2 mating , siab kev tawm tsam thiab kev tawm tsam rau phenylamides (Shaw et al., 2007). Daim ntawv qhia tib hom tau pom nyob hauv cov qauv sau hauv 2004 hauv Netherlands thiab sab qaum teb Fabkis, tawm tswv yim tias nws tau tsiv mus rau Tebchaws Askiv los ntawm Tebchaws Europe txuas ntxiv, tej zaum muaj noob qos yaj ywm (Cooke li al., 2007). Txoj kev tshawb ntawm cov genome ntawm cov neeg sawv cev ntawm cov kab mob no tau qhia txog qib siab ntawm polymorphism ntawm nws cov theem (los ntawm 2016, tus lej ntawm nws cov subclonal hloov tau mus txog 340) thiab qhov tseem ceeb ntawm kev hloov pauv hauv theem ntawm cov noob qhia, incl. effector noob thaum raug mob ntawm cov nroj tsuag (Cooke et al., 2012; Cooke, 2017). Cov yam ntxwv no, nrog rau kev ncua sijhawm ntev ntawm cov xyoob ntoo biotrophic, tuaj yeem ua rau muaj zog ntau ntxiv ntawm 13_A2 thiab nws lub peev xwm kis tau txawm tias qos hom tiv taus kab lig.
Hauv ob peb xyoos tom ntej no, lub caj dab tau nthuav dav mus thoob plaws cov teb chaws Northwestern Europe (Great Britain, Ireland, Fabkis, Belgium, Netherlands, Lub Tebchaws Yelemees) nrog kev hloov chaw nyob ib puag ncig dhau los 1_A1, 2_A1, 8_A1 (Montarry et al. 2010; Gisi li al. , 2011; Van den Bosch et al., 2011; Ncuav qab zib, 2015; Khaub noom, 2017). Raws li lub vev xaib www.euroblight.net, qhov sib koom ntawm 13_A2 nyob rau hauv cov neeg ntawm cov tebchaws no nce mus txog 60-80% thiab ntau dua; lub xub ntiag ntawm cov cim hluav taws xob no tseem tau sau tseg hauv qee lub tebchaws ntawm Eastern thiab yav qab teb Europe. Txawm li cas los xij, xyoo 2009-2012. 13_A2 poob nws txoj haujlwm tseem ceeb hauv Great Britain thiab Fabkis, yielding mus rau 6_A1 kab (8_A1 hauv Ireland), thiab nyob rau hauv Netherlands thiab Belgium nws tau hloov kho qee yam los ntawm genotypes 1_A1, 6_A1, thiab 33_A2 (Cooke et al., 2012; Cooke, 2017; Stellingwerf, 2017).
Txog rau hnub tim, kwv yees li 70% ntawm Western European cov pej xeem ntawm P. infestans yog monoclonal. Raws li lub vev xaib vev xaib www.euroblight.net, hom tshuaj tseem ceeb nyob hauv cov tebchaws ntawm North-Western Europe (UK, Fabkis,
Netherlands, Belgium) nyob twj ywm, kwv yees li ntawm cov sib npaug ntawm cov sib npaug, 13_A2 thiab 6_A1, thiab tom kawg ua tsis tshwm sim sab nraum thaj chaw tshwj xeeb (tshwj tsis yog Tebchaws Ireland), tab sis twb muaj tsawg kawg 58 subclones (Cooke, 2017). Kev hloov pauv 13_A2 nyob rau hauv tus naj npawb pom nyob hauv lub teb chaws Yelemees, thiab tseem pom nyob rau hauv cov tebchaws Central thiab yav qab teb thaj tsam. Cim 1_A1 ua ib feem tseem ceeb ntawm cov neeg ntawm Belgium thiab ib nrab yog Netherlands thiab Fabkis. Zaj Cim 8_A1 tau ruaj khov nyob rau hauv cov pej xeem European nyob rau theem ntawm 3-6%, tshwj tsis yog Ireland, qhov chaw uas nws tau tuav nws txoj haujlwm ua thawj coj thiab tau faib ua ob subclones (Stellingwerf, 2017). Thaum kawg, xyoo 2016, muaj ib qho kev hloov pauv ntxiv ntawm qhov tshwm sim ntawm cov genotypes tshiab 36_A2 thiab 37_A2, sau thawj zaug xyoo 2013-2014; txog rau hnub tim, cov kab mob genotypes muaj nyob hauv Netherlands thiab Belgium thiab ib nrab nyob rau Fabkis thiab Lub Tebchaws Yelemees, ib yam li nyob rau yav qab teb Tebchaws Askiv (Cooke, 2017). Kwv yees li 20-30% ntawm Western European cov pej xeem yog sawv cev los ntawm cov cim genotypes txhua xyoo.
Tsis zoo li Western Europe, los ntawm lub sijhawm 13_A2 tus qauv tshwm sim, cov neeg ntawm Qaum Teb Europe (Sweden, Norway, Denmark, Finland) tau sawv cev tsis yog los ntawm kab kab, tab sis los ntawm ntau cov cim ntawm genotypes (Brurberg li al.,
Xyoo 2011). Nyob rau lub sijhawm muaj kev sib kis nquag ntawm 13_A2 hauv Western Europe, qhov pom ntawm cov cim ntawm no hauv Scandinavia tsis tau pom txog thaum 2011, thaum nws tau pom thawj zaug hauv North Jutland (Denmark), qhov twg feem ntau muaj cov qos yaj ywm loj hlob nrog kev siv nquag ntawm metalaxyl-muaj cov kab mob fungicides (Nielsen li al., 2014). Raws li www.euroblight.net, keeb kwm 13_A2 kuj tseem raug kuaj hauv ntau qhov qauv los ntawm Norway thiab Denmark hauv 2014 thiab hauv ntau cov piv txwv Norwegian hauv xyoo 2016; ntxiv rau, hauv 2013, lub sijhawm ntawm cov cim 6_A1 hauv ib qho nyiaj tsawg tau muab sau tseg hauv Finland. Qhov laj thawj tseem ceeb rau qhov ua tsis tiav ntawm 13_A2 thiab lwm cov kab hauv pob zeb hauv kev sib tw ntawm Scandinavia raug suav tias yog huab cua huab cua sib txawv ntawm thaj av no los ntawm cov teb chaws nyob rau Western Europe.
Ntxiv nrog rau qhov tseeb uas txias lub caij ntuj sov thiab lub caij ntuj no txias ua rau muaj kev ciaj sia ntawm oospores ntau dua li kev noj qab haus huv mycelium (Sjöholm et al., 2013), av cia rau huab cua hauv lub caij ntuj no (uas feem ntau tsis tshwm sim hauv cov teb chaws sov ntawm Western Europe) ua rau muaj kev sib txuam ntawm oospores kev cog qoob loo thiab cog. qos, uas txhim kho lawv txoj haujlwm ua lub hauv paus ntawm tus kab mob thawj zaug (Brurberg li al., 2011). Nws tseem yuav tsum tau sau tseg tias, nyob rau sab qaum teb kev mob, kev tsim kab mob kis los ntawm oospores nthuav dav txoj kev loj hlob ntawm cov tub raj muaj kab mob, uas thaum kawg txwv tsis pub muaj tus thawj ntawm txawm ntau dua kev sib cav, tab sis tom qab tsim cov kab mob clonal (Yuen, 2012). Tus qauv ntawm cov neeg feem coob kawm ntawm P. infestans hauv cov teb chaws ntawm Eastern Europe (Tebchaws Poland, Tebchaws Baltic) tau zoo sib xws rau hauv tebchaws Scandinavia.
Ob hom kev xaiv tus poj niam tseem muaj nyob ntawm no, thiab feem ntau ntawm cov genotypes txiav txim siab los ntawm SSR tsom xam yog qhov tshwj xeeb (Chmielarz et al., 2014; Runno-Paurson li al., 2016). Raws li nyob rau sab qaum teb Europe, kev faib tawm ntawm cov kab clonal (feem ntau ntawm 13_A2 caj ces) tsis cuam tshuam rau cov pej xeem hauv zos ntawm cov kab mob, uas khaws cov theem siab ntawm ntau haiv neeg nrog qhov tsis muaj cov lus tshaj tawm txoj kab.
Qhov pom ntawm 13_A2 qee zaus pom nyob rau hauv cov teb muaj ntau hom qos. Hauv Lavxias, qhov xwm txheej tau tsim los ntawm qhov zoo sib xws. Kev tsom xam ntawm P. infestans cais nyob rau xyoo 2008-2011 hauv 10 thaj chaw sib txawv ntawm European feem ntawm Lavxias, qhia txog qib siab ntawm kev sib txawv genotypic thiab kev ua tiav tsis muaj qhov tsis txaus ntseeg nrog European cov kab xaum (Statsyuk li al., 2014). Ntau xyoo tom qab, kev tshawb fawb ntawm P. infestans cov qauv sau nyob hauv thaj av Leningrad xyoo 2013-2014 qhia qhov sib txawv ntawm lawv thiab cov genotypes los ntawm thaj av no tau txheeb pom hauv cov kev tshawb fawb dhau los. Hauv ob txoj kev tshawb nrhiav, tsis pom sab hnub poob European European genotypes (Beketova li al., 2014; Kuznetsova li al., 2016).
Cov caj ces muaj ntau haiv neeg ntawm cov pej xeem nyob sab hnub tuaj ntawm P. infestans thiab qhov uas tsis muaj cov kab clonal nyob rau hauv lawv yuav cuam tshuam nrog ntau yam. Ua ntej, ib yam li nyob rau Qaum Teb Europe, cov huab cua kev nyab xeeb ntawm cov teb chaws suav hais tias yog ua los ntawm kev tsim oospores yog qhov tseem ceeb ntawm kev kis tus kab mob (Ulanova li al., 2010; Chmielarz li al., 2014). Thib ob, ib qho tseem ceeb ntawm cov qos yaj ywm tsim hauv cov teb chaws no tau loj hlob ntawm cov liaj teb ntiav me, feem ntau nyob ib puag ncig los ntawm hav zoov lossis lwm yam kev cuam tshuam rau qhov kev txav dawb ntawm cov khoom sib kis (Chmielarz li al., 2014). Raws li txoj cai, qos yaj ywm zus nyob rau hauv cov xwm txheej zoo li tsis tau kho nrog tshuaj, thiab kev xaiv ntawm ntau yam yog ua raws li lawv cov blight tsis kam, i.e. tsis muaj kev xaiv ua kom nruj rau kev txhoj puab heev thiab tiv taus hlau hlau, uas ua rau tsis haum cov tshuaj tiv thaiv genotypes, xws li 13_A2, ntawm qhov zoo dua lwm cov genotypes (Chmielarz li al., 2014). Thaum kawg, vim thaj av me me, lawv cov tswv feem ntau tsis xyaum cog qoob loo, cog qos yaj ywm rau xyoo nyob rau hauv tib qho chaw, uas ua rau muaj kev txuam nrog kev tsim hluav taws xob tsis sib luag (Runno-Paurson et al., 2016; Elansky, 2015; Elansky et al. ., 2015).
Asia
Txog thaum tsis ntev los no, tus qauv ntawm P. infestans cov neeg nyob hauv Asia tseem tsis tshua nkag siab. Nws tau paub tias nws yog sawv cev tsuas yog los ntawm kab ntau dhau, thiab cov txiaj ntsig ntawm kev sib deev rov qab los ntawm kev tshwm sim ntawm cov genotypes tshiab yog tsawg heev. Yog li, piv txwv li, hauv xyoo 1997-1998. Nyob rau cov tebchaws Esxias ntawm tebchaws Russia (Siberia thiab Far East), cov neeg muaj kabmob pathogen tau sawv cev los ntawm tsuas yog peb tus genotypes uas muaj qhov tshwm sim dhau los ntawm SIB-1 genotype (Elansky et al., 2001). Muaj cov clonal pathogen kab tau pom nyob hauv cov teb chaws xws li Tuam Tshoj, Nyiv, Kaus Lim Kauslim, Philippines, thiab Taiwan (Koh li al., 1994; Chen li al., 2009). Txoj kab xaim US-1 tau hla dhau ib thaj chaw loj ntawm Asia nyob rau xyoo 90s - thaum ntxov 2000s. yuav luag txhua qhov chaw pib hloov los ntawm lwm cov genotypes, uas, nyeg, muab txoj kev rau tus tshiab. Feem ntau, kev hloov pauv ntawm cov qauv thiab kev txuam ntawm cov neeg nyob hauv cov tebchaws Asian tau txuam nrog kev tsiv teb tsaws ntawm cov genotypes tshiab los ntawm sab nraud. Yog li, hauv Nyij Pooj, tshwj xeeb ntawm JP-3 caj ces, tag nrho lwm cov genotypes Nyij Pooj uas tshwm sim tom qab Asmeskas-1 (JP-1, JP-2, JP-3) muaj ntau dua los yog tsawg cov pov thawj sab nraud (Akino et al., 2011) ... Tam sim no muaj peb lub ntsiab ntawm cov neeg tseem ceeb nyob rau hauv Suav teb, uas muaj thaj chaw thaj chaw meej; Tsis muaj ib qho muaj lossis noob muaj zog txaus los ntawm cov neeg no (Guo et al., 2010; Li li al., 2013b). Cim 13_A2 tshwm sim rau thaj chaw ntawm Tuam Tshoj hauv nws cov xeev yav qab teb (Yunnan thiab Sichuan) xyoo 2005-2007, thiab xyoo 2012-1014. tau pom hauv sab qaum teb sab hnub tuaj ntawm lub teb chaws (Li li al., 2013b). Hauv Is Nrias teb, 13_A2 tshwm sim txawm tias nyob rau tib lub sijhawm zoo li nyob rau hauv Suav teb, feem ntau yuav nrog cov noob qos yaj ywm (Chowdappa et al., 2015), thiab xyoo 2009-2010. ua rau muaj mob loj heev epiphytotic txog lig blight ntawm cov txiv lws suav nyob rau sab qab teb ntawm lub teb chaws, tom qab ntawd nws tau kis rau cov qos yaj ywm thiab xyoo 2014 tau ua rau muaj kev npau taws rau cov neeg sab hnub poob hauv West Bengal, uas ua rau muaj kev puas tsuaj thiab kev tua tus kheej ntawm ntau tus neeg ua liaj ua teb hauv zos (Fry, 2016).
Teb chaws Africa
Txog xyoo 2008-2010 kev kawm paub tshawb fawb txog tus kab P. infestans hauv tebchaws Africa tsis tau coj los ua. Lub sijhawm no, cov neeg African ntawm P. infestans tuaj yeem muab faib ua ob pawg, thiab qhov kev faib tawm no tau pom meej meej nrog qhov tseeb ntawm qhov tseem ceeb ntawm kev coj cov noob qos yaj ywm los ntawm Tebchaws Europe.
Hauv North Africa, uas nquag nqa cov noob qos yaj ywm los ntawm Tebchaws Europe, hom A2 mating yog dav sawv cev nyob rau hauv yuav luag txhua thaj chaw, uas muab qhov kev xav ntawm qhov tshwm sim ntawm cov genotypes tshiab uas yog los ntawm kev sib daj sib deev (Corbière et al., 2010; Rekad et al., 2017). Tsis tas li ntawd, hauv Algeria, muaj cov genotypes 13_A2, 2_A1, thiab 23_A1 tau sau tseg nrog kev tshaj tawm ntawm cov thawj ntawm lawv, nrog rau kev maj mam txo qis hauv cov kev sib faib ntawm cov cim genotypes kom tiav kev ploj (Rekad et al., 2017). Hauv kev sib piv rau thaj tsam ntawm thaj av, hauv Tunisia (tshwj tsis yog lub nroog qaum teb sab qaum teb), cov neeg muaj kabmob loj yog sawv cev los ntawm A1 mating ntau yam (Harbaoui li al., 2014).
Clonal kab NA-01 yog hom ntawm no. Nyob rau hauv dav dav, qhov kev faib ua feem ntawm kab clonal nyob rau hauv cov pej xeem tsuas yog 43%. Nyob rau sab hnub tuaj thiab yav qab teb Africa, qhov twg ntim ntawm cov noob ntshuam yog me me xwb (Fry li al., 2009), P. infestans yog sawv cev los ntawm tsuas yog ob qho clonal A1-hom kab, Asmeskas-1 thiab KE-1, thiab tom kawg ua rau hloov chaw qub ntawm cov qos yaj ywm ( Pule et al., 2012; Njoroge li al., 2016). Txog rau hnub tim, ob qho tib si ntawm cov genotypes muaj ib qho tshwm sim muaj pes tsawg ntawm subclonal hloov.
Australia
Thawj qhov kev tshaj tawm xov xwm txog blight qos yaj ywm hauv Australia hnub tim rov qab rau xyoo 1907, thiab thawj zaug epiphytotia, txawm tias yog los nag hlob heev hauv lub hli sov, tau tshwm sim xyoo 1909-1911. (Drenth li al., 2002). Nyob rau hauv dav dav, txawm li cas los, lig blight tsis muaj qhov tseem ceeb nyiaj txiag tseem ceeb rau lub teb chaws. Tshaj tawm ntawm cov blight lig, ua rau cov huab cua ua kom muaj huab cua sov, tshwm sim tsis ntau tshaj ib zaug txhua 5-7 xyoo thiab tau pib ua thaj chaw feem ntau nyob rau sab qaum teb Tasmania thiab nruab nrab Victoria. Nyob rau hauv kev sib txuas nrog cov saum no, cov ntawv tshaj tawm tau kawm rau txoj kev tshawb nrhiav ntawm cov qauv ntawm Australian cov pej xeem ntawm P. infestans yog xyaum tsis tuaj. Cov ntaub ntawv muaj tseeb zaum kawg los ntawm 1998-2000. (Drenth li al., 2002). Raws li cov sau phau ntawv, cov pej xeem Victoria yog ib tus kab clonal US-1.3, uas tau lees paub tias tsis muaj tseeb txog kev tsiv teb tsaws chaw ntawm cov qauv no los ntawm Tebchaws Meskas. Cov piv txwv Tasmanian tau muab cais ua AU-3, uas txawv ntawm cov genotypes uas tau muaj nyob rau lub sijhawm ntawd hauv lwm qhov chaw ntawm lub ntiaj teb.
Cov yam ntxwv ntawm kev txhim kho ntawm blight lig nyob rau hauv Russia
Hauv Tebchaws Europe, kev kis tus kabmob nrog cov noob kab mob, oospores uas dhau mus ua av, zoo li zoosporangia coj los ntawm cua los ntawm cov nroj tsuag cog los ntawm overwintered cov tub luam nyob rau xyoo tas los cov teb ("cog" paj ntoo), lossis ntawm cov kem culled phau ntawv rau cia ntawm tubers. Ntawm no, cov nroj tsuag zus nyob rau heaps ntawm pov tseg tubers raug suav hais tias yog feem ntau txaus ntshai qhov chaw ntawm tus kab mob. muaj, tus naj npawb ntawm cov qe tawm yog feem ntau tseem ceeb, thiab zoosporangia tuaj yeem nqa los ntawm lawv dhau mus deb. Qhov seem ntawm cov chaw (oospores, "tuaj pab dawb" nroj tsuag) tsis txaus ntshai, vim tias nws tsis yog kev coj los cog ntawm cov ntoo hauv thaj chaw qub ntau zaus ntau dua ib zaug txhua txhua 3-4 xyoo. Kev kis tus kab mob los ntawm cov noob ntxaum kuj tseem tsawg vim yog lub noob zoo tswj tau.
Nyob rau hauv dav dav, tus nqi ntawm inoculum nyob rau hauv European pejxeem yog tsawg, thiab yog li ntawd nce nyob rau hauv lub phaum mob yog qeeb qeeb thiab muaj peev xwm yuav ntse tswj siv tshuaj fungicidal npaj. Lub luag haujlwm tseem ceeb hauv cov xwm txheej European yog kev tawm tsam rau kev sib kis kab mob nyob rau theem thaum qhov zaws ntawm zoosporangia los ntawm cov nroj tsuag cuam tshuam tau pib.
Hauv Lavxias, qhov xwm txheej dhau los sib txawv. Feem ntau ntawm cov qos thiab lws suav qoob loo yog cog hauv cov tsev ntiav me; kev tiv thaiv kev ntsuas yog ob qho tib si tsis ua rau lawv txhua, lossis kev kho kom haum fungicidal tau nqa tawm hauv tus naj npawb tsis txaus thiab pib tom qab qhov tsos pom ntawm blight lig rau saum. Yog li ntawd, cov tsev cog qoob loo ntiag tug ua lub hauv paus uas kis tus kab mob, los ntawm cov zoosporangia uas nqa los ntawm cov cua mus rau cov chaw ua lag luam. Qhov no tau lees paub los ntawm peb cov kev soj ntsuam ncaj qha hauv Moscow, Bryansk, Kostroma, Ryazan thaj chaw: kev puas tsuaj rau cov nroj tsuag hauv cov vaj tsev ntiav yog pom txawm tias ua ntej pib fungicide kho ntawm kev ua lag luam plantings. Tom qab ntawd, qhov kis mus rau thaj teb loj yog raug khi los ntawm kev siv cov tshuaj fungicidal, thaum nyob hauv cov vaj tsev ntiav muaj kev txhim kho sai ntawm blight lig.
Nyob rau hauv cov ntaub ntawv tsis raug lossis "peev nyiaj" kev kho mob ntawm cov ntoo cog kev lag luam, foci ntawm lig blight kuj tshwm sim hauv thaj teb; tom qab ntawd lawv nquag tsim kho, npog thaj tsam loj dua (Elansky, 2015). Kev kis rau hauv cov tsev ntiav muaj qhov cuam tshuam loj ntawm kev sib kis hauv thaj chaw ua lag luam. Hauv txhua lub qos-cog thaj tsam ntawm Lavxias, thaj chaw nyob los ntawm cov qos yaj ywm hauv tsev ntiav muaj ntau zaus ntau dua li thaj tsam tag nrho ntawm cov liaj teb ntawm cov neeg tsim khoom loj. Nyob rau hauv xws li ib puag ncig, ntiav cog vaj yuav saib tau raws li lub ntiaj teb inoculum kev pab rau cov teb coj mus muag. Cia peb sim txheeb xyuas cov khoom uas yog cov xeeb ceem ntawm cov genotypes ntawm lim nyob rau hauv ntiag tug lub vaj.
Cog cov tsis muaj noob thiab kev cais tawm ntawm cov kab mob cov qos yaj ywm, cov noob txiv lws suav tau txais los ntawm cov tsim txom txawv teb chaws, kev cog qoob loo ntev thiab qos yaj ywm rau ntawm tib thaj chaw, kev kho cov tshuaj tua kab tsis raug lossis lawv tsis tiav ua rau muaj mob loj heev nyob rau hauv kev lag luam ntiag tug, qhov tshwm sim ntawm yog dawb hla kev, hybridization thiab tsim ntawm oospores nyob rau hauv ntiav vaj. Raws li qhov tshwm sim, muaj ntau qhov sib txawv genotypic ntawm cov kab mob raug soj ntsuam, thaum yuav luag txhua qhov txawv txawv hauv nws cov cim (Elansky li al., 2001, 2015). Cog cov noob qos yaj ywm ntawm ntau cov caj ces ua rau nws tsis zoo li cov kab uas tshwj xeeb rau cov kab tshwj xeeb yuav tawm tuaj. Cov hom raug xaiv nyob rau hauv xws li cov kis no txawv los ntawm lawv cov versatility hauv kev sib raug zoo nrog cov cuam tshuam ntau yam, feem ntau ntawm lawv muaj qhov ze rau cov siab ntawm cov virulence noob. Qhov no yog qhov sib txawv ntawm cov kab ke ntawm cov "clonal kab" feem ntau rau cov teb loj ntawm cov chaw ua liaj ua teb nrog cov txheej txheem tsim kom muaj kev tiv thaiv tiv thaiv blight lig. "Clonal kab" (thaum txhua hom kab mob lig blight pathogen hauv lub tshav pob yog sawv cev los ntawm ib lossis ntau qhov genotypes) yog qhov chaw nyob hauv cov teb chaws uas cog qoob loo yog nqa tshwj xeeb los ntawm cov liaj teb loj: Tebchaws USA, Tebchaws Netherlands, Denmark, thiab lwm yam Hauv Tebchaws Askiv, Ireland, Poland, qhov twg hauv tsev neeg thaj av ib txwm tseem coj raws li ib txwm muaj. qos yaj ywm loj hlob, muaj kuj tseem muaj ntau dua genotypic ntau haiv neeg hauv vaj. Thaum xaus ntawm lub xyoo pua 20, "cov kab txuas pob txha" tau nrov thoob plaws Asia thiab Far Eastern ntawm Russia (Elansky et al., 2001), uas pom meej meej vim siv tib hom qos yaj ywm tshwj xeeb rau kev cog ntoo. Tsis ntev los no, qhov xwm txheej hauv cov cheeb tsam no tseem pib hloov pauv ntawm kev nce hauv genotypic ntau haiv neeg.
Cov tsis muaj kev kho mob hnyav nrog fungicidal npaj muaj lwm, qhov tsim nyog tau ncaj qha - tsis muaj qhov txuam nrog ntawm cov tawv ntoo tsis haum hauv lub vaj. Tseeb, peb cov txiaj ntsig pom tau tias cov hlau-hlau tsis tshua pom kev pom muaj tsawg dua nyob rau hauv cov chaw ntiav tshwj xeeb tshaj li hauv kev cog ntoo.
Qhov sib thooj ntawm cov qos yaj ywm thiab cov txiv lws suav cog, ib txwm muaj rau cov tsev ntiav, ua kom yooj yim rau kev tsiv ntawm cov kab mob nruab nrab ntawm cov qoob loo no, vim qhov ntawd, ntawm kaum xyoo dhau los, ntawm txoj kev sib cais los ntawm cov qos yaj ywm, qhov sib npaug ntawm txoj kev nqa cov noob rau kev tawm tsam rau ntau hom txiv lws suav (T1), yav dhau los yam ntxwv tsuas yog rau " txiv lws suav "lim ntshav. Txoj kev mob ua ke nrog T1 cov noob feem ntau mob siab rau ntawm qos yaj ywm thiab txiv lws suav.
Nyob rau hauv xyoo tsis ntev los, lig blight on lws suav pib tshwm sim nyob rau hauv ntau kis ntxov tshaj rau qos yaj ywm. Cov txiv ntseej muaj peev xwm kis tau los ntawm oospores hauv av, lossis oospores tam sim no hauv cov noob txiv lws suav lossis ua raws li lawv (Rubin et al., 2001). Hauv 15 lub xyoo dhau los, coob leej pheej yug cov noob ntoo pheej yig, feem ntau yog tuaj txawv teb chaws, tau tshwm sim hauv khw muag khoom, thiab feem ntau ntawm cov neeg tsim khoom me tau hloov mus siv lawv. Cov noob tuaj yeem nqa cov hlab ntshav nrog cov genotypes ib txwm muaj ntawm thaj tsam ntawm lawv qhov kev loj hlob. Yav tom ntej, cov genotypes no suav nrog cov txheej txheem kev sib deev hauv tsev ntiav, uas ua rau muaj kev rov tshwm sim ntawm genotypes tshiab kiag li.
Yog li, nws tuaj yeem hais tias cov tsev ntiav yog lub ntiaj teb "lub lauj kaub rau" hauv lub ntiaj teb, uas yog los ntawm kev sib pauv ntawm cov khoom hauv caj ces, cov genotypes uas twb muaj lawm tau ua tiav thiab tag nrho cov tshiab tshwm sim. Ntxiv mus, lawv xaiv tshwm sim nyob rau hauv tej yam kev mob uas txawv heev ntawm cov uas tsim rau cov qos yaj ywm hauv cov liaj teb loj: tsis muaj cov xov ntawm fungicidal, kev tsis sib xws ntawm cov nroj tsuag, qhov tseem ceeb ntawm cov nroj tsuag cuam tshuam los ntawm ntau hom kab mob thiab kab mob sib kis, sib thooj rau cov txiv lws suav thiab cov noog tsaus ntuj, nquag hla kev thiab oospore tsim, tau rau oospores ua kab ke uas yog qhov chaw kis rau lwm xyoo.
Tag nrho cov no ua rau muaj ntau qhov ntau heev genotypic ntau haiv ntawm tiaj nraum qaum tsev neeg. Nyob rau hauv cov xwm txheej ntawm epiphytotics hauv zaub vaj, lig blight kis tau sai heev thiab cov nyiaj ntawm cov noob kab mob tso tawm, ya mus rau thaj chaw ua lag luam nyob ze. Txawm li cas los xij, tau nkag mus rau thaj chaw ua lag luam nrog cov txheej txheem tseeb ntawm kev ua liaj ua teb thev naus laus zis thiab kev tiv thaiv tshuaj, cov noob kab tsuag uas tau tuaj txog tau tsis muaj sijhawm los pib mob epiphytotics hauv thaj teb, uas yog vim tsis muaj cov kab txuas los tiv thaiv cov kab mob fungicides thiab tshwj xeeb rau cov khoom siv ntau hom.
Lwm qhov chaw ntawm thawj inoculum tej zaum yuav kis kabmob kaw rau hauv kev lag luam seedlings. Cov tub no tau loj hlob, raws li txoj cai, hauv cov teb uas muaj cov thev naus laus zis zoo thiab muaj cov tshuaj tiv thaiv zoo. Cov genotypes ntawm cov cais tawm uas kis tau lub raj tau yoog rau kev txhim kho ntawm lawv tus kheej ntau yam. Cov hom kab no muaj kev phom sij rau kev lag luam cog ntau dua li cov hauv paus hauv vaj hauv tsev. Cov txiaj ntsig ntawm peb qhov kev tshawb fawb kuj txhawb qhov kev xav no. Cov pej xeem cais tawm los ntawm cov liaj teb loj nrog kev soj ntsuam tshuaj kom zoo thiab kev siv tshuab ua liaj ua teb zoo tsis txawv hauv qhov muaj ntau hom genotypic Feem ntau cov no yog ob peb qho kab clonal uas nquag txhoj puab heev.
Cov kab mob los ntawm cov khoom lag luam hauv cov noob tuaj yeem nkag mus rau cov neeg nyob hauv cov vaj zaub thiab tau koom nrog cov txheej txheem uas yuav tshwm sim hauv lawv. Txawm li cas los xij, nyob hauv ib lub vaj zaub, lawv cov kev sib tw yuav qis dua li hauv thaj chaw ua lag luam, thiab tsis ntev lawv yuav tsum tsis muaj nyob rau hauv txoj hauv kev ntawm kab, tab sis lawv cov noob yuav siv tau hauv "vaj" pejxeem.
Tus kab mob kis tau ntawm "cog qoob loo" cov nroj tsuag thiab ntawm cov txhab ntawm culled thaum lub sijhawm sau qoob tsis muaj feem xyuam rau Russia, vim tias Nyob rau hauv lub ntsiab qos-loj hlob thaj chaw ntawm Russia, sib sib zog nqus lub caij ntuj no av khov yog pom, thiab cov nroj tsuag los ntawm cov tub ntxhais uas muaj wintered hauv cov av tsis tshua muaj kev loj hlob. Ntxiv mus, raws li peb cov kev sim qhia, lig blight pathogen tsis muaj sia nyob ntawm qhov tsis kub tsis zoo txawm tias ntawm cov tub ntxhais uas tau tuav lawv txoj kev muaj peev xwm. Nyob rau hauv thaj chaw qhuav, qhov chaw uas cog qoob loo rau thaum ntxov, xyaum blight lig yog qhov tsis tshua muaj vim lub caij qhuav thiab kub pheej loj.
Yog li, peb tab tom soj ntsuam kev faib P. cov neeg tsawg mus rau hauv “thaj chaw” thiab “vaj” coob. Txawm li cas los xij, nyob rau hauv xyoo tas los no, cov txheej txheem tau muaj kev ceeb toom ua rau muaj kev sib txuam thiab txhais lus ntawm cov genotypes los ntawm cov neeg no.
Ntawm lawv, ib qho tuaj yeem sau tseg qhov nce ntawm kev paub ntawm cov neeg tsim khoom me me, tshwm sim ntawm cov pob khoom me me ntawm cov qos yaj ywm noob, kis ntawm fungicidal npaj hauv cov pob me me, thiab cov tsis ntshai ntawm "kev ua tshuaj" los ntawm cov pej xeem.
Qhov xwm txheej tshwm sim thaum, ua tsaug rau kev ua ub ua no ntawm ib tus neeg xa khoom, tag nrho cov zos tau cog nrog cov noob ntawm tib cov ntau yam thiab muab nrog cov pob me me ntawm tib cov tshuaj tua kab. Nws tuaj yeem xav tias cov qos yaj ywm ntawm tib yam ntau yam yuav pom ntawm cov chaw cog ntoo ua lag luam nyob ze.
Ntawm qhov tod tes, qee cov tuam txhab kev lag luam tshuaj tua tsiaj tau txhim kho cov phiaj xwm kev siv tshuaj "kev siv nyiaj". Hauv qhov no, tus naj npawb ntawm cov kev kho mob pom zoo yog qhov tsis txaus ntseeg thiab qhov pheej yig tshaj fungicides muaj, thiab qhov tseem ceeb tsis yog los ntawm kev tiv thaiv kev txhim kho ntawm blight lig txog rau txiav cov plaub hau, tab sis ntawm qee qhov kev ncua hauv epiphytoty thiaj li yuav nce cov txiaj ntsig. Xws li cov qauv kev lag luam yog kev ncaj ncees thaum cog cov qos yaj ywm los ntawm cov khoom siv noob qis-qib, thaum nyob rau hauv hauv paus ntsiab lus tsis muaj lus nug ntawm tau txais txiaj ntsig zoo. Txawm li cas los xij, hauv qhov no, hauv kev sib piv rau cov neeg vaj, cov qib keeb kwm keeb kwm ntawm cov qos tau pab xaiv cov tshwj xeeb ntawm kev sib tw physiological, uas yog qhov txaus ntshai heev rau ntau yam no.
Feem ntau, cov nyiam rau txoj kev sib hloov ntawm "vaj" thiab "teb" txoj kev ntawm cov qos yaj ywm tsim tawm zoo li peb txaus ntshai. Txhawm rau tiv thaiv lawv cov txiaj ntsig tsis zoo, ob leeg hauv tsev thiab kev lag luam, nws yuav tsum tau tswj hwm ob qho kev lees paub ntawm cov noob qos yaj ywm thiab cov kab mob ntawm cov fungicides pub rau cov tswv ntiag tug hauv cov ntim me, nrog rau kev soj qab taug qab cov phiaj xwm kev tiv thaiv thiab kev siv cov fungicidal npaj hauv kev lag luam.
Hauv thaj chaw ntawm kev lag luam ntiag tug, muaj kev tsim kho ntawm tsis tsuas lig blight, tab sis kuj Alternaria. Feem ntau cov tswv ntawm cov liaj teb ntiag tug tsis siv tshwj xeeb los tiv thaiv Alternaria, yuam kev kev loj hlob ntawm Alternaria rau lub ntuj wilting ntawm cov nplooj ntoo lossis kev txhim kho ntawm blight lig. Yog li ntawd, nrog kev txhim kho ntau ntawm Alternaria ntawm cov tau ntxim nyiam, cov phiaj hauv tsev neeg tuaj yeem ua lub hauv paus ntawm inoculum rau kev cog ntoo.
Cov txheej txheem ntawm cov hloov pauv
Txoj kev hloov kom muaj txheej txheem
Txij li thaum qhov tshwm sim ntawm kev hloov pauv yog ib qho txheej txheem sib txawv mus nrog cov tsawg zaus, qhov tshwm sim ntawm kev hloov pauv ntawm txhua qhov chaw nyob yog nyob ntawm ntau zaus ntawm kev hloov pauv ntawm cov chaw no thiab qhov loj ntawm cov pejxeem. Thaum kawm qhov tsawg zaus ntawm kev hloov pauv ntawm P. infestans hom kab mob tsawg tsawg, cov naj npawb ntawm cov noob qoob loo uas tau xaiv rau cov khoom noj muaj txiaj ntsig zoo nyob rau tom qab kho nrog tshuaj lom neeg lossis lub cev hloov chaw feem ntau txiav txim siab. Raws li tuaj yeem pom los ntawm cov ntaub ntawv tshaj tawm nyob rau hauv Table 8, kev hloov pauv ntawm tib hom ntawm tib lub txawv ntawm qhov chaw nyob txawv tuaj yeem sib txawv los ntawm ntau qhov kev txiav txim ntawm qhov ntau. Kev hloov pauv loj hauv kev hloov pauv mus rau kev tiv taus hlau hlau tuaj yeem yog ib qho laj thawj ntawm kev sib xyaw ua ke ntawm cov kab uas cuam tshuam nrog nws nyob rau hauv cov xwm.
Kev nquag ntawm qhov tshwm sim los ntawm qhov hloov chaw, suav raws qhov kev sim ntsuas, tsis yog ib txwm piv rau cov txheej txheem uas tshwm sim hauv cov neeg hauv ntuj, rau cov laj thawj hauv qab no:
1. Nrog asynchronous nuclear fissions, nws yog tsis muaj peev xwm kwv yees qhov ntau zaus ntawm kev hloov pauv ib zaug ib lub cim nuclear. Yog li, feem ntau cov kev sim tsuas muab cov ntaub ntawv ncaj qha txog qhov ntau zaus ntawm kev hloov pauv, yam tsis muaj qhov txawv ntawm ob qho kev hloov pauv thiab ib qho xwm txheej tom qab mitosis.
2. Cov kev hloov pauv ib leeg feem ntau txo qhov sib npaug ntawm cov genome, yog li ntawd, nrog rau kev kis ntawm cov cuab yeej tshiab, qhov kev nyab xeeb ntawm cov kab mob tsawg dua. Feem ntau ntawm kev sim uas tau txais kev hloov pauv muaj txo kev ua phem thiab tsis tau sau tseg rau cov neeg nyob hauv ntuj. Yog li, qhov sib txheeb ntawm qhov sib npaug ntawm qhov tsis haum ntawm P. infestans hloov mus rau phenylamide fungicides thiab kev loj hlob ntawm ib qho khoom nruab nrab yog nruab nrab (-0,62), thiab qhov tsis kam ntawm fungicides thiab nruj ntawm cov nplooj qos (-0,65) (Derevyagina li al. , 1993), uas qhia hais tias lub cev muaj zog tsis sib haum. Kev sib hloov tsis kam rau qhov tsis kam dimethomorph kuj tau nrog poob qis dua ntawm kev muaj peev xwm (Bagirova li al., 2001).
3. Feem ntau ntawm qhov tshwm sim thiab qhov kev hloov pauv tau cuam tshuam thiab tsis tshwm sim lawv tus kheej phenotypically hauv kev sim, tab sis ua ib qho kev zais ntawm kev hloov pauv ntawm cov neeg ib txwm muaj. Hom kab mob sib xyaw cais nyob rau hauv kev sim tshuaj sim nqa kev hloov pauv tseem ceeb los sis ib nrab hloov pauv (Kulish and Dyakov, 1979). Thaj, nuclear diploidy piav qhia txog qhov ua tsis tau tiav kom tau txais kev hloov pauv nyob hauv qab ntawm UV irradiation uas ua rau muaj kev phom sij rau yav dhau los tiv thaiv ntau yam (McKee, 1969). Raws li tus sau sau cov kev suav, xws li kev hloov pauv tuaj yeem tshwm sim nrog cov zaus ntawm tsawg dua 1: 500000. Txoj kev hloov ntawm kev hloov pauv rau ib homozygous, phenotypically qhia hauv lub xeev tuaj yeem tshwm sim vim yog kev sib deev lossis kev sib deev rov los ua ke (saib hauv qab). Txawm li cas los xij, txawm hais tias qhov kev hloov pauv no tuaj yeem npog qhov tseem ceeb ntawm cov tsiaj loj hauv cov tsiaj qus hauv cenotic (multinucleated) mycelium thiab phenotypically tsau tsuas yog thaum tsim cov mononuclear zoospores.
Cov Lus 8. Zaus ntawm P. infestans hloov mus rau kev loj hlob-thaiv cov tshuaj hauv qab kev ua ntawm nitrosomethylurea (Dolgova, Dyakov, 1986; Bagirova li al., 2001)
Cuam | Kev sib hloov tsawg zaus |
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 |
Hlauaxil | 6,9 x 10-6 |
Cov pejxeem ntau thiab tsawg kuj ua si lub luag haujlwm uas txiav txim siab tshwm sim ntawm kev hloov chaw uas tshwm sim. Hauv cov neeg nyob coob heev, uas cov lej ntawm N> 1 / a, qhov twg tus lej ntawm qhov hloov pauv, kev hloov pauv tsis ua rau muaj qhov txawv txav (Kvitko, 1974).
Cov kev xam pom tau hais tias nrog kev pheej hmoo nruab nrab ntawm cov qos yaj ywm cog (35 me ntsis hauv cov nroj tsuag), 8x1012 cov noob kab ntsig yog tsim txhua hnub rau ntawm ib thaj chaw (hectak (Dyakov and Suprun, 1984). Thaj, xws li cov pejxeem muaj txhua qhov kev hloov pauv uas tso cai los ntawm hom kev sib pauv ntawm txhua qhov chaw nyob. Txawm hais tias muaj kev sib hloov tsis tshua muaj tshwm sim, muaj tshwm sim nrog zaus ntawm 10-9, yuav kis tau los ntawm ib txhiab tus neeg tawm ntawm lab tus neeg nyob ntawm ib thaj av ntawm ib thaj av ntawm cov qos yaj ywm. Txog kev hloov pauv tau tshwm sim nrog ntau zaus (piv txwv, 10-6), nyob rau hauv cov neeg zoo li no, ntau cov kev sib hloov tau tshwm sim txhua hnub (ib txhij ntawm ob qhov chaw), i.e. kev hloov pauv yuav hloov rov ua dua tshiab.
Tsiv teb tsaws
Txog P. infestans, ob lub ntsiab tseem ceeb ntawm kev tsiv teb tsaws chaw tau paub: kom kaw kev nrug deb (hauv thaj chaw cog qoob loo lossis thaj teb ze) los ntawm kis zoosporangia los ntawm huab cua dej tsaws tsag lossis dej nag, thiab mus rau qhov chaw nyob ntev - nrog cog cov taub lossis thauj cov txiv lws suav. Thawj txoj hau kev muab rau kev nthuav dav ntawm cov mob, thib ob - kev tsim foci tshiab hauv qhov chaw nyob deb ntawm thawj.
Kev sib kis ntawm cov kab mob nrog cov txiv lws suav thiab txiv hmab txiv ntoo tsis tsuas yog pab txhawb kev kis tus kabmob rau hauv cov chaw tshiab, tab sis tseem yog qhov tseem ceeb ntawm kev muaj caj ces ntawm cov neeg. Nyob hauv thaj av Moscow, cov qos yaj ywm raug zus, coj los ntawm ntau thaj tsam ntawm Lavxias thiab Western Europe. Cov txiv lws suav yog coj los ntawm cov cheeb tsam yav qab teb ntawm Russia (thaj av Astrakhan, thaj chaw Krasnodar, North Caucasus). Cov noob soob, uas tseem tuaj yeem ua qhov chaw kis tus kab mob (Rubin et al., 2001), kuj tseem tuaj ntawm cov cheeb tsam yav qab teb ntawm Russia, Suav, Tebchaws Europe thiab lwm lub tebchaws.
Raws li kev suav los ntawm E. Mayr (1974), kev hloov pauv ntawm cov neeg hauv nroog tau tshwm sim los ntawm kev hloov pauv tsis tshua muaj ntau tshaj 10-5 toj ib lub zos, thaum nyob hauv cov pejxeem qhib, qhov sib pauv vim yog cov txee tawm ntawm cov noob tsawg kawg yog 10-3 - 10-4.
Kev tsiv teb tsaws rau hauv cov kis tau tus kab mob yog lub luag haujlwm rau kev nkag ntawm P. infestans mus rau Tebchaws Europe, kis mus rau txhua thaj tsam hauv ntiaj teb uas qos yaj ywm loj tuaj; lawv tau ua rau muaj kev hloov pauv hauv cov pej xeem. Blight lig rau cov qos yaj ywm tau tshwm sim rau thaj chaw ntawm Lavxias teb sab yuav luag tib lub sijhawm nrog nws cov tsos hauv Western Europe.
Txij li thaum tus kab mob tau pom thawj zaug xyoo 1846-1847 hauv Baltic States thiab tsuas yog nyob rau xyoo tom qab kis tau nyob hauv Belarus thiab thaj tsam qaum teb qaum teb ntawm Russia, nws cov tebchaws Western Western tau pom tseeb. Thawj qhov chaw ntawm blight lig nyob rau hauv Lub Ntiaj Teb Qub yog tsis pom tseeb. Cov kev xav tau tsim los ntawm Fry et al. (Fry et al., 1992; Fry, Goodwin, 1995, Goodwin et al., 1994) qhia tias tus cab no yog tuaj ntawm Mexico mus rau North America, qhov twg nws kis tau hla qoob loo, thiab tom qab ntawd tau thauj mus rau Western Europe. (Daim duab 7).
Raws li qhov tshwm sim ntawm kev pheej rov ua dua qub (ob qhov cuam tshuam ntawm "bottleneck"), ib qho clones tau txais mus rau Tebchaws Europe, cov xeeb ntxwv ntawm cov uas ua rau muaj kev sib kis thoob plaws thoob plaws thaj av ntawm Lub Ntiaj Teb Qub, uas cov qos yaj ywm loj tuaj. Raws li pov thawj rau qhov kev xav no, cov sau phau ntawv hais, ua ntej, qhov ubiquitous tshwm sim tsuas yog ib hom kev sib koom ua ke (A1) thiab, qhov thib ob, homogeneity ntawm hom kev ntsuas ntawm cov hom kev kawm los ntawm ntau thaj tsam (tag nrho lawv yog los ntawm cov cim molecular, suav nrog 2 isozyme loci, DNA fingerprinting pattern, thiab cov qauv ntawm mitochondrial DNA zoo ib yam, thiab sib raug rau cov clone US-1 tau piav qhia hauv Asmeskas). Txawm li cas los xij, qee cov ntaub ntawv ua rau kev ua xyem xyav txog tsawg kawg ntawm qee qhov kev ua tiav ntawm qhov kev xav ua tsis tau. Kev soj ntsuam ntawm P. infestans mitochondrial DNA cais los ntawm herbarium qos yaj ywm kis thaum thawj thawj zaug mob epiphytotic thaum xyoo 40 tau qhia tias lawv sib txawv hauv cov qauv ntawm mitochondrial DNA los ntawm clone US-1, uas, yog li ntawd, tsawg tsis yog tib qho kev kis tus mob hauv Tebchaws Europe (Ristaino li al, 2001).
Qhov teeb meem blight lig tau mob hnyav dua nyob rau hauv 80s ntawm XX xyoo pua. Cov kev hloov hauv qab no tau tshwm sim:
1) Qhov nruab nrab quab yuam ntawm cov pejxeem tau nce, uas tau coj, tshwj xeeb, mus rau txoj kev sib kis dav dav ntawm cov teeb meem tshwm sim ntau tshaj plaws ntawm cov blight lig - kev puas tsuaj rau cov plaub thiab qia.
2) Muaj qhov hloov pauv nyob rau lub sijhawm lig ntawm blight on qos yaj ywm - txij lub Xya Hli txog rau Lub Xya Hli pib thiab txawm txog rau thaum Lub Rau Hli.
3) Hom A2 mating, uas yav dhau los tsis tuaj rau hauv Ntiaj Teb Qub, tau dhau los ua ubiquitous.
Cov kev hloov pauv tau ua ntej los ntawm ob qho xwm txheej: kev siv loj heev ntawm kev siv tshuaj tshiab fungicide metalaxyl (Schwinn thiab Staub, 1980) thiab qhov tshwm sim ntawm Mexico ua lub ntiaj teb xa tawm cov qos yaj ywm (Niederhauser, 1993). Raws li qhov no, ob qho laj thawj rau cov neeg hloov pauv tau muab tso ua ntej - kev hloov pauv ntawm hom mating nyob rau hauv kev cuam tshuam ntawm metalaxyl (Ko, 1994) thiab cov lus qhia loj ntawm cov kab mob tshiab nrog cov kab mob kis los ntawm Mexico (Fry thiab Goodwin, 1995). Txawm hais tias kev sib txuam ntawm cov mating hom nyob rau hauv kev cuam tshuam ntawm metalaxyl tau txais tsis yog los ntawm Ko, tab sis kuj nyob rau hauv kev ua haujlwm nqa tawm ntawm Moscow State University laboratory (Savenkova, Chherepennicova-Anikina, 2002), qhov thib ob kev xav yog preferable. Nrog rau cov tsos ntawm hom ob ntawm kev ua niam txiv, qhov kev hloov pauv loj tau tshwm sim nyob rau hauv cov genotypes ntawm Lavxias P. infestans leeg, suav nrog hauv kev tsis sib luag (isozyme thiab RFLP loci), zoo li hauv cov qauv ntawm mitochondrial DNA. Qhov nyuaj ntawm cov kev hloov pauv no tsis tuaj yeem piav qhia los ntawm kev ua ntawm metalaxyl; es tsis txhob muaj cov neeg tuaj txawv teb chaws tuaj tshiab tuaj hauv Mexico, uas, muaj kev tawm tsam ntau dua (Kato et al., 1997), tau tsiv tawm ntawm cov tub rog qub (US-1), ua neeg nyob hauv cov neeg. Kev hloov pauv ntawm kev txuam ntawm cov pej xeem nyob sab Europe tau tshwm sim nyob rau lub sijhawm luv luv - txij xyoo 1980 txog 1985 (Fry et al., 1992). Ntawm thaj chaw ntawm thaj chaw qub USSR, "cov kab mob tshiab" tau pom nyob hauv cov ntawv sau los ntawm Estonia hauv 1985, uas yog, ntxov dua hauv tebchaws Poland thiab Lub Tebchaws Yelemees (Goodwin et al., 1994). Lub sijhawm dhau los "tawv qub kis-US-1" hauv tebchaws Russia tau cais los ntawm ib tus kab mob lws suav hauv cheeb tsam Moscow hauv xyoo 1993 (Dolgova li al., 1997). Tsis tas li ntawd nyob rau Fabkis, cov kab mob "laus" tau pom hauv kev cog txiv lws suav txog thaum 90s thaum ntxov, uas yog, tom qab lawv tau ploj mus ntev ntawm cov qos yaj ywm (Leberton thiab Andrivon, 1998). Hloov P. Cov kab mob qis tshaj cuam tshuam rau ntau qhov kev ua haujlwm, nrog rau cov kev coj ua uas tseem ceeb, thiab nce qhov ua rau muaj kev phom sij.
Kev sib deev rov qab los
Hais txog kev sib deev rov qab los ua kom muaj qhov sib txawv, nws yog qhov tsim nyog, ua ntej, qhov muaj ob hom kev sib tham nyob hauv cov neeg nyob hauv ib qho sib piv uas ze rau 1: 1, thiab, ob, lub xub ntiag ntawm cov pejxeem thawj pauv.
Qhov ntsuas ntawm kev sib piv yog ntau yam nyob rau ntau hom neeg thiab txawm nyob hauv ntau xyoo hauv tib neeg pej xeem (Cov Lus 9,10, 90) Cov laj thawj rau cov kev hloov pauv hnyav li no ntawm cov mating ntau hom hauv cov neeg (xws li, hauv tebchaws Russia lossis Ixayees nyob rau thaum 2002s ntawm lub xyoo pua xeem) tsis paub, tab sis nws ntseeg tias qhov no yog vim muaj kev qhia ntxiv ntawm cov pob zeb sib tw (Cohen, XNUMX).
Qee cov ntaub ntawv tsis ncaj qhia qhia tias cov txheej txheem ntawm kev sib deev hauv qee xyoo thiab hauv qee thaj chaw:
1) Kev tshawb fawb ntawm cov neeg los ntawm thaj av Moscow tau qhia tias hauv 13 cov neeg nyob hauv qhov kev faib ua feem ntawm A2 hom kev ua niam txiv yog tsawg dua 10%, tag nrho cov kev sib txawv ntawm kev tshuaj ntsuam tau muab xam rau peb isozyme loci yog 0,08, thiab hauv 14 cov neeg nyob hauv qhov kev faib ua feem ntawm A2 tshaj 30%, noob caj noob ces yog ob zaug siab (0,15) (Elansky li al., 1999). Yog li, qhov muaj peev xwm ntau dua ntawm cov neeg sib deev, ntau dua cov caj ces ntawm cov neeg.
2) Qhov sib txheeb ntawm qhov sib piv ntawm mating hom nyob rau hauv cov pejxeem thiab qhov siv ntawm oospore tsim tau pom nyob hauv tebchaws Israel (Cohen et al., 1997) thiab hauv Holland
(Flier li al., 2004). Peb qhov kev tshawb fawb tau qhia tias, nyob rau hauv cov neeg nyob rau hauv uas cais tawm nrog A2 mating hom lees rau 62, 17, 9, thiab 6%, oospores tau pom nyob rau hauv 78, 50, 30, thiab 15% ntawm cov nplooj ntsuas uas tau soj ntsuam (muaj 2 lossis ntau qhov chaw).
Cov qauv uas muaj 2 lossis ntau qhov chaw tsis ntau yuav tsum muaj oospores dua li cov piv txwv nrog 1 qhov nqaij (32 thiab 14% ntawm cov hnoos qeev), (Apryshko li al., 2004).
Oospores tau muaj ntau dua nyob hauv nplooj ntawm nruab nrab thiab qis txheej ntawm cov qos yaj ywm cog (Mytsa et al., 2015; Elansky li al., 2016).
3) Hauv qee thaj chaw, tau nrhiav pom cov neeg caj ceg txawv, tshwm sim uas cuam tshuam nrog kev sib deev sib deev. Yog li, hauv tebchaws Poland hauv 1989 thiab hauv Fabkis xyoo 1990, muaj homozygous rau ntshav qabzib-6-
ntsev yog isomerase (GPI 90/90). Txij li yav dhau los tsuas yog 10/90 heterozygotes tau ntsib rau 100 xyoo, homozygosity raug ntaus los ntawm kev sib daj sib deev (Sujkowski li al., 1994). Hauv Colombia (Tebchaws USA), cais tawm sib txuas A2 nrog GPI 100/110 thiab A1 nrog GPI 100/100 yog ib qho muaj, txawm li cas los xij, qhov kawg ntawm 1994 lub caij (Lub Yim Hli 16 thiab Cuaj Hlis 9), cov kab mob sib xyaw nrog cov tshuaj tiv thaiv genotypes (A1 GPI 100/110) thiab A2 GPI 100/100) (Miller li al., 1997).
4) Hauv qee cov pejxeem los ntawm Poland (Sujkowski li al., 1994) thiab North Caucasus (Amatkhanova li al., 2004), kev faib tawm ntawm cov ntiv tes DNA loci thiab allozyme protein loci sib raug rau Hardy-Weinberg kev faib tawm, uas qhia tau
txog qhov siab sib koom ntawm kev koom tes ntawm kev sib deev rov ua dua kom txawv txav ntawm cov neeg. Hauv lwm thaj chaw ntawm Russia, tsis muaj kev sib tiv tauj mus rau Hardy-Weinberg kev faib tawm hauv cov neeg tau pom, tab sis qhov pom ntawm cov kab mob sib txuas tau pom, qhia txog qhov muaj peev xwm ntawm cov pob txha luam tawm (Elansky et al., 1999).
5) Kev sib txawv ntawm caj ces (GST) nruab nrab ntawm cov kab mob sib txawv nrog cov sib txawv ntawm cov khub (A1 thiab A2) tau qis dua ntawm cov neeg sib txawv (Sujkowski li al., 1994), uas qhia txog kev sib daj sib deev.
Nyob rau tib lub sijhawm, qhov kev koom ua ke ntawm kev sib deev rov qab rau cov neeg sib txawv tsis tuaj yeem ua ntau. Qhov kev koom tes no tau suav rau cov pejxeem ntawm cheeb tsam Moscow (Elansky et al., 1999). Raws li cov kev suav ntawm Lewontin (1979), "recombination, uas tuaj yeem tsim cov hloov pauv tshiab los ntawm ob lub zos uas muaj ntau zaus tsis dhau cov khoom ntawm lawv cov kev txwv sib kis, ua tau zoo yog tias qhov tseem ceeb ntawm heterozygosity rau ob qho sib npaug twb tau siab lawm."
Nrog rau qhov sib piv ntawm ob hom kev ua khub, uas ib txwm muaj rau cheeb tsam Moscow, sib npaug rau 4: 1, qhov tso tawm zaus yuav yog 0,25. Qhov tshwm sim uas hla cov kab mob tsis txaus ntseeg yuav yog qhov tsis haum rau ob ntawm peb qhov kev kawm isozyme loci hauv cov neeg kawm tshawb nrhiav yog 0,01 (2 leeg tawm ntawm 177). Yog li ntawd, qhov tshwm sim ntawm qhov tshwm sim ntawm ob chav heterozygotes raws li kev qhia tawm yuav tsum tsis pub tshaj lawv cov khoom sib npaug los ntawm qhov tshwm sim ntawm kev hla (0,25x0,02x0,02) = 10-4, i.e. kev sib deev recombinants feem ntau tsis poob rau hauv kev kawm piv txwv ntawm cov raug mob. Cov kev xam no tau ua rau cov pej xeem los ntawm cheeb tsam Moscow tus yam ntxwv txawv txav ntawm qhov siab. Hauv cov neeg monomorphic zoo li cov Siberian, cov txheej txheem kev sib deev, txawm tias nws tshwm sim hauv cov neeg cais, tsis tuaj yeem cuam tshuam lawv cov caj ces sib txawv.
Ntxiv rau, P. infestans yog tus cwj pwm los ntawm kev mob ntsws ntau dhau nyob hauv meiosis, uas ua rau muaj qhov tsis nco qab (Carter et al., 1999). Cov kev ua txhaum cai no txo qis ntawm kev ua me nyuam ntawm hybrids.
Parasexual recombination, mitotic gene hloov dua siab tshiab
Hauv kev sim rau cov splicing ntawm P. infestans lim nrog kev hloov pauv hauv kev kam rau cov kev loj hlob sib txawv inhibitors, kev rov tshwm sim ntawm misolates tsis kam rau ob lub inhibitors tau pom (Shattock thiab Shaw, 1975; Dyakov, Kuzovnikova, 1974; Kulish, Dyakov,
Xyoo 1979). Cov kab mob ua rau ob txoj kev loj hlob inhibitors tshwm sim los ntawm heterokaryotization ntawm mycelium, thiab hauv qhov no lawv tau muab pov tseg thaum kev tsim kho los ntawm mononuclear zoospores (Judelson, Ge Yang, 1998), lossis tsis tau muab pov tseg rau hauv cov xeeb ntxwv monozoosporous, vim lawv muaj tetraploid (txij li qhov pib cais tawm yog diploid) nuclei (K , Xyoo 1979). Heterozygous diploids cais tsawg kawg vim yog haploidization, chromosome nondisjunction, thiab mitotic hla dhau (Poedinok li al., 1982). Qhov ntau zaus ntawm cov txheej txheem no tuaj yeem tau nce nrog kev pab ntawm qee yam kev ua ntawm heterozygous diploids (piv txwv, UV irradiation ntawm germinating spores).
Txawm hais tias qhov tsim ntawm kev noj qab haus huv hybrids nrog ob qho kev tawm tsam tshwm sim tsis tsuas yog nyob rau hauv vitro, tab sis kuj nyob rau hauv qos tub kis mob nrog kev sib xyaw ntawm kev hloov tsiaj (Kulish li al., 1978), nws yog qhov nyuaj rau kev ntsuam xyuas lub luag haujlwm ntawm parasexual recombination nyob rau hauv tiam ntawm genotypes tshiab hauv cov neeg. Qhov ntau zaus ntawm segregants tsim vim haploidization, nondisjunction ntawm chromosomes thiab mitotic hla dhau yam tsis muaj teebmeem tshwj xeeb yog kev tsis pom zoo (tsawg dua 10-3).
Qhov tshwm sim ntawm homozygous segregants ntawm heterozygous hom yuav tau los ntawm ob txoj kev hla dhau thiab kev hloov pauv ntawm cov kab mob hloov pauv, uas hauv P. sojae tshwm sim nrog zaus ntawm 3 x 10-2 txog 5 x 10-5 toj ib thaj chaw, nyob ntawm ib hom kab mob (Chamnanpunt li al. , Xyoo 2001).
Txawm hais tias qhov ntau zaus ntawm kev tshwm sim ntawm heterokaryons thiab heterozygous diploids hloov tawm qhov kev xav tsis meej (nce txog kaum tawm feem pua), cov txheej txheem no tsuas yog tshwm sim thaum cov kab kev hloov pauv tau los ntawm tib hom kab mob spliced. Thaum siv cov nyias muaj nyias sib cais los ntawm cov xwm, heterokaryotization tsis tshwm sim (los yog tshwm sim nrog qhov tsis tshua muaj ntau zaus) vim muaj cov khoom noj tsis haum (Poedinok thiab Dyakov, 1981; Anikina et al., 1997b; Cherepennikova-Anikina li al., 2002). Yog li, lub luag haujlwm ntawm parasexual recombination tuaj yeem raug txo kom tsawg ntxiv rau kev tso tseg nyob rau hauv heterozygous nuclei thiab kev hloov pauv ntawm cov neeg ib leeg mus rau ib lub xeev homozygous tsis muaj txheej txheem kev sib deev. Txoj kev no yuav yog qhov tseem ceeb ntawm cov kabmob sib kis nrog kev hloov pauv hnyav lossis ib nrab ntawm cov khoom siv fungicide tsis sib xws. Nws txoj kev hloov mus rau lub xeev homozygous vim yog tus txheej txheem parasexual yuav nce lub zog ntawm cov cab kuj ntawm kev hloov pauv (Dolgova, Dyakov, 1986).
Introgression ntawm cov noob
Heterothallic hom Phytophthora muaj peev xwm los cuam tshuam nrog kev tsim ntawm cov xov paj ntoo (saib Vorob'eva thiab Gridnev, 1983; Sansome li al., 1991; Veld li al., 1998). Lub ntuj hybrid ntawm ob hom Phytophthora yog li txhoj puab heev uas nws tua ntau txhiab leej neeg hauv tebchaws Askiv (Brasier li al., 1999). P. infestans tuaj yeem tshwm sim nrog lwm hom ntawm cov genus (P. erythroseptica, P. nicotianae, P. Cactorum, thiab lwm yam) ntawm cov tswv tsev dav thiab hauv av, tab sis tsis muaj ntaub ntawv me me hauv cov ntawv nyeem txog kev muaj peev xwm sib cais. Hauv qab kuaj qhov mob, cov kab mob nruab nrab ntawm P. infestans thiab P. Mirabilis tau txais (Goodwin thiab Fry, 1994).
Cov Lus 9. Qhov feem ntawm P. infestans hom mob nrog A2 mating hom nyob hauv ntau lub teb chaws ntawm lub ntiaj teb nyob rau lub sijhawm txij xyoo 1990 txog 2000 (raws li cov ntaub ntawv ntawm cov ntaub ntawv qhib thiab cov vev xaib www.euroblight.net, www.eucablight.org)
lub teb chaws | 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) | ||||
Fabkis | 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) | ||||||||||
Qaum teb. Ireland | 10 (41) | 9 (58) | 1 (106) | 0 (185) | 0 (18) | 0 (56) | 0 (35) | 0 (26) | |||
Netherlands | 7 (41) | 5 (276) | 24 (377) | 44 (353) | 23 (185) | ||||||
Norway | 25 (446) | 28 (156) | 8 (39) | 18 (257) | 38 (197) | ||||||
Peru | 0 (34, 1984 -86) | 0 (287, 1997-98) | 0 (112) | 0 (66) | |||||||
Poland | 19 (180) | 21 (142) | 33 (256) | 26 (149) | 35 (70) | ||||||
Scotland | 25 (147) | 11 (163) | 22 (189) | 5 (22) | |||||||
Sweden | 25 (263) | 62 (258) | 49 (163) | ||||||||
Wales | 0 (16) | 7 (97) | 0 (48) | 0 (25) | |||||||
Kauslim teb | 36 (42) | 10 (130) | 15 (98) | ||||||||
Tuam Tshoj | 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) |
Rooj 10. Qhov feem ntawm P. infestans hom mob nrog A2 mating hom nyob hauv ntau lub teb chaws ntawm lub ntiaj teb nyob rau lub sijhawm txij xyoo 2000 txog 2011
lub teb chaws | 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 koom pheej | 35 (31) | 54 (64) | 38 (174) | 12 (80) | |||||||
P "RμSЂRјR ° RЅRёSЏ | 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 | ||||||||
Fabkis | 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 | |||||||
Qaum teb. Ireland | 0 (38) | 0 (58) | 0 (40) | 0 (24) | 5 (54) | 0 (18) | 27 (578) | 45 (239) | 36 (213) | 82 (60) | 10 (80) |
Netherlands | 66 (24) | 93 (15) | 91 (11) | ||||||||
Norway | 39 (328) | 3 (115) | 12 (19) | ||||||||
Peru | 0 (36) | ||||||||||
Poland | 25 (46) | 10 (30) | 85 (20) | 38 (44) | 75 (66) | 55 (56) | 65 (35) | 72 (81) | 85 (21) | ||
Scotland | 3 (213) | 2 (474) | 24 (135) | 86 (337) | 88 (386) | 74 (172) | |||||
Sweden | 60 (277) | 39 (87) | |||||||||
Slovakia | 0 (36) | 14 (26) | 62 (26) | 0 (26) | |||||||
Wales | 25 (12) | 68 (106) | 80 (88) | 92 (143) | 75 (45) | ||||||
Kauslim teb | 46 (26) | ||||||||||
Brazil | 0 (49) | 0 (30) | |||||||||
Tuam Tshoj | 10 (30) | 0 (6) | 0 (6) | ||||||||
Nyab Laj teb | 0 (294, 2003-04) | ||||||||||
Uganda | 0 (8) |
Lub zog ntawm cov qauv ntawm genotypic ntawm cov neeg
Kev hloov pauv ntawm cov keeb kwm genotypic ntawm P. cov kab mob ua rau muaj peev xwm tshwm sim nyob rau hauv tus neeg tsiv teb tsaws ntawm cov pob zeb tshiab los ntawm lwm cheeb tsam, kev ua liaj ua teb (kev hloov pauv ntawm ntau yam, daim ntawv thov ntawm fungicides), thiab huab cua. Kev cuam tshuam sab nraud cuam tshuam rau cov pob zeb sib txawv ntawm ntau theem ntawm lub neej hauv lub neej; yog li ntawd, cov neeg txhua xyoo muaj kev hloov pauv hloov raws qhov muaj peev xwm ntawm cov noob yuav raug xaiv, vim muaj kev hloov pauv ntawm txoj haujlwm tseem ceeb ntawm cov dej nyab thiab xaiv.
Cawv ntawm ntau yam
Cov hom tshiab nrog cov muaj txiaj ntsig zoo rau ntsug tsis kam (R-noob) yog qhov muaj peev xwm xaiv tau, xaiv cov clones nrog cov virulence genes hauv P. infestans. Thaum tsis muaj nonspecific tsis kam nyob rau hauv cov qos ntau yam, uas inhibits qhov kev loj hlob ntawm cov pathogen cov pej xeem, cov txheej txheem ntawm kev hloov lub pob zeb tseem ceeb hauv cov pejxeem tshwm sim sai. Yog li, tom qab kis tau rau thaj tsam ntawm Moscow thaj tsam ntawm Domodedovsky ntau yam, uas muaj cov R3 tsis kam cov noob, qhov zaus ntawm clones virulent rau cov hom no tau nce ntawm 0,2 txog 0,82 hauv ib xyoos (Dyakov thiab Derevjagina, 2000).
Txawm li cas los xij, kev hloov pauv ntawm ntau zaus ntawm cov kab mob virulence (pathotypes) hauv cov pejxeem muaj tshwm sim tsis yog nyob rau hauv kev cuam tshuam ntawm kev cog qoob loo ntau yam. Piv txwv li, hauv Belarus txog xyoo 1977, cov pob zeb nrog virulence noob 1 thiab 4 muaj tswv yim, uas tau tshwm sim los ntawm kev cog qoob loo ntawm cov qos yaj ywm nrog cov tsis sib xws R1 thiab R4 (Dorozhkin, Belskaya, 1979). Txawm li cas los xij, qhov kawg ntawm xyoo 70 ntawm lub xyoo pua XX, cov qog tau tshwm sim nrog cov caj ces sib txawv thiab lawv cov kev sib txuas ua ke, thiab cov kev sib xyaw tsis sib xws yeej tsis tau siv rau hauv kev tua cov qos yaj ywm (ntxiv rau virulence noob) (Ivanyuk li al., 2002). Qhov laj thawj rau qhov tshwm sim ntawm cov pob zeb no, pom meej, yog vim tias cov neeg tsiv teb tsaws chaw mus rau Tebchaws Europe ntawm cov khoom kis tau los ntawm Mexico nrog cov qos ntoo. Nyob hauv tsev, cov pob zeb no tsim tsis tsuas yog rau cov qos yaj ywm cog qoob loo, tab sis kuj rau ntawm cov tsiaj qus coj ntau hom kev tiv thaiv ntawm cov noob;
Raws li rau ntau yam nrog nonspecific tsis kam, lawv, los ntawm kev txo tus nqi ntawm kev ua me nyuam ntawm cov kab mob, ncua qhov kev hloov pauv ntawm nws cov neeg, uas, raws li twb tau hais, yog ib qho haujlwm ntawm tus naj npawb. Txij li kev ua nruj ua tsiv yog polygenic, cov pob txha uas muaj ntau dua ntawm cov neeg ua haujlwm rau "kev ua phem" ntau zuj zus sai dua cov neeg loj. Yog li no, cov haiv neeg muaj kev kub ntxhov tsis yog cov khoom siv ntawm kev yoog rau ntau hom tsis sib xws, tab sis, ntawm qhov tsis sib xws, ntau dua yuav raug kuaj pom hauv cov cog ntawm cov qhov muaj kev cuam tshuam ntau yam uas yog cov muaj ntau ntawm cov cab cab.
Yog li, hauv tebchaws Russia cov haiv neeg muaj coob leej ntau tus ntawm P. Cov kab mob me me tau pom hauv cov cheeb tsam ntawm txhua xyoo epiphytoties (cov neeg tuaj ntawm Sakhalin, Leningrad, thiab Bryansk cheeb tsam) Cov kev nruj ntawm cov neeg no tau ua rau muaj siab dua cov neeg Mev (Filippov et al., 2004).
Ib qho ntxiv, tsawg dua oospores tsim nyob rau hauv cov nplooj ntawm cov hom tsis muaj zog dua li ntawm cov raug tau yooj yim (Hanson thiab Shattock, 1998), uas yog, qhov tsis muaj peev xwm ntawm ntau yam kuj txo cov cab muaj peev xwm rov ua dua tshiab thiab muaj peev xwm hloov lwm txoj hauv kev.
Cawv ntawm fungicides
Fungicides tsis tsuas yog txo cov phytopathogenic fungi, i.e. cuam tshuam rau qhov ntau ntawm cov yam ntxwv ntawm lawv cov neeg, tab sis lawv kuj tuaj yeem hloov qhov nquag ntawm cov tib neeg genotypes, i.e. cuam tshuam qhov muaj txiaj ntsig ntawm cov pej xeem. Ntawm qhov ntsuas tseem ceeb tshaj plaws ntawm cov pej xeem hloov pauv nyob rau hauv kev cuam tshuam ntawm fungicides yog cov hauv qab no: kev hloov pauv ntawm cov tshuaj fungicides, kev hloov pauv ntawm kev txhoj puab heev thiab virulence, thiab kev hloov pauv ntawm cov kab hauv kev sib tua.
Cawv ntawm fungicides ntawm kev tawm tsam thiab nruj ntawm cov neeg
Cov theem ntawm cov cawv ntawd tau txiav txim siab, ua ntej txhua yam, los ntawm hom kev siv tshuaj tua kab, uas tuaj yeem muab faib ua cov tsis muaj zog, polysite, oligosite thiab monosite.
Lub qub suav nrog feem ntau tiv tauj fungicides. Kev tawm tsam rau lawv (yog tias nws ua tau txhua lub sijhawm) raug tswj los ntawm coob tus ntawm cov noob tsis tshua muaj zog. Cov khoom no txiav txim siab qhov tsis pom ntawm kev hloov pauv ntawm cov neeg ntawm lub cev tom qab kho nrog fungicides (txawm hais tias nyob rau qee qhov kev sim qee qhov nce ntxiv hauv kev tawm tsam tau txais). Cov pej xeem fungal tshwj tseg tom qab txau nrog tiv thaiv fungicides muaj ob pawg kab mob:
1) Txoj hlab ntsha khaws cia rau hauv thaj chaw ntawm cov nroj tsuag tsis kho nrog cov tshuaj. Txij li thaum tsis muaj kev sib cuag nrog lub fungicide, kev txhoj puab heev thiab tawm tsam ntawm cov kab no tsis hloov.
2) Txoj hlab ntsha hauv kev sib cuag nrog tus neeg tua hluav taws, qhov siab ntawm uas nyob ntawm cov ntsiab lus ntawm kev sib cuag tau qis dua kev tuag. Raws li tau hais los saum toj no, kev tawm tsam ntawm feem ntawm cov pejxeem no tseem tsis hloov, txawm li cas los xij, vim muaj qee qhov ua rau lub fungicide txawm tias nyob rau hauv sublethal concentration ntawm cov metabolism hauv lub xovtooj ntawm fungal, kev tawm dag zog dav dav thiab nws cov parasitic tiv thaiv, kev txhoj puab heev, txo qis (Derevyagina thiab Dyakov, 1990).
Yog li, txawm tias ib feem ntawm cov pejxeem uas tsis tau tuag, pom kev sib cuag nrog lub fungicide, muaj qhov kev ua kom tsis muaj zog thiab tsis tuaj yeem yog lub hauv paus ntawm epiphytotics. Yog li, kev ua tib zoo saib, uas txo qis ntau zaus ntawm cov neeg tsis sib haum nrog fungicide, yog qhov xwm txheej rau kev ua tiav ntawm kev tiv thaiv kev tiv thaiv. Kev tawm tsam oligosite fungicides yog tswj los ntawm ob peb lub noob ntxiv.
Kev hloov pauv ntawm txhua tus noob ua rau qee qhov nce ntxiv ntawm qhov ua haujlwm tiv thaiv, thiab feem ntau ntawm kev tuaj yeem tiv thaiv vim yog qhov sib ntxiv ntawm kev hloov pauv. Yog li ntawd, qhov nce nyob rau hauv tsis kam tshwm sim hauv cov kauj ruam. Ib qho piv txwv ntawm cov kauj ruam kev nce qib ntawm kev tawm tsam yog kev hloov pauv nyob rau hauv kev tawm tsam fungicide dimethomorph, uas tau siv dav los tiv thaiv qos yaj ywm los ntawm blight lig. Dimethomorph tsis kam yog polygenic thiab ntxiv rau. Ib qho kev hloov pauv ib nyuag qhov siab ntxiv.
Txhua qhov kev hloov pauv zuj zus txo lub hom phiaj loj thiab, yog li ntawd, qhov ntau zaus ntawm kev hloov pauv tshiab (Bagirova li al., 2001). Qhov nce ntawm nruab nrab tsis kam ntawm cov pejxeem tom qab ntau yam kev kho mob nrog oligosite fungicide tshwm sim stepwise thiab maj mam. Tus nqi ntawm cov txheej txheem no tau txiav txim los ntawm tsawg kawg yog peb yam: kev hloov pauv ntawm cov tsis kam ntawm cov noob, qhov sib npaug ntawm qhov tsis kam (qhov sib piv ntawm cov tshuaj tua kab mob ntawm qhov muaj peev xwm tiv taus nyob rau hauv kev sib raug zoo nrog cov rhiab) thiab qhov cuam tshuam ntawm kev hloov pauv hauv cov kev tawm tsam ntawm kev tawm dag zog.
Qhov nquag ntawm kev tshwm sim ntawm txhua qhov kev sib hloov tom qab muaj qis dua qhov ua ntej; yog li ntawd, cov txheej txheem muaj tus cwj pwm damping (Bagirova li al., 2001). Txawm li cas los xij, yog tias cov txheej txheem sib txuas (sib deev lossis sib deev) tshwm sim nyob rau hauv cov pejxeem, tom qab ntawd nws yog qhov muaj peev xwm los muab cov kev sib txawv ntawm cov niam txiv tso rau hauv ib qho kev sib txuas thiab sib txuas ntxiv. Yog li, cov pej xeem panmix tau txais kev tawm tsam sai dua li cov neeg txom nyem, thiab nyob rau tom kawg, cov neeg tsis muaj kev tsis sib haum xeeb sai dua li cov pej xeem sib cais los ntawm cov teeb meem no. Hauv qhov no, qhov pom muaj ntawm cov neeg tsis sib xws hauv qhov sib txawv ntawm cov sib xyaw ua ke nrawm nrawm cov txheej txheem tau txais cov txheej txheem tiv taus oligosite fungicides.
Qhov thib ob thiab thib peb yam tsis ua rau kom muaj kev txuam nrog sai ntawm cov kab dimethomorph-resistant nyob rau hauv ntau haiv neeg. Txhua qhov kev hloov pauv tom qab kwv yees ob npaug ntawm qhov kev tawm tsam, uas yog qhov tsis txaus ntseeg, thiab tib lub sijhawm txo qhov kev loj hlob ntxiv hauv qhov chaw tsis muaj zog thiab kev tawm tsam (Bagirova li al., 2001; Qia, Kirk, 2004). Tej zaum yog vim li cas thiaj tsis muaj kev tawm tsam los ntawm cov kab mob P. infestans, txawm tias sau los ntawm cov nroj tsuag qos tau kho nrog dimethomorph.
Ib tug pej xeem kho nrog tus mob oligosite fungicide yuav tseem muaj ob pawg ntawm cov kab mob: cov neeg uas tsis tau ntsib nrog tus neeg tua tsiaj, thiab yog li ntawd tsis tau hloov cov yam ntxwv thaum pib (yog tias muaj kev tawm tsam pom ntawm cov pab pawg no, lawv yuav tsis tuaj yeem vim muaj ntau dua kev sib tw thiab kev sib tw ntawm cov neeg muaj peev xwm), thiab leeg mob chwv nrog sublethal ntau ntawm fungicide. Nws yog ntawm lub tom kawg uas tsub zuj zuj ntawm cov kab kev tawm tsam yog qhov ua tau, vim hais tias ntawm no lawv muaj qhov zoo dua qhov tsis nkag siab.
Yog li no, thaum siv cov tshuaj oligosite fungicides, nws tsis yog li kev kho mob kom zoo uas yog qhov tseem ceeb raws li lub siab xav ntawm cov tshuaj, ob peb zaug siab dua li cov koob tshuaj tuag, vim tias nrog stepwise mutagenesis, qhov pib tsis kam ntawm hom hloov pauv muaj tsawg.
Thaum kawg, kev hloov pauv nyob hauv kev tawm tsam rau monosite fungicides yog qhov qhia tau zoo heev, uas yog, ib qho kev hloov pauv tuaj yeem tshaj tawm cov qib siab ntawm kev tiv thaiv kom ua tiav qhov kev poob tawm. Yog li ntawd, qhov nce ntawm qhov tsis kam ntawm cov pej xeem tshwm sim sai heev.
Ib qho piv txwv ntawm cov fungicides no yog phenylamides, suav nrog cov tshuaj tua kab mob uas feem ntau siv, metalaxyl. Kev hloov pauv ntawm kev tsis taus rau nws tshwm sim nrog lub siab, thiab cov neeg kawm ntawv nyob rau hauv kev hloov pauv muaj ntau heev - nws ntau dhau ntawm cov kab mob rhiab los ntawm qhov chaw ntawm ib txhiab lossis ntau dua (Derevyagina li al., 1993). Txawm hais tias qhov kev loj hlob ntawm lub cev thiab kev nruj kev tsiv ntawm cov kev hloov pauv tau txo qis rau keeb kwm yav dhau los ntawm kev tuag ntawm cov raug mob los ntawm cov kab mob fungic, tus naj npawb ntawm cov pej xeem tiv thaiv tau nce zuj zus thiab, nyob rau tib lub sijhawm, nws kev txhoj puab heev yog nce ntxiv. Yog li no, tom qab ob peb xyoos ntawm kev siv cov tshuaj tua hluav taws, kev txhoj puab heev ntawm kev tawm tsam tsis tuaj yeem tsis tuaj yeem sib npaug xwb qhov kev txhoj puab los ntawm kev nkag siab, tab sis kuj dhau nws (Derevyagina, Dyakov, 1992).
Ua rau ntawm kev sib deev rov los
Txij li feem ntau tshwm sim ntawm A2 mating hom hauv P. infestans cov neeg tau txais kev sib tw nrog kev siv hlau ntau dhau los tiv thaiv blight lig, nws tau xav tias metalaxyl ntxias mating hom hloov dua siab tshiab. Hauv P. parasitica, xws li hloov dua siab tshiab nyob rau hauv kev nqis tes ntawm Chloroneb thiab metalaxyl tau sim sim (Ko, 1994). Ib nqe lus ntawm qhov nruab nrab nrog qhov tsis tshua muaj siab ntawm metalaxyl ua rau muaj qhov tshwm sim ntawm homothallic cais los ntawm ib hom kab ntawm P. infestans rhiab rau metalaxyl nrog mating hom A1 (Savenkova thiab Cherepnikova-Anikina, 2002). Thaum lub sijhawm txuas ntxiv ntawm cov ntawv xov xwm nrog kev sib zog ntau dua ntawm metalaxyl, tsis yog ib leeg cais ntawm A2 hom kev ua ke tau tshawb pom, txawm li cas los xij, feem ntau cais, thaum hla nrog A2 cais, tsis txhob muaj oospores, tsim ugly mycelium tsub zuj zus thiab tau huv. Cov lus hla ntawm cov kab kev cuam tshuam muaj hom A2 mating hom ntawm kev tshaj xov xwm nrog lub siab ntawm cov hlau hlau tau tso cai rau peb kuaj peb hom kev hloov mating hom: 1) ua kom tsis muaj menyuam thaum hla nrog A1 thiab A2 cais tawm; 2) homotallism (kev tsim ntawm oospores hauv monoculture); 3) kev hloov pauv ntawm A2 mating type rau A1. Yog li, metalaxyl tuaj yeem ua rau cov kev hloov pauv ntawm hom kev sib raug zoo hauv P. infestans cov neeg muaj peev xwm thiab, yog li, kev sib daj sib deev hauv lawv.
Ua rau ntawm vegetative recombination
Qee cov noob rau tshuaj tua kab mob ua kom muaj zog nce ntxiv ntawm hyphal heterokaryotization thiab nuclear diploidization (Poedinok thiab Dyakov, 1981). Raws li tau sau tseg ua ntej, kev hloov heterokaryotization ntawm hyphae thaum lub sijhawm fusion ntawm cov sib txawv ntawm P. cov kab mob tshwm sim tsis tshua muaj tshwm sim vim muaj qhov tshwm sim ntawm kev noj haus tsis sib xws hauv cov kab mob no. Txawm li cas los xij, caj ces ntawm kev tawm tsam rau qee cov tshuaj tua kab mob tuaj yeem muaj kev phiv, qhia tawm hauv kev kov yeej kev noj haus tsis sib xws. Cov cuab yeej no tau muaj los ntawm 1S-1 mutant streptomycin resistance gene. Lub xub ntiag ntawm cov kev hloov pauv zoo li no hauv cov neeg thaj av ntawm phytophthora tuaj yeem nce cov kev hloov ntawm cov noob ntawm cov kab mob sib txawv thiab ua kom nrawm rau kev yoog raws txhua tus pejxeem mus rau ntau yam tshiab lossis fungicides.
Qee yam fungicides thiab tshuaj tua kab mob tuaj yeem cuam tshuam qhov ntau zaus ntawm mitotic recombination, uas tseem tuaj yeem hloov pauv genotype hauv cov neeg. Cov dav siv fungicide benomyl khi rau beta-tubulin, cov protein los ntawm cov microtubules ntawm cytoskeleton tau tsim, thiab yog li cuam tshuam cov txheej txheem ntawm kev sib cais ntawm chromosome hauv lub anaphase ntawm mitosis, nce lub cev ntawm mitotic recombination (Hastie, 1970).
Lub fungicide para-fluorophenylalanine, siv los kho cov kab mob Dutch hauv elms, muaj cov khoom ntiag tug. Para-fluorophenylalanine tau nce ntau ntxiv ntawm qhov rov ua dua nyob hauv heterozygous diploids P. infestans (Poedinok li al., 1982).
Cyclic kev hloov pauv ntawm cov keeb kwm ntawm cov neeg nyob hauv lub neej hloov pauv P. infestans
Kev loj hlob puag ncig los ntawm P. infestans hauv cheeb tsam huab cua sov muaj 4 ntu.
1) Lub sijhawm ntawm kev nthuav dav ntawm kev nthuav dav ntawm cov pejxeem (polycyclic theem) nrog ntau tiam neeg luv. Cov theem no feem ntau pib thaum Lub Xya Hli thiab kav ntev txog 1,5-2 hli.
2) Cov theem ntawm kev txwv qhov kev loj hlob ntawm cov pejxeem vim qhov txo qis ntawm qhov kev faib ua feem tsis muaj qhov tsis zoo lossis qhov pib ntawm huab cua tsis zoo. Theem no hauv kev ua liaj ua teb uas ua kom ntxov ua ntej lub caij nplooj zeeg cov khoom tawm tshem tawm hauv lub voj voog txhua xyoo.
3) Cov theem ntawm lub caij ntuj no hauv lub raj, nrog rau qhov kev txo qis ntawm cov neeg loj vim tias kis tau tus mob ntawm lub cev, qhov kev txhim kho qeeb ntawm kev kis tus kab mob hauv lawv, qhov tsis muaj qhov rov kis tau ntawm cov tub ntxhais, ua rau lwj thiab culling ntawm cov cuam tshuam rau hauv qab ntawm cov khoom cia.
4) Cov theem ntawm txoj kev loj hlob qeeb hauv av thiab ntawm cov nplooj ntoo (monocyclic theem), nyob rau hauv uas lub sijhawm ntawm tiam tuaj yeem ncav cuag ib hlis lossis ntau dua (lig Lub Tsib Hlis - Lub Xya Hli pib). Feem ntau nyob rau lub sijhawm no, kab mob nplooj yog qhov nyuaj los nrhiav tau txawm tias muaj kev soj ntsuam tshwj xeeb.
Theem ntawm exponential neeg loj hlob (theem polycyclic)
Ntau qhov kev soj ntsuam (Pshedetskaya, Kozubova, 1969; Borisenok, 1969; Osh, 1969; Dyakov, Suprun, 1984; Rybakova, Dyakov, 1990) tau qhia tias thaum pib ntawm epiphytoty, qis-virulent thiab ib qho me me zuj zus tuaj, uas tom qab hloov los ntawm ntau dua virulent thiab txhoj puab heev. tus nqi ntawm kev loj hlob ntawm cov nruj ntawm cov pej xeem yog qhov siab dua, tsis kam tiv thaiv ntau yam ntawm cov tswv tsev cog.
Raws li cov neeg loj zuj zus, qhov tseem ceeb ntawm ob qho ntawm ib qho tseem ceeb ntawm cov noob tau qhia rau hauv kev lag luam ntau yam (R1-R4) thiab xaiv tsis nruab nrab (R5-R11) nce. Yog li, nyob rau hauv cov pejxeem nyob ze Moscow hauv 1993, qhov nruab nrab virulence los ntawm lig Lub Xya Hli txog rau Lub Yim Hli-Yim Hli tau nce los ntawm 8,2 rau 9,4, thiab qhov loj tshaj tau pom rau qhov xaiv tsis muaj virulence gene R5 (los ntawm 31 txog 86% ntawm cov virulent clones) (Smirnov, 1996) )).
Ib qho kev txo qis ntawm kev loj hlob ntawm ib haiv neeg yog nrog los ntawm qhov kev txo qis hauv kev ua si ntawm pawg neeg parasitic. Yog li ntawd, nyob rau lub xyoo muaj kev ntxhov siab, ob qho tag nrho cov neeg sib tw thiab qhov sib npaug ntawm cov kev sib tw sib tw muaj tsawg dua nyob hauv epiphytotic (Borisenok, 1969). Yog tias ntawm qhov siab ntawm huab cua epiphytotic pauv hloov mus rau qhov tsis txaus siab rau lub ntsej muag blight thiab qos yaj ywm txo qis, qhov kev ua kom muaj zog ntawm cov virulent thiab txhoj puab heev tau txo qis (Rybakova li al., 1987).
Qhov nce ntawm cov zaus ntawm cov noob muaj feem cuam tshuam rau virulence thiab nruj ntawm cov pejxeem yuav yog vim muaj kev xaiv ntau virulent thiab txhoj puab heev hauv cov pejxeem sib xyaw. Txhawm rau qhia kev xaiv, ib txoj hauv kev rau kev tsom xam ntawm kev tsis sib hloov tau tsim, uas tau ua tiav hauv chemostat cov neeg muaj poov xab (Adams li al., 1985) thiab Fusarium graminearum (Wiebe li al., 1995).
Kev hloov pauv ntawm cov tsis tuaj yeem tiv thaiv blasticidin S hauv cov neeg thaj teb ntawm P. infestans tau txo qis thaum uas tig mus nrog kev loj hlob ntawm kev quab yuam ntawm cov pejxeem, uas qhia qhov hloov pauv ntawm cov pob zeb tseem ceeb thaum lub sijhawm kev loj hlob ntawm cov pejxeem (Rybakova li al., 1987).
Lub caij ntuj no nyob rau theem nyob rau hauv tubers
Thaum lub caij ntuj no hauv cov qos yaj ywm, virulence thiab nruj ntawm P. cov kab mob tsawg dua, thiab qhov virulence tsawg dua qhov tshwm sim qeeb dua li kev quab yuam (Rybakova thiab Dyakov, 1990). Pom zoo, nyob rau hauv cov xwm txheej uas ua tau zoo rau txoj kev loj hlob sai ntawm cov pejxeem loj (r-xaiv), "ntxiv" virulence noob thiab kev txhoj puab heev yog qhov tseem ceeb, yog li kev txhim kho ntawm epiphytotics yog nrog xaiv cov feem ntau virulent thiab nruj clones. Nyob rau hauv qhov xwm txheej ntawm qhov kev tsis txaus siab ntawm ib puag ncig, thaum tsis yog tus nqi luam, tab sis qhov kev mob siab ntawm qhov muaj qhov tsis zoo (K-xaiv) ua lub luag haujlwm tseem ceeb, "ntxiv" noob ntawm virulence thiab kev txhoj puab heev txo kev tawm dag zog, thiab clones nrog cov noob no yog thawj qhov kev tuag tawm, yog li qhov nruab nrab quab yuam thiab virulence ntawm cov neeg yog poob.
Tej theem nroj tsuag hauv av
Qeb qeb no yog kev paub tsis meej tshaj plaws hauv kev ua neej (Andrivon, 1995). Nws cov kev ua neej tau tshaj tawm txoj kev xav - vim tsis muaj cov ntaub ntawv hais txog dab tsi tshwm sim rau lub pathogen dhau ib lub sijhawm ntev (qee zaum ntau tshaj li ib hlis) - los ntawm kev tshwm sim ntawm cov noob qos rau qhov pom ntawm thawj cov pob ntawm tus kab mob rau lawv. Raws kev soj ntsuam thiab kev sim, tus cwj pwm ntawm cov pwm hauv lub sijhawm no ntawm lub neej tau rov kho dua tshiab (Hirst thiab Stedman, 1960; Boguslavskaya, Filippov, 1976).
Sporulation ntawm fungus tuaj yeem ua rau ntawm cov kab mob muaj kab mob hauv cov av. Lub txiaj ntsig spores nthuav dav nrog hyphae, uas tuaj yeem ua khoom noj rau lub sijhawm ntev hauv av. Lub Koom Haum Me Nyuam Yaus (tsim ntawm cov tub ntxhais) thiab cov theem nrab (ntawm mycelium hauv av) cov noob kab mob sawv mus rau cov av saum npoo av los ntawm capillary currents, tab sis tau txais lub peev xwm kis tau cov qos yaj ywm tsuas yog tom qab nws cov nplooj qis qis qis thiab nkag mus rau hauv cov av saum npoo av. Xws li nplooj (namely, thawj cov pob ntawm tus kab mob pom ntawm lawv) tsis ua rau tam sim ntawd, tab sis tom qab lub caij nyoog loj hlob thiab kev loj hlob ntawm cov qos saum.
Yog li, hauv lub neej ua haujlwm ntawm P. infestans, cov zaub cog qoob loo ntawm theem tuaj yeem tseem muaj. Yog hais tias nyob rau hauv cov theem ntawm cov kab mob ntawm lub neej mus ncig ua phem yog qhov tseem ceeb tshaj plaws ntawm kev ua kom muaj zog, tom qab ntawd hauv kev xaiv saprotrophic theem yog txo kev txo cov kab mob parasitic, raws li kev sim qhia rau qee cov kab mob phytopathogenic (saib Carson, 1993). Yog li no, nyob rau theem ntawm lub voj voog no, cov khoom siv txhoj puab heev yuav tsum ploj ntau qhov tseeb. Tab sis txog tam sim no tseem tsis tau muaj kev sim ncaj qha mus rau kev pom zoo cov kev xav saum toj no.
Cov caij nyoog hloov pauv tsis cuam tshuam rau cov kab mob tseem ceeb los ntawm P. infestans, tab sis kuj tiv thaiv cov kab mob fungicides, uas loj hlob nyob hauv theem polycyclic (thaum muaj mob epiphytoties), thiab poob qis thaum lub caij ntuj no cia (Derevyagina li al., 1991; Kadish thiab Cohen, 1992). Kev poob siab tshwj xeeb hauv kev tawm tsam cov hlau quav yeeb tau pom nyob nruab nrab ntawm kev cog ntoo ntawm cov cuam tshuam thiab cov tsos ntawm thawj cov pob ntawm tus kab mob hauv lub tshav pob.
Intraspecific tshwj xeeb thiab nws cov kev hloov pauv
P. Cov kab mob ua rau muaj kabmob sib kis hauv ob qho qoob loo tseem ceeb, qos yaj ywm thiab lws suav. Epiphytoties rau ntawm cov qos yaj ywm tau pib sai sai tom qab lub fungus nkag mus rau thaj chaw tshiab. Lub yeej ntawm cov txiv lws suav kuj tau sau tseg sai sai tom qab qhov tsos pom ntawm kev kis tus kab mob rau cov qos yaj ywm, tab sis epiphytoties ntawm lws suav tau sau tseg tsuas yog ib puas xyoo tom qab - hauv nruab nrab ntawm XNUMX caug xyoo. Ntawm no yog qhov Hallegli thiab Niederhauser sau txog qhov yeej txiv lws suav hauv tebchaws Asmeskas
(1962): “Kwv yees li 100 xyoo tom qab muaj mob loj heev thaum xyoo 1845, muaj tsawg tsawg lossis yuav luag tsis muaj peev xwm ua tau kom muaj ntau hom txiv lws suav. Txawm hais tias lig blight thawj zaug tau sau tseg rau cov txiv lws suav thaum ntxov li xyoo 1848, nws tsis yog qhov khoom ntawm lub siab ntawm cov neeg yug tsiaj ntawm cov nroj tsuag no kom txog thaum muaj kev sib kis muaj zog ntawm tus kabmob xyoo 1946. Nyob rau qhov chaw ntawm teb chaws Russia lig blight ntawm lws suav tau sau npe nyob rau hauv 60th caug xyoo. "Tau ntev, cov kws tshawb nrhiav tsis them nyiaj rau tus kab mob no, vim nws tsis ua rau muaj kev puas tsuaj rau kev lag luam. Tab sis xyoo 70 thiab 1979s. Xyoo pua puv XNUMX xyoo txog lub caij soob cov txiv lws suav muaj cai pom nyob rau hauv Soviet Union, feem ntau nyob hauv thaj av Lower Volga, Ukraine, North Caucasus, Moldova ... ”(Balashova, XNUMX).
Txij thaum ntawd los xij, lws suav blight los ntawm lig blight tau dhau los ua ib xyoos ib zaug, tau kis thoob plaws txhua qhov chaw ntawm kev lag luam thiab vaj tsev qoob loo thiab ua rau muaj kev lag luam muaj kev puas tsuaj rau cov qoob loo no. Dab tsi tshwm sim? Vim li cas thiaj li pom thawj zaug ntawm cov cab rau ntawm qos yaj ywm thiab mob epiphytotic ntawm cov qoob loo no tshwm sim yuav luag tib lub sijhawm, thiab yog vim li cas nws siv sijhawm li xyoo pua rau epiphytotic tshwm rau cov txiv lws suav? Cov kev sib txawv no pab txhawb tus neeg Mev es tsis yog Neeg Asmeskas Qab Teb qhov muaj tus kab mob. Yog tias hom tsiaj Phytophthora infestans tsim los ua cov cab ntawm Mexican tuber-bearing hom ntawm cov kua roj Solanum, tom qab ntawd nws nkag siab tias vim li cas cog qos yaj ywm koom nrog tib ntu ntawm cov genus vim tias hom neeg Mev tau cuam tshuam ntau heev, tab sis vim yog qhov tsis muaj kev sib koom tes nrog cov cab, uas tsis txhim kho cov tswv yim ntawm kev tshwj xeeb thiab tsis tseem ceeb.
Lws suav nyob rau ntawm ntu sib txawv ntawm cov genus, hom kev sib pauv tau muaj qhov sib txawv ntawm cov hom tuberous, yog li ntawd, txawm hais tias lub txiv lws suav tsis nyob sab nraum cov zaub mov tshwj xeeb ntawm P. infestans, qhov kev siv ntau ntawm nws qhov kev puas tsuaj tsis txaus rau qhov kev lag luam muaj kev puas tsuaj loj.
Qhov tshwm sim ntawm epiphytoties rau ntawm lws suav yog vim muaj kev hloov caj ces ntawm cov cab loj, uas tau nce ntau qhov nws yoog (pathogenicity) thaum kab mob parasitized. Peb ntseeg tias daim foos tshiab tshwj xeeb rau kev ntsuas cov txiv lws suav yog T1 haiv neeg tau piav qhia los ntawm M. Gallegly, cuam tshuam ntau yam ntawm txiv lws suav (Liab Cherry, Ottawa), tiv taus T0 haiv neeg thoob plaws ntawm qos yaj ywm (Gallegly, 1952). Thaj, qhov hloov pauv (lossis ib hom kev hloov pauv) uas tau hloov T0 mus rau hauv kev sib tw T1 thiab ua rau muaj qhov tshwm sim ntawm cov pob zeb ua kom nruj rau lub yeej ntawm cov txiv lws suav. Raws li feem ntau tshwm sim, qhov nce ntawm pathogenicity rau ib tus tswv tsev tau nrog nws txo qis rau lwm tus, uas yog, thaum pib, tseem tsis tau ua tiav qhov kev paub tshwj xeeb tseem ceeb - rau cov qos yaj ywm (haiv neeg T0) thiab rau txiv lws suav (haiv neeg T1).
Dab tsi yog ua pov thawj rau qhov kev xav no?
- Muaj tshwm sim ntawm cov qos yaj ywm thiab txiv lws suav. Ntawm cov nplooj txiv lws suav, Kev sib tw T1 muaj ntau tshaj, thaum nyob ntawm qos nplooj nws yog qhov tsis tshua muaj. Raws li S.F.Bagirova thiab T.A. Oreshonkova (tsis raug tshaj tawm) hauv cheeb tsam Moscow xyoo 1991-1992, qhov tshwm sim ntawm T1 kev sib tw hauv cov nroj tsuag qos yog 0%, thiab nyob rau tsob txiv lws suav - 100%; hauv 1993-1995 - 33% thiab 90%, feem; xyoo 2001 - 0% thiab 67%. Cov ntaub ntawv zoo sib xws tau muaj nyob hauv tebchaws Israel (Cohen, 2002). Cov kev sim nrog kev kis tus kab mob qos nrog kev sib cais ntawm T1 haiv neeg thiab kev sib xyaw ntawm kev sib cais T0 thiab T1 qhia tau hais tias kev sib cais ntawm kev sib tw T1 tsis zoo rau hauv cov tub ntxhais thiab tau hloov los ntawm kev sib cais ntawm T0 haiv neeg (Dyakov li al., 1975; Rybakova, 1988).
2) Kev loj hlob ntawm T1 kev sib tw hauv cov nroj tsuag soob. Thawj tus kab mob ntawm cov nplooj soob yog nqa los ntawm kev sib cais ntawm T0 haiv neeg, uas ua tus thawj hauv qhov tsom xam ntawm kev kis tus kab mob hauv thawj cov pob tsim rau ntawm nplooj. Qhov no pom tseeb cov txheej txheem lees txais feem ntau ntawm cov kab mob sib kis: qhov loj ntawm kev kis tus kab mob los ntawm cov qos yaj ywm yog tsim los ntawm T0 haiv neeg, txawm li cas los xij, muaj tsawg tus T1 clones tau khaws cia hauv cov qos yaj ywm, ib zaug ntawm cov txiv lws suav, tshem tawm T0 haiv neeg thiab nthuav mus rau thaum kawg ntawm lub sijhawm epiphytotic. Nws tseem muaj peev xwm hais tias muaj lwm txoj hauv kev ntawm txiv lws suav kab mob kis nrog T1 haiv neeg, uas tsis muaj zog li cov qos qos thiab nplooj, tab sis tas li. Yog li no, qhov chaw no muaj qhov tsis muaj zog txog kev hloov caj ces ntawm cov neeg kis tau cov txiv lws suav, tab sis tom qab ntawd txiav txim siab qhov kev txuam nrog T1 haiv neeg (Rybakova, 1988; Dyakov li al., 1994).
3) Kev txhoj puab heev rau cov qos yaj ywm thiab txiv lws suav. Cov kab mob sib kis ntawm cov txiv lws suav thiab qos nplooj nrog cais tawm ntawm kev sib tw T0 thiab T1 pom tias yav dhau los ua nruj rau cov qos yaj ywm ntau dua rau cov txiv lws suav, thiab cov tom kawg tau nruj rau cov txiv lws suav ntau dua rau cov qos yaj ywm. Cov kev sib txawv no tau nthuav tawm hauv kev hloov chaw ntawm kev sib cais ntawm cov "tsis muaj" ntawm cov haiv neeg sib xyaw thaum cov nplooj ntoo nplooj ntoo hauv tsev cog khoom (D'yakov et al., 1975) thiab hauv thaj av (Leberton li al., 1999); qhov sib txawv ntawm qhov sib kis tsawg kawg nkaus, lub sijhawm latency, qhov loj me ntawm cov kis kab mob thiab spore ntau lawm (Rybakova, 1988; Dyakov li al., 1994; Legard et al., 1995; Forbes li al., 1997; Oyarzun li al., 1998; Leberton li al., 1999) al., 2000; Vega-Sanchez li al., 2002; Knapova, Gisi, 2004; Sussuna li al., XNUMX).
Cov kev ua phem ntawm cais ntawm T1 haiv neeg rau cov hom txiv lws suav uas tsis muaj cov noob tsis kam yog qhov muaj peev xwm zoo li ntawd cov cais tawm ntawm cov nplooj raws li cov khoom noj khoom haus nruab nrab tsis muaj necrotizing cov ntaub so ntswg mob (Dyakov li al., 1975; Vega-Sanchez li al., 2000).
4) Virulence rau cov qos yaj ywm thiab txiv lws suav. Kev sib tw T1 cuam tshuam rau cov txiv lws suav hom nrog cov Ph1 tsis kam, thaum lub T0 kev sib tw tsis muaj peev xwm kis cov hom no, i.e. muaj ib rab me paj paim thiab. Hauv kev hais txog qhov sib txawv
R-cov noob ntawm cov qos yaj ywm muaj feem cuam tshuam; cov kab mob sib cais los ntawm cov nplooj soob tsis muaj qhov npau taws dua li cov kab mob "qos" (Cov Lus 11).
5) Qhov nruab nrab kos xim. Kev soj ntsuam ntawm cov cim ntsuas ntawm qhov nruab nrab ntawm cov neeg ntawm P. infestans parasitizing rau qos yaj ywm thiab txiv lws suav kuj tseem ua pov thawj rau qhov kev xav tau ntau txoj kev xaiv. Hauv Brazilian cov neeg coob ntawm P. infestans, nplooj soob cais rau cov kab clonal US-1, thiab cov uas los ntawm nplooj qos yog los ntawm BR-1 kab (Suassuna li al., 2004). Hauv Florida (Tebchaws Asmeskas), txij li xyoo 1994, clone US-90 tau pib ua rau cov qos yaj ywm (nrog kev tshwm sim ntau dua 8%), thiab clones US-11 thiab US-17 ntawm cov txiv lws suav, thiab cov tom kawg cais tau nruj rau cov txiv lws suav ntau dua rau cov qos yaj ywm (Weingartner , Tombolato, 2004). Cov kev sib txawv tseem ceeb hauv cov keeb kwm keeb kwm (DNA cov ntiv tes) hauv cov qos yaj ywm thiab txiv lws suav sib cais tau tsim rau 1200 P. cov kab mob sib kis tau tsawg dua hauv Tebchaws Asmeskas txij xyoo 1989 txog xyoo 1995 (Deahl li al., 1995).
Siv tus txheej txheem AFLP ua rau nws tuaj yeem cais 74 hom kab sau los ntawm cov qos thiab lws suav nplooj hauv 1996-1997. hauv Fab Kis thiab Switzerland, nyob hauv 7 pawg. Cov qos thiab txiv lws suav pom tsis meej meej pom tsis meej, tab sis cov kab mob "qos" yog cov caj ces txawv dua li cov "lws suav". Cov qub tau pom nyob rau hauv tag nrho xya pawg, thiab tom kawg, tsuas yog hauv plaub, uas qhia qhov tshwj xeeb ntau ntawm cov caj dab tom kawg (Knapova and Gisi, 2002).
6) Cov txheej txheem ntawm kev rho tawm. Yog hais tias cov neeg ntawm tus cab no ntawm ob tus tswv tsev cov tsiaj hom hloov zuj zus mus rau txoj kev nqaim ntawm kev ua haujlwm tshwj xeeb rau lawv tus kheej "tus tswv tsev", tom qab ntawd ntau yam ua ntej- thiab postmeiotic cov kev tshwm sim uas tiv thaiv kev sib deev caj ces sib pauv (Dyakov thiab Lekomtseva, 1984).
Ntau cov kev tshawb fawb tau tshawb xyuas cov txiaj ntsig ntawm lub hauv paus ntawm niam txiv lim ntshav ntawm kev ua haujlwm ntawm kev ua kom zoo. Thaum hla cov hom kab sib cais los ntawm ntau cov sib txawv ntawm cov genus Solanum hauv Ecuador (Oliva et al., 2002), nws pom tias cov kab mob nrog mating hom A2 los ntawm Solanaceae qus (cov kab rov tav EC-2) yog qhov phem tshaj nrog cov kab mob los ntawm cov txiv lws suav (kab EC. -3), thiab feem ntau cuam tshuam zoo nrog cov kab mob qos (EC-1).
Txhua tus me nyuam tsiaj pom tias tsis yog-tus kabmob. Tus sau phau ntawv ntseeg tias qhov feem pua tsawg ntawm kev sib xyaw thiab kev txo qis ntawm cov kab mob tseem ceeb hauv kev ua ke yog vim muaj cov postmeiotic cov txheej txheem ntawm kev ua me nyuam ntawm cov neeg muaj mob.
Hauv kev sim ntawm Bagirova li al. (1998), coob tus qos thiab lws suav hom tau hla nrog cov khoom ntawm T0 thiab T1 kev sib tw. Cov muaj txiaj ntsig zoo tshaj plaws yog cov khaub lig ntawm T1xT1 hom raug rho tawm los ntawm soob (36 oospores hauv lub microscope teb ntawm kev saib, 44% ntawm oospore germination), qhov tsawg tshaj plaws yog crosses ntawm T0xT1 kev sib tw cais los ntawm cov sib txawv (tsawg tus naj npawb ntawm kev tsim kho thiab cov nroj tsuag oospores, qhov ntau ntawm kev tsim txom thiab kev tsim kho oospores) ... Qhov ua tau zoo ntawm cov ntoo khaub lig sib cais ntawm T0 kev sib cais ntawm cov qos yaj ywm tau muaj nruab nrab. Txij li lub cev tseem ceeb ntawm kev sib tw ntawm T0 cuam tshuam rau cov qos yaj ywm, nws muaj kev ntseeg tau txog lub caij ntuj no - cov qos yaj ywm, vim qhov tseem ceeb ntawm oospores raws li lub caij ntuj no kis rau cov neeg los ntawm cov qos yaj ywm muaj tsawg. Daim ntawv soob ua "txiv lws suav" muaj peev xwm lub caij ntuj no ntawm txiv lws suav hauv kev ua cov oospores (saib hauv qab) thiab yog li muaj qhov ua tau zoo ntawm kev sib deev. Vim tias nws qhov muaj peev xwm ntau, T1 kis tau ib lub peev xwm ywj pheej rau thawj tus kab mob hauv cov txiv lws suav. Cov txiaj ntsig tau los ntawm Knapova li al. (Knapova li al., 2002) tuaj yeem txhais tib yam nkaus. Hla cov kab mob sib cais los ntawm cov qos yaj ywm nrog lim los ntawm txiv lws suav tau muab ntau tshaj plaws ntawm oospores - 13,8 ib ntu mm. nruab nrab (nrog kev sib kis ntawm 5-19) thiab ib nrab feem pua ntawm kev kis tus neeg ntawm oospores (6,3 nrog kev sib kis ntawm 0-24). Cov kab hla kev sib txawv ntawm cov txiv lws suav tau muab qhov feem pua qis ntawm oospores (7,6 nrog kev sib kis ntawm 4-12) nrog qhov feem pua siab tshaj plaws ntawm lawv cov noob (10,8). Cov kab sib hla ntawm txoj hlua sib cais ntawm cov qos yaj ywm tau muab cov xov tooj ntawm nruab nrab oospores (8,6 nrog cov ntaub ntawm qhov sib txawv - 0-30) thiab qhov feem pua qis tshaj ntawm kev tawm ntawm oospores (2,7). Yog li, lim los ntawm cov qos yaj ywm muaj zog tsawg dua li cov los ntawm cov txiv lws suav, tab sis cov kws txhais lus ntu tsis muab cov txiaj ntsig tsis zoo dua li cov kev mob siab. Nws yog tau hais tias qhov sib txawv nrog cov ntaub ntawv saum toj no los ntawm Bagirova li al. tau piav qhia los ntawm qhov tseeb hais tias cov kws tshawb nrhiav Lavxias tau ua haujlwm nrog cov kab sib txawv cais nyob rau thaum ntxov 90s ntawm lub xyoo pua nees nkaum, thiab cov kws tshawb nrhiav Swiss - nrog cov kab mob cais tawm nyob rau hauv lub lig 90s.
Lub hauv paus rau kev qes qis tej zaum yuav yog qhov heteroploidy ntawm tus leeg. Yog tias nyob hauv cov neeg hauv tebchaws Mexico, qhov kev sib deev thiab thawj kis nrog oospore progeny yog ib txwm muaj, feem ntau ntawm cov kab kev kawm ntawm P. Infestans yog diploid, tom qab ntawd hauv cov teb chaws ntawm Ntiaj Teb Qub intrapopulation polymorphism ntawm ploidy tau pom (di-, tri- thiab kab mob tetraploid, nrog rau heterokaryotic leeg nrog heteroploid nuclei) , thiab cov leeg muaj ntau hom kev sib tham, i.e. ob leeg muaj peev xwm, muaj qhov sib txawv hauv nuclear ploidy (Therrien li al., 1989, 1990; Whittaker li al., 1992; Ritch, Daggett, 1995). Kev ua kom sib txawv ntawm nuclei hauv antheridia thiab oogonia tuaj yeem yog qhov laj thawj ntawm kev yug qis.
Raws li rau nuclear kev sib pauv ntawm hyphae thaum anastomoses, qhov no tiv thaiv los ntawm kev noj haus tsis sib xws, uas tawg asexual coob rau ntau cov caj ces raug rho tawm (Poedinok thiab Dyakov, 1987; Gorbunova li al., 1989; Anikina li al., 1997b).
7) Kev sib txuam ntawm cov neeg. Cov ntaub ntawv saum toj no qhia tau hais tias kev sib xyaw ntawm "qos" thiab "soob" P. infestans leeg tau muaj. Reciprocal rov kis tus kab mob ntawm cov tswv tsev sib txawv kuj ua tau, txawm hais tias txo qis kev txhoj puab heev.
Kev tshawb ntawm cov pej xeem cov cim hauv cov kev sib cais los ntawm cov kab qos thiab txiv lws suav uas nyob ib sab hauv xyoo 1993 qhia tau hais tias kwv yees li ib feem peb ntawm cov cais tawm ntawm cov txiv lws suav nplooj tau pauv los ntawm cov teb cog qoob loo nyob ze (Dolgova li al., 1997). Raws li txoj kev xav, nws yuav xav tias kev sib txawv ntawm cov neeg ntawm ob lub tswv yim yuav nce thiab coj mus rau qhov tshwm sim ntawm cov ntaub ntawv tshwj xeeb (f.sp. qos thiab f.sp. soob), tshwj xeeb tshaj yog txij li oospores tuaj yeem mob siab rau cov nroj tsuag khib nyiab (Drenth et al., 1995 ; Bagirova, Dyakov, 1998) thiab cov noob txiv lws suav (Rubin et al., 2001). Thiaj li, cov txiv lws suav tam sim no muaj cov txiaj ntsig ntawm lub caij nplooj ntoo hlav tshiab ywj siab ntawm qos tubers.
Txawm li cas los xij, txhua yam tshwm sim txawv. Kev hla dhau nrog oospores tau tso cai rau cov cab ua kom tsis txhob muaj qhov nqaim theem tshaj plaws hauv nws lub neej kev ua neej - monocyclic theem ntawm cov nroj tsuag hauv av, thaum lub sijhawm cov cab no txo qis, uas yog maj mam rov qab los rau hauv polycyclic theem nyob rau lub caij ntuj sov.
Rooj ntawv 11. Feem ntau ntawm cov virulence noob rau cov qos txawv ntawm cov kab mob sib txawv hauv hom P. infestans hom
lub teb chaws | Xyoo | Qhov nruab nrab pes tsawg ntawm virulence noob hauv hom | sau | |
los ntawm cov qos yaj ywm | los ntawm txiv lws suav | |||
Fabkis | 1995 | 4.4 | 3.3 | Leberton li al., 1999 |
1996 | 4.8 | 3.6 | Leberton, Andrivon, 1998 | |
Fab Kis, Switzerland | 1996-97 | 6.8 | 2.9 | Knapova, Gisi, 2002 |
United States | 1989-94 | 5 | 4.8 | Goodwin li al., 1995 |
USA, Zap. Washington | 1996 | 4.6 | 5 | Dorrance li al., 1999 |
1997 | 6.3 | 3.5 | " | |
Ecuador | 1993-95 | 7.1 | 1.3 | Oyarzun li al., 1998 |
Ixayees | 1998 | 7 | 4.8 | Cohen, 2002 |
1999 | 6 | 5.7 | " | |
2000 | 6.7 | 6.1 | " | |
Russia, Mosk. cheeb tsam | 1993 | 8.9 | 6.7 | Smirnov, 1996 |
Russia, thaj chaw sib txawv | 1995 | 9.4 | 8 | Kozlovskaya thiab lwm tus. |
1997 | 9.2 | 9.2 | " | |
2000 | 8.7 | 4.8 | " |
Thawj zoosporangia thiab zoospores, uas tsim tawm oospores, muaj qhov siab ntawm kev ua haujlwm parasitic, tshwj xeeb tshaj yog yog hais tias oospores tau tsim parthenogenetically nyob rau hauv kev cuam tshuam ntawm pheromones ntawm ib qho kev cuam tshuam nrog hom sib txawv ntawm kev sib raug zoo. Yog li, cov khoom kis tau ntawm cov txiv lws suav loj hlob los ntawm cov noob kis tau tus mob oospores yog cov kab mob heev rau ob lub txiv lws suav thiab qos.
Cov kev hloov pauv no ua rau lwm tus pejxeem rov txhim kho dua, qhia raws li cov kev hloov pauv tseem ceeb hauv qab ntu kis los ntawm kev pom:
- Raug kis cov txiv lws suav tau dhau los ua qhov tseem ceeb ntawm thawj qhov kev kis kab mob ntawm cov qos yaj ywm (Filippov, Ivanyuk, cov lus ntawm tus kheej).
- Epiphytoties ntawm cov qos yaj ywm tau pib pom thaum ntxov txij Lub Rau Hli, txog li ib hlis ua ntej tshaj li niaj zaus.
- Hauv cov nroj tsuag qos, qhov feem pua ntawm kev sib tw T1 nce, uas tau pom yav dhau los muaj qhov tsis txaus (Ulanova et al., 2003).
- Hom mob cais los ntawm cov nplooj soob tsis txawv ntawm kab mob ntawm cov qos rau virulence ntawm qos txawv ntawm virulence noob thiab pib dhau los "hom" kab mob hauv kev ua phem tsis yog rau cov txiv lws suav, tab sis kuj rau cov qos yaj ywm (Lavrova et al., 2003; Ulanova li al. , 2003).
Yog li, hloov ntawm divergence, muaj kev sib txuam ntawm cov neeg, qhov tshwm sim ntawm ib tus tib neeg ntawm ob tus tswv tsev cog nrog qhov siab thiab nruj rau ob hom.
xaus
Yog li, txawm tias ntau tshaj li 150 xyoo ntawm kev kawm siab ntawm P. infestans, hauv kev kawm txog tsiaj txhu, suav nrog cov pejxeem ntawm cov kab mob no ua rau muaj tus kabmob tseem ceeb tshaj plaws ntawm kev cog qoob loo solanaceous, ntau yam tseem tsis tau paub. Nws tsis paub meej tias qhov kev tso cai ntawm qee tus neeg ntawm lub neej mus los li cas, cuam tshuam dab tsi ntawm caj ces thiab caj dab, qhov sib piv ntawm kev muaj me nyuam thiab cov kab mob sib txawv ntawm cov kev ua me nyuam ntawm cov pej xeem, ua li cas txoj kev noj tsis taus txoj kev tau txais los ntawm caj ces, dab tsi yog lub luag haujlwm ntawm cov qos yaj ywm thiab cov txiv lws dab tsi cuam tshuam rau cov pej xeem cov qauv ntawm cov cab. Txog tam sim no, cov teeb meem tseem ceeb ntawm cov tswv yim xws li cov kev tshuaj ntsuam caj ceg rau kev hloov pauv kev txhoj puab ntawm cov cab los yog yaig ntawm nonspecific qos tsis kam tsis tau daws. Nrog kev sib sib zog nqus thiab nthuav dav ntawm kev tshawb fawb txog cov qos lig blight, cov cab no tawm tsam cov kev cov nyom tshiab rau cov kws tshawb nrhiav. Txawm li cas los xij, kev txhim kho ntawm kev muaj peev xwm sim, qhov tshwm sim ntawm txoj kev tshiab ntawm kev tawm tswv yim tshiab rau kev tswj hwm nrog cov noob thiab cov protein ua rau peb cia siab rau qhov kev vam meej ntawm cov lus nug.
Cov ntsiab lus tau luam tawm nyob rau hauv phau ntawv journal "Kev Tiv Thaiv Qos Yaj Zwm" (No. 3, 2017)