S.N. Elansky, L. Yu. Kokaeva, N.V. Statsyuk, Yu.T. Anayankha Zamgululi
Mau oyamba
Oomycete Phytophthora infestans (Mont.) De Bary, yemwe amayambitsa matenda oopsa mochedwa, matenda ofunikira kwambiri a mbatata ndi tomato, adakopa chidwi cha ofufuza ochokera kumayiko osiyanasiyana kwazaka zopitilira zana limodzi ndi theka. Mwadzidzidzi kuwonekera ku Europe pakati pa zaka za XNUMXth, kudadzetsa mliri wa mbatata womwe wakumbukirabe m'mibadwo yambiri.
Mpaka pano, amatchedwa "bowa wanjala yaku Ireland". Pafupifupi zaka zana limodzi kuchokera pomwe miliri yoyamba idapezeka, mitundu ya mbatata yaku Mexico yolimbana ndi vuto lakumapeto idapezeka, njira zodutsira ndi mbatata zomwe zidalimidwa zidapangidwa (Muller, 1935), ndipo mitundu yoyamba yolimbana ndi ziwopsezo idapezeka (Pushkarev, 1937). Komabe, atangoyamba kumene kulima malonda, mafuko a tizilombo toyambitsa matenda omwe anali oopsa amtundu wosagwirizana anasonkhana. ndipo kuyambitsidwa kwa majini atsopanowa kuchokera ku mbatata zakuthengo zaku Mexico kukhala mitundu idayamba kuchepa mphamvu.
Kulephera kugwiritsa ntchito monogenic (ofukula) kukana kukakamiza obereketsa kuti ayang'ane njira zovuta kugwiritsa ntchito nonspecific polygenic (yopingasa) kukana. M'zaka zaposachedwa, mafuko aukali kwambiri ayamba kudziunjikira mwa anthu amodzi a tiziromboti, ndikupangitsa kukokoloka kwa ngakhale kukana kwapadera. Kubwera kwa tizilombo toyambitsa matenda kwadzetsa mavuto pakugwiritsa ntchito mankhwala oteteza mbatata.
Chifukwa cha kusiyana kwakukulu pakati pa oomycetes ndi bowa mumankhwala, ultrastructure ndi metabolism, fungicides, makamaka systemic, yomwe imagwiritsidwa ntchito kuteteza zomera ku matenda ambiri am'fungulo, siyothandiza ku oomycetes.
Chifukwa chake, poteteza mankhwala ku vuto lakumapeto, zingapo (mpaka maulendo 12 pa nyengo kapena kupitilira apo) kupopera mbewu mankhwalawa ndi kukonzekera kukhudzana ndi zochitika zosiyanasiyana. Gawo losintha linali kugwiritsa ntchito phenylamides, yomwe ndi poizoni wa oomycetes ndikufalikira mwadongosolo muzomera. Komabe, kugwiritsidwa ntchito kwawo mwachangu kunadzetsa kuchuluka kwa mitundu yolimbana ndi mafangasi (Davidse et al., 1981), zomwe zidasokoneza chitetezo cha mbewu. P. infestans ndiye kachilombo kokha kamene kali m'dera lotentha, zoyipa zomwe kulima organic sizingathetsedwe popanda kugwiritsa ntchito njira zodzitetezera (Van Bruggen, 1995).
Zomwe tafotokozazi zikufotokozera chidwi chomwe ofufuza ochokera kumayiko osiyanasiyana amapereka pophunzira za P. infestans kuchuluka, mphamvu zakukula kwawo ndi kapangidwe ka majini, komanso njira zosinthira majini.
Nthawi yamoyo wa R. INFESTANS
Oomycete Phytophthora infestans amapanga tinthu tating'onoting'ono ta mycelium tokhala ndi haustoria mkati mwa masamba a mbatata. Kudya masamba a tsamba, kumayambitsa mapangidwe amdima, omwe amatembenuka wakuda ndikuwola nyengo yamvula. Ndi kugonjetsedwa kwamphamvu, tsamba lonse limamwalira. Pakadutsa nthawi yodyetsa, zotuluka zimapangidwa pa mycelium - sporangiophores - zomwe zimakula panja kudzera mu stomata. M'nyengo yonyowa, amapanga pachimake choyera mozungulira madera omwe ali pansi pamasamba. Kumapeto kwa sporangiophores, zoosporangia zooneka ngati mandimu zimapangidwa, zomwe zimatuluka ndikunyamulidwa ndi mvula (mkuyu 1). Kugwera m'madontho amadzi pamwamba pa tsamba la mbatata, sporangia imera ndi malo osungira a 6-8, omwe, atayenda kwakanthawi, amakhala ozunguliridwa, okutidwa ndi chipolopolo ndikumera ndi chubu cha mphukira. Mphukira imalowa kudzera mu stomata mpaka minofu ya masamba. Nthawi zina, sporangia amatha kukula mu chubu chokula mwachindunji m'masamba a masamba. Pazifukwa zabwino, nthawi yoti matenda apatsidwe mpaka kupangidwa kwatsopano ndi masiku 3-4 okha.
Akakhala pansi ndikusefera panthaka, sporangia amatha kupatsira ma tubers. Matenda a tubers okhudzidwa kwambiri amawola panthawi yosungirako; mwa omwe akhudzidwa pang'ono, kachilomboka kangapitirirebe mpaka nyengo yotsatira. Kuphatikiza apo, wothandizila wa caucative wa vuto lakumapeto amatha kupitilira nthawi yozizira ngati ma oospores (mipanda yolimba yopumulira zogonana) m'nthaka pazinyalala zambewu ndi mbewu za phwetekere. Oospores amapangidwa pazomera zamoyo pomwe mitundu yamitundu yosiyanasiyana ikumana ndi chinyezi chochuluka. Mu kasupe, kuphulika kwa asexual kumapangidwa pa tubers yomwe ili ndi kachilomboka komanso pa zotsalira zazomera zokhala ndi ma oospores; zoospores zimalowa m'nthaka ndikuyambitsa matenda am'munsi mwa masamba. Nthawi zina, mycelium imatha kukula kuchokera ku tuber yomwe ili ndi kachilomboka pambali yobiriwira ya mbewuyo ndipo nthawi zambiri imawonekera kumtunda kwa tsinde.
Kusiyanitsa kwakukulu pakati pa oomycetes ndi mafangayi ambiri kumakhala pakukula kwa diplophase m'moyo wawo ndimasewera a meiosis ndikumera kwa zygotes (oospores) popanda kutulutsa nyukiliya kochepetsera. Izi, kuphatikiza ma dipolar heterotallism m'malo mwa amuna kapena akazi okhaokha, zikuwoneka kuti zingathandize kugwiritsa ntchito oomycetes njira zophunzirira kuchuluka kwa ma eukaryote apamwamba (kusanthula kwa panmixia ndi kugawa magawo a anthu, kuthamanga kwa intra- ndi kulowerera, ndi zina zambiri). Komabe, pali zinthu zitatu zomwe sizimalola kusamutsiratu njirazi pophunzira za P. infestans anthu.
1. Pamodzi ndi oospores a haibridi, oospores yodzipangira yokha komanso yopanga gawo limodzi imapangidwa mwa anthu (Fife ndi Shaw, 1992; Anikina et al., 1997a; Savenkova, Cherepnikoba-Anirina, 2002; Smirnov, 2003), komanso kuchuluka kwakapangidwe kake kokwanira kukhala kokwanira pa zotsatira za mayeso.
2. Kugonana kwa P. infestans kumathandizira kwambiri pakukula kwa kuchuluka kwa anthu, chifukwa mafangayi amabereka makamaka ndi masamba, omwe amapanga zoposa 90% yazotsatira zakusanthula kwa mtundu wa mating pogwiritsa ntchito njira yachikhalidwe pa chopatsa thanzi ... nyengo yokula ndimibadwo ingapo ya asexual sporulation (chitukuko cha matenda a polycyclic). Ziwombankhanga zimathandiza kwambiri kuteteza thupi m'nyengo yomwe zomera zobiriwira kulibe (m'nyengo yozizira) komanso mu matenda oyamba a mbande. Kenako, nthawi yachilimwe, kubala kwama clonal kumachitika ndikuwonjezeka kapena, kapena, kuchepa kwa kuchuluka kwamatombo omwe amabwera chifukwa chakukumbutsanso zachiwerewere, komwe kumatsimikizika makamaka pakusankha kosinthidwa kwambiri. Chifukwa chake, kuchuluka kwa matanthwe amodzi mwa anthu koyambirira ndi kumapeto kwa epiphytotic atha kukhala osiyana kotheratu.
3. Makombedwe omwe anafotokozedwawa ndi omwe amapezeka kwa anthu wamba a P. infestans kwawo, Central America. M'madera ena adziko lapansi, machitidwe azakugonana sanadziwike kwazaka zopitilira 100; chotupa cha mycelium m'matumba a mbatata omwe anali ndi kachilombo anali nthawi yachisanu. Kuzungulira kwa moyo kunali kovuta kwambiri, ndipo kufalikira kunali koyang'ana m'chilengedwe: matenda ochokera ku ma tubers omwe ali ndi kachilombo kamodzi adadutsa m'masamba, ndikupanga mawonekedwe oyambilira a matendawa, omwe amatha kuphatikizika pakukula kwakukulu kwa matendawa.
Chifukwa chake, mmadera ena pakhoza kukhala kusinthana kwamachitidwe azakugonana ndi asexual, pomwe ena - azisangalalo zokha.
Chiyambi cha P. INFESTANS
P. infestans adapezeka ku Europe kumapeto kwa theka loyambirira la 1991th century. Popeza mbatata imapezeka kumpoto chakum'mawa kwa South America, zimaganiziridwa kuti tizilomboti tidachokera kumeneko kupita ku Europe panthawi ya mchere waku Chile. Komabe, maphunziro omwe adachitika ku Rockefeller Center station ya mbatata ku Toluca Valley, Mexico adakakamiza kuti malingalirowa asinthidwe (Niederhauser 1993, XNUMX).
1. M'chigwa cha Toluca, mitundu ya mbatata ya m'deralo (Solanum demissum, S. bulbocastanum, ndi zina) imakhala ndi mitundu yosiyanasiyana ya majini osagwirizana motsutsana ndi milingo yayikulu, yomwe imasonyeza kusinthika kwakanthawi ndi tiziromboti. Mitundu yaku South America, kuphatikiza mbatata za mbewu, ilibe majini otsutsana.
2. M'chigwa cha Toluca, mumadzipatula ndi mitundu ya A1 ndi A2, chifukwa chake kuchuluka kwa P. infestans kuli ponseponse; ali kudziko lakwawo kwa mbatata zolimidwa, South America, tiziromboti timafalikira mwachilengedwe.
3. M'chigwa cha Toluca, mumakhala miliri yoopsa yapachaka yoopsa mochedwa. Chifukwa chake, pakati pa ofufuza aku North America (University of Cornell), malingaliro onena za Mesoamerica (Central America) monga malo obadwira mbatata phytophthora akhazikitsidwa (Goodwin et al., 1994).
Ofufuza aku South America sagwirizana ndi izi. Amakhulupirira kuti mbatata yolimidwa ndi tizilombo toyambitsa matenda a P. infestans ali ndi kwawo komweko - South America Andes. Anathandizira malingaliro awo mwa kafukufuku wamagulu pofufuza ma DNA polymorphisms a mitochondrial genome (mtDNA) ndi majini anyukiliya a RAS ndi β-tubulin (Gomez-Alpizar et al., 2007). Adawonetsa kuti zovuta zomwe zimasonkhanitsidwa kuchokera kumadera osiyanasiyana padziko lapansi zimachokera ku mizere itatu yamakolo, yomwe (onse atatu) amapezeka ku South America Andes. Andean haplotypes ndi mbadwa za mizere iwiri: olekanitsidwa ndi mzere wakale kwambiri wa mtDNA amapezeka pa zakutchire Solanaceae kuchokera pagawo la Anarrhicomenum ku Ecuador, pomwe gawo lachiwiri limapezeka pa mbatata, tomato, ndi nightshades zakutchire. Ku Toluca, ngakhale ma haplotypes osowa kwenikweni amachokera pamzera umodzi wokha, ndikusintha kwamitundu yamavuto a Toluca (kuchepa kwamalo amalo ena osinthika) akuwonetsa kuyambitsa kwamphamvu chifukwa chakungoyenda kwaposachedwa.
Kuphatikiza apo, mtundu watsopano P. andina udapezeka ku Andes, morphologically and genetetic similar to P. infestans, omwe, malinga ndi olemba, amaloza ku Andes ngati malo otentha kwambiri mu mtundu wa Phytophthora. Pomaliza, ku Europe ndi United States, P. Infestans amakhala ndi mizere yonse ya Andes, pomwe ku Toluca m'modzi yekha.
Bukuli lidalimbikitsa kuyankha kuchokera pagulu la ofufuza ochokera kumayiko osiyanasiyana omwe adachita ntchito zambiri zoyesera kuti abwereze kafukufuku wakale (Goss et al., 2014). Pogwira ntchitoyi, choyamba, ma DNA opangidwa ndi microsatellite ambiri adagwiritsidwa ntchito kuphunzira ma polymorphisms a DNA; kachiwiri, pakuwunika masango, njira zosamukira, nthawi yosiyana kwa anthu, ndi zina zambiri. Mitundu yapamwamba kwambiri idagwiritsidwa ntchito (ziwerengero za F, ziwerengero za ku Bayesi, ndi zina zambiri) ndipo, chachitatu, kuyerekezera sikunangogwiritsidwa ntchito ndi mitundu ya Andesan P. andina, momwe mtundu wosakanizidwa unakhazikitsidwa (P. infestans x Phytophthora sp.) . Chifukwa cha kusanthula uku, zidawonetsedwa mosapita m'mbali kuti gawo loyambira la mtengo wamtundu wa mitundu yonse yamtundu wa Phytophthora lomwe lidatengedwa mu kafukufukuyu, kupatula mtundu wosakanizidwa wa P. andina, ndi amtundu wa Mexico, ndipo mayendedwe osunthira ali ndi malangizo Mexico - Andes, osati mosemphanitsa, ndipo chiyambi chake chimagwirizana ndi aku Europe chikoloni cha New World (zaka 2012-300 zapitazo). Chifukwa chake, kutuluka kwa mitundu ya P. infestans, yodziwika bwino pakugonjetsedwa kwa mbatata, kunachitika mu chipinda chachiwiri chachilengedwe chokhazikitsa tuberous solanaceae, i.e. ku Central America.
Chibadwa cha P. INFESTANS
Mu 2009, gulu lapadziko lonse la asayansi linatsata mtundu wonse wa P infestans genome (Haas et al, 2009), kukula kwake kunali 240 MB. Izi ndizowirikiza kangapo kuposa mitundu yofanana kwambiri ya P. sojae (95 Mb), yomwe imayambitsa mizu yovunda ya soya, ndi P. Ramorum (65 Mb), zomwe zimakhudza mitengo yamtengo wapatali monga thundu, beech ndi ina. Zomwe zapezeka zikuwonetsa kuti genome ili ndi mitundu yambiri yamakanema obwereza - 74%. Genome ili ndi mitundu 17797 yolembera mapuloteni, ambiri mwa iwo omwe ali majini omwe amakhudzidwa ndimachitidwe am'manja, kuphatikiza kubwereza kwa DNA, kusindikiza ndikumasulira mapuloteni.
Kuyerekeza kwa majini amtundu wa Phytophthora kuwulula bungwe lachilendo la genome, lomwe limapangidwa ndimitundu yazomwe zimasungidwa, momwe kuchuluka kwa majini kumakhala kochulukirapo, ndipo zomwe zimachitika mobwerezabwereza ndizotsika, ndi zigawo zina zomwe zimakhala ndi mitundu yosasamala ya majini, okhala ndi kuchuluka kwa majini ochepa komanso okhala ndi zigawo zambiri zobwereza. Ma Conservative block amapanga 70% (12440) mwa mitundu yonse ya P. infestans yolemba ma protein. Pakati pokhazikika, majini nthawi zambiri amakhala otalikirana ndi 604 bp. M'madera omwe amakhala pakati pa zotchinga, mtunda wa intergenic ndi wokulirapo (3700 bp) chifukwa cha kuchuluka kwa zinthu zobwereza. Makina obisalira mwachangu omwe amapezeka mwachangu amapezeka m'magawo osauka.
Kusanthula motsatira kwa P. Infestans genome kunawonetsa kuti pafupifupi gawo limodzi mwa magawo atatu a matupi athu ndi zinthu zosinthika. Matupi a P. infestans amakhala ndi mabanja osiyanasiyana mosiyanasiyana kuposa ma genome ena odziwika. Ambiri mwa ma infestans a P. amachokera kubanja lachi Gypsy.
Mu P. infestans genome, kuchuluka kwakukulu kwamabanja amtundu wa gene omwe amapezeka mu pathogenesis adadziwika. Gawo lalikulu laiwo limakhazikitsa mapuloteni othandizira omwe amasintha momwe thupi limagwirira ntchito ndikuthandizira pakukhudzidwa. Amagwera m'magulu awiri akulu: opoplastic effectors, omwe amakhala m'malo amkati mwa ma cell (apoplast), ndi ma cytoplasmicactor, omwe amalowa m'maselo kudzera pa haustoria. Zoyambitsa za apoplastic zimaphatikizapo ma enzyme obisika a hydrolytic monga ma protease, lipases ndi glycosylases omwe amawononga maselo am'mimba; zoletsa zoteteza ku michere yodzitchinjiriza; ndi poizoni wa necrotizing monga mapuloteni onga a Nep1 (NPLs) ndi ma Pcf onga mapuloteni ang'onoang'ono a cysteine (SCRs).
Mitundu ya P. infestans yogwiritsira ntchito ndi yochulukirapo ndipo nthawi zambiri imakhala yayikulu kuposa mitundu yopanda tizilombo. Odziwika bwino ndi ma cytoplasmic effectors RXLR ndi Crinkler (CNR). Zomwe zimayambitsa ma cytoplasmic a oomycetes ndi mapuloteni a RXLR. Mitundu yonse yamtundu wa RXLR yomwe yapezeka pakadali pano ili ndi gulu la amino-terminal Arg-XLeu-Arg, pomwe X ndi amino acid. Zotsatira za kafukufukuyu, akuti akuti pali mitundu 563 ya RXLR mu P. infestans genome, yomwe ndi 60% kuposa ya P. sojae ndi P. ramorum. Pafupifupi theka la majeremusi a RXLR mu P. infestans genome ndimitundu yodziwika. Othandizira a RXLR ali ndi mayendedwe osiyanasiyana. Mwa iwo, mabanja amodzi akulu ndi 150 adadziwika. Mosiyana ndi proteni yayikulu, majini oyambitsa RXLR nthawi zambiri amakhala m'magawo osauka am'badwa komanso olemera obwereza a genome. Zinthu zamagetsi zomwe zimatsimikizira kukula kwa maderawa zimathandizira kupangidwanso m'matendawa.
Othandizira a Cytoplasmic CRN adadziwika koyambirira mu P. infestans zolemba zomwe zimalemba ma peptide am'mimba. Chiyambire kupezeka kwawo, ndizochepa zomwe zimadziwika ponena za banja la ochita izi. Kufufuza kwa mtundu wa P. Infestans kunawulula banja lalikulu la majini a CRN 196, omwe ndi akulu kwambiri kuposa a P. sojae (100 CRN) ndi P. ramorum (19 CRN). Monga ma RXLRs, ma CRN ndi ma protein osasintha ndipo amakhala ndi nkhokwe ya N-terminal LFLAK yotetezedwa kwambiri (50 amino acid) ndi dera loyandikana ndi DWL lokhala ndi majini osiyanasiyana. Ma CRN ambiri (60%) amakhala ndi peputayidi wazizindikiro.
Kuthekera kwa ma CRN osiyanasiyana kusokoneza njira zamagetsi za omwe akukhala nawo kwawerengedwa. Pakuwunika kwa necrosis yazomera, kuchotsedwa kwa mapuloteni a CRN2 kunapangitsa kuti zidziwike dera la C-terminal lomwe lili ndi ma 234 amino acid (malo 173-407, DXG domain) ndikupangitsa kufa kwa cell. Kufufuza kwa majeremusi a P. infestans CRN kuwulula zigawo zinayi zosiyana za ma C-terminal, zomwe zimayambitsanso kufa kwamkati mwa chomeracho. Izi zikuphatikiza madera omwe atchulidwa kumene a DC (P. Infestans ali ndi majini 18 ndi ma pseudogenes 49), komanso D2 (14 ndi 43) ndi madera a DBF (2 ndi 1) omwe amafanana ndi protein kinases. Mapuloteni a madera a CRN omwe amafotokozedwa mu chomera amasungidwa (pakalibe ma peptide azizindikiro) m'selo yazomera ndikulimbikitsa kufa kwama cell pogwiritsa ntchito makina amkati. Zotsatira zina 255 zokhala ndi magawo a CRN mwina sizigwira ntchito ngati majini.
Kuwonjezeka kwa kuchuluka ndi kukula kwa mabanja amtundu wa RXLR ndi CRN mwina mwina chifukwa cha kuphatikizanso kosagwiritsa ntchito mankhwala osokoneza bongo. Ngakhale kuti genome ili ndi zinthu zambiri zogwiritsira ntchito, palibenso umboni weniweni wosintha kwa majini oyambitsa.
Njira zomwe zimagwiritsidwa ntchito pophunzira kuchuluka kwa anthu
Kafukufuku wamitundu ya anthu pano akutengera kusanthula zikhalidwe zoyera zamavuto ake. Kuwunikiridwa kwa anthu osapatula mbewu zoyera kumachitikanso pazifukwa zina, monga, kuphunzira zaukali wa anthu kapena kupezeka kwa mitundu yolimbana ndi fungicides mmenemo (Filippov et al., 2004; Derevyagina et al., 1999). Kafukufuku wamtunduwu amaphatikizapo kugwiritsa ntchito njira zapadera, zomwe zimafotokozedwa zomwe sizingachitike pakuwunikaku. Pofufuza kuyerekezera kwa mitundu, njira zingapo zimagwiritsidwa ntchito potengera kusanthula kwa DNA kapenanso kuphunzira za mawonetseredwe a phenotypic. Kuyerekeza kuyerekezera kwa anthu kuyenera kuthana ndi anthu ambiri odzipatula, zomwe zimapereka zofunikira pazomwe amagwiritsa ntchito. Mwachidziwikire, ayenera kukwaniritsa zofunikira izi (Cooke, Lees, 2004, Mueller, Wolfenbarger, 1999):
- kukhala wotsika mtengo, wosavuta kuyigwiritsa ntchito, safuna nthawi yayikulu, kutengera ukadaulo womwe ulipo (mwachitsanzo, PCR);
- Ayenera kupanga zida zambiri zodziyimira palokha mokwanira;
- ali ndi kubereka kwakukulu;
- gwiritsani ntchito kuchuluka kwa minofu kuti mufufuze;
- khalani achindunji pa gawo lapansi (kuipitsidwa komwe kulipo pachikhalidwe sikuyenera kukhudza zotsatira);
- safuna kugwiritsa ntchito njira zowopsa komanso mankhwala owopsa.
Tsoka ilo, palibe njira zomwe zikugwirizana ndi magawo onsewa. Pofufuza mofananamo zamavuto m'masiku athu ano, njira zimagwiritsidwa ntchito potengera kusanthula kwa mikhalidwe ya phenotypic: virulence kwa mitundu ya mbatata ndi phwetekere (mitundu ya mbatata ndi phwetekere), mtundu wa mating, mtundu wa peptidase isoenzymes ndi glucose-6-phosphate isomerase, ndikuwunika kwa kapangidwe ka DNA: kutalika kwa polymorphism choletsa chidutswa (RFLP), chomwe nthawi zambiri chimaphatikizidwa ndi kafukufuku wosakanizidwa RG 57, kusanthula kubwereza kwa microsatellite (SSR ndi InterSSR), kukulitsa ndi zoyambira mwachisawawa (RAPD), kukulitsa kwa zidutswa zoletsa (AFLP), kukulitsa ndi zoyambira zomwe zimafanana ndi mafoni (mwachitsanzo, Inter SINE PCR), kutsimikiza kwa mitochondrial DNA haplotypes.
Kufotokozera mwachidule za njira zowerengera poyerekeza zamavuto omwe amagwiritsidwa ntchito ndi P. Infestans
Makhalidwe a phenotypic
Mitundu ya "Mbatata"
Mitundu ya "mbatata" ndimomwe amafufuzidwa komanso kugwiritsidwa ntchito. Mitundu "yosavuta ya mbatata" imakhala ndi jini imodzi yamatenda owopsa, "ovuta" - osachepera awiri. Black et al. (1953), powerengera mwachidule chidziwitso chonse chomwe adapeza, adapeza kuti mtundu wa phytophthora umatha kupatsira mbewu ndi jini / majini otsutsana ndi P. infestans virulence gene / majini, ndipo adapeza mitundu 1, 2, 3, ndi 4 yomwe imafalitsa mbewu ndi majini R1, R2, R3 ndi R4, motsatana, i.e. kulumikizana pakati pa tiziromboti ndi wolandirayo kumachitika molingana ndi jini la mfundo ya jini. Kuphatikiza apo, Black, potenga nawo mbali Gallegly ndi Malcolmson, adapeza mitundu yotsutsa R5, R6, R7, R8, R9, R10 ndi R11 komanso mitundu yofananira (Black, 1954; Black & Gallegly, 1957; Malcolmson & Black, 1966; Malcolmson, 1970).
Pali chidziwitso chokwanira pamtundu wamatenda omwe amachokera kumadera osiyanasiyana. Popanda kusanthula tsatanetsataneyu, tiziwonetsa njira yokhayo: pomwe mitundu yomwe ili ndi mitundu yatsopano yolimbana nayo kapenanso kuphatikiza kwake idagwiritsidwa ntchito, poyamba panali kufooka kwa vuto lakumapeto, koma kenako mitundu yomwe imafanana ndi majeremusi oyenera idawonekera ndikusankhidwa ndikuphulika kwa vuto lakumapeto kuyambiranso. Vutoli lenileni motsutsana ndi majeremusi anayi oyambilira (R4-R1) silinkawonekeranso m'magulu omwe anasonkhanitsidwa asanayambe kulima mitundu ndi majiniwa, koma kuchuluka kwa tizilombo toyambitsa matenda tidakulirakulira pomwe tizilomboto tidasakazidwa pamitundu yonyamula majini. Chibadwa 4-5, komano, chinali chofala kwambiri pamatola (Shaw, 11).
Kafukufuku wokhudza kuchuluka kwa mitundu yosiyanasiyana m'nyengo yokula, yomwe idachitika kumapeto kwa zaka za m'ma 1980, idawonetsa kuti kumayambiriro kwa matendawa, ndimatenda okhala ndiukali komanso ma 1-2 majeremusi omwe ali ndi kuchuluka kwa anthu.
Kuphatikiza apo, ndikukula kwa vuto lakumapeto, kuchuluka kwa miyala yoyambayo kumachepa ndipo kuchuluka kwa mafuko "ovuta" okhala ndiukali kwambiri kumakulirakulira. Zomwe zimachitika kumapeto kwa nyengo zifika 100%. Mukasunga ma tubers, kuchepa kwamphamvu kumachepetsa komanso kutayika kwamitundu yabwinobwino. Mphamvu zakusinthira kwamakina zimatha kuchitika mosiyanasiyana mosiyanasiyana (Rybakova & Dyakov, 1990). Komabe, maphunziro athu mu 2000-2010 awonetsa kuti mitundu yovuta imapezeka kuyambira koyambirira kwa ma epiphytotic pakati pamavuto omwe amakhala kutali ndi mbatata ndi tomato. Izi mwina chifukwa cha kusintha kwa anthu a P. Infestans ku Russia.
Pofika 1988-1995, kupezeka kwa "superraces" ndi mitundu yonse kapena pafupifupi mitundu yonse ya virulence m'malo osiyanasiyana kudafika 70-100%. Izi zidadziwika, mwachitsanzo, ku Belarus, zigawo za Leningrad ndi Moscow, ku North Ossetia ndi ku Germany (Ivanyuk et al., 2002a, 2002b; Polityko, 1994; Schober-Butin et al., 1995).
Mitundu ya "phwetekere"
M'mabzala a phwetekere, mitundu iwiri yokha yokana kulimbana ndi zovuta mochedwa idapezeka - Ph2 (Gallegly & Marvell, 1) ndi Ph1955 (Al-Kherb, 2). Monga momwe zimakhalira pamipikisano ya mbatata, kulumikizana pakati pa tomato ndi P. infestans kumachitika motsatira mibadwo. Mpikisano wa T1988 umayambitsa mitundu yomwe ilibe majini otsutsana (mitundu yambiri yogwiritsidwa ntchito m'makampani), mtundu wa T0 umayambitsa mitundu ndi mtundu wa Ph1 (Ottawa), ndipo mpikisano wa T1 umayambitsa mitundu ndi mtundu wa Ph2.
Ku Russia, pafupifupi T0 yokha idapezeka pa mbatata; T0 idapambana tomato koyambirira kwa nyengo, koma pambuyo pake idasinthidwa ndi mpikisano wa T1 (Dyakov et al., 1975, 1994). Pambuyo pa 2000, T1 pa mbatata mwa anthu ambiri idayamba kuchitika koyambirira kwa nthawi ya epiphytotic. Ku United States, mitundu ya mbatata sinali tizilombo toyambitsa matenda ku phwetekere, komanso mitundu T0, T1, ndi T2, pomwe T1 ndi T2 zimadalira tomato (Vartanian & Endo, 1985; Goodwin et al., 1995).
Mtundu wa mating
Pakafukufukuyu, mitundu ya tester (reference) yamitundu yodziwika yokwatirana - A1 ndi A2 amafunika. Kudzipatula koyesedwako kumayamwa ndi iwo awiriawiri m'mipukutu ya Petri yokhala ndi oat agar medium. Pambuyo pakusakaniza kwa masiku 10, mbale zimayesedwa ngati kulibe kapena kupezeka kwa oospores pakatikati pazolumikizana ndi zovuta. Pali zosankha 4: mtunduwo ndi wa mtundu wa mating A1, ngati wapanga ma oospores ndi woyeserera A2, kupita ku A2, ngati ungapangire oospores ndi woyesa A1, kupita ku A1A2, ngati ungapangire oospores ndi oyesa onse, kapena wosabala (00), ngati sapanga oospores wopanda woyesa (magulu awiri omalizawa ndi osowa).
Kuti mudziwe mitundu yamatenda mwachangu, zoyesayesa zidapangidwa kuti azindikire zigawo za matupi athu zomwe zimakhudzana ndi mtundu wa mating, ndi cholinga chowagwiritsanso ntchito kuti adziwe mtundu wa PCR. Chimodzi mwazoyesera zoyambirira kuti adziwe tsambalo chidachitika ndi ofufuza aku America (Judelson et al., 1995). Pogwiritsa ntchito njira ya RAPD, adakwanitsa kuzindikira dera la W16 lomwe limalumikizidwa ndi mtundu wa mating mwa ana awiriwa omwe adadutsa ndikukhazikitsa zopangira 24-bp pakukweza kwake (W16-1 (5'-AACACGCACAAGGCATATAAATGTA-3 ') ndi W16-2 (5 -GCGTAATGTAGCGTAACAGCTCTC-3 ') Pambuyo poletsa zomwe PCR idachita ndi zoletsa enzyme HaeIII, zinali zotheka kusiyanitsa zodzipatula ndi mitundu iwiri ya A1 ndi A2.
Kuyesanso kwina kopezera zolembera za PCR kuti adziwe mitundu yamatenda kunachitika ndi ofufuza aku Korea (Kim, Lee, 2002). Adazindikira zopangidwa pogwiritsa ntchito njira ya AFLP. Zotsatira zake, ma primers angapo a PHYB-1 (patsogolo) (5'-GATCGGATTAGTCAGACGAG-3 ') ndi PHYB-2 (5'-GCGTCTGCAAGGCCTTT-3') adapangidwa, kulola kukulitsa dera la genome lomwe limalumikizidwa ndi mtundu wa mating A2. Pambuyo pake, adapitiliza ntchitoyi ndikupanga zoyambira 5 'AAGCTATACTGGGACAGGGT-3' (INF-1, patsogolo) ndi 5'-GCGTTCTTTCGTATTACCAC-3 '(INF-2), kulola kukulitsa dera la Mat-A1, mawonekedwe amtundu wokhala ndi mtundu wa mating A1. Kugwiritsa ntchito mitundu ya PCR yozindikira mating kukuwonetsa zotsatira zabwino pakuphunzira kwa P. infestans anthu ku Czech Republic (Mazakova et al., 2006), Tunisia (Jmour, Hamada, 2006), ndi madera ena. Mu labotale yathu (Mytsa, Elansky, yosasindikizidwa), 34 P. infestans tizilombo ta matenda omwe amapezeka kumatenda odwala ndi ziwalo za phwetekere m'malo osiyanasiyana ku Russia (Kostroma, Ryazan, Astrakhan, ndi Moscow) adasanthulidwa. Zotsatira zakusanthula kwa PCR pogwiritsa ntchito zoyambira zingapo zopitilira 90% zidagwirizana ndi zotsatira za kusanthula kwamtundu wa mating malinga ndi njira yachikhalidwe pamagawo azakudya.
Gulu 1. Kusiyanasiyana kwa kukana mkati mwazitsulo la Sib 1 (Elansky et al., 2001)
Zitsanzo zosonkhanitsira malo | Chiwerengero chazokha zomwe zidawunikidwa | Chiwerengero cha zovuta (S), zofooka zosagwira (SR) ndi zosagwira (R) zovuta, ma PC (%) | ||
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 |
Mzinda wa Yekaterinburg | 15 | 8 (53) | 1 (7) | 6 (40) |
O. Sakhalin | 66 | 0 | 0 | 66 (100) |
Dera la Omsk | 18 | 0 | 0 | 18 (100) |
Kukaniza kwa Metalaxyl monga chikhomo cha anthu
Kumayambiriro kwa zaka za m'ma 1980, kufalikira kwamphamvu kwamatenda akuchedwa chifukwa cha mitundu yolimbana ndi metalaxyl P. infestans kunadziwika m'malo osiyanasiyana. Minda yama mbatata m'maiko ambiri yatayika kwambiri (Dowley & O'Sullivan, 1981; Davidse et al., 1983; Derevyagina, 1991). Kuyambira pamenepo, m'maiko ambiri padziko lapansi, kuwunika mosalekeza kupezeka kwa mitundu yolimbana ndi phenylamide mwa anthu a P. infestans kwachitika. Kuphatikiza pa kuwunika koyenera kwa chiyembekezo chogwiritsa ntchito mankhwala okhala ndi phenylamide, kupanga njira zodzitetezera komanso kuneneratu za epiphytoties, kukana mankhwalawa kwakhala chimodzi mwazomwe zimagwiritsidwa ntchito kwambiri pofufuza poyerekeza kuchuluka kwa tizilomboto. Komabe, kugwiritsidwa ntchito kwa kukana kwa metalaxyl m'maphunziro owerengera anthu kuyenera kuchitidwa poganizira kuti: 1 - maziko a kukana sanatsimikizidwebe bwinobwino, 2 - kukana metalaxyl ndichikhalidwe chodalira chomwe chingasinthe kutengera kugwiritsa ntchito phenylamides, 3 - zosiyana kuchuluka kwa chidwi cha mitundu ya metalaxyl mkati mwa mzere umodzi (tebulo. 1).
Makonda a isozymes
Zolemba za Isozyme nthawi zambiri sizimayenderana ndi zakunja, zimawonetsa cholowa cha Mendelian ndipo ndizodziwika bwino, zomwe zimalola kusiyanitsa pakati pa homo- ndi heterozygotes. Kugwiritsiridwa ntchito kwa mapuloteni monga zolembera zam'magazi kumapangitsa kuti athe kuzindikira kukonzanso kwamitundu yayikulu, kuphatikiza kusintha kwa chromosomal ndi genomic, komanso m'malo mwa amino acid osakwatira.
Kafukufuku wama electrophoretic wamapuloteni awonetsa kuti michere yambiri imapezeka m'zinthu zazing'onozing'ono zomwe zimasiyana mosiyanasiyana pamagetsi. Zigawozi zimachokera pakukhazikitsa mitundu ingapo yama michere ndi ma loci osiyanasiyana (isozymes kapena isozymes) kapena ma alleles osiyanasiyana amalo omwewo (allozymes kapena alloenzymes). Ndiye kuti, isozymes ndi mitundu yosiyanasiyana ya enzyme imodzi. Mitundu yosiyanasiyana imakhala ndi zochitika zofananira, koma imasiyana pang'ono m'malo amodzi amino acid mu peptide komanso woyang'anira. Kusiyanaku kumawululidwa panthawi ya electrophoresis.
Pakafukufuku wa P. infestans strains, mawonekedwe a isoenzymes a mapuloteni awiri, peptidase ndi glucose-6-phosphate isomerase, amagwiritsidwa ntchito (enzyme iyi ndi monomorphic mwa anthu aku Russia, chifukwa chake, njira zophunzirira zake sizikupezeka pantchitoyi). Kuti muwalekanitse mu isozymes m'munda wamagetsi, mapuloteni okonzedwa kuchokera kuzinthu zophunziridwa amagwiritsidwa ntchito pa mbale ya gel yomwe imayikidwa mumagetsi. Kuchuluka kwa mapuloteni amtundu uliwonse mu gel osakaniza kumadalira kuchuluka kwake ndi kulemera kwake; chifukwa chake, pamagetsi, kuphatikiza kwa mapuloteni kumagawika m'magawo awiri, omwe amatha kuwonetsedwa pogwiritsa ntchito utoto wapadera.
Kafukufuku wa peptidase isoenzymes amachitika pa cellulose-acetate, wowuma kapena ma polyacrylamide gels. Njira yabwino kwambiri ndiyo njira yogwiritsira ntchito mapadi a acetate opangidwa ndi Helena Laboratories Inc. Sichifuna kuchuluka kwa zida zoyeserera, zimalola munthu kupeza magulu osiyanasiyana pa gel pambuyo pa electrophoresis ya enzyme loci, kukhazikitsa kwake sikutanthauza nthawi yayikulu komanso ndalama zakuthupi (mkuyu 2).
Chidutswa chochepa cha mycelium chimasamutsidwa mu 1,5 ml microtube, madontho 1-2 a madzi osungunulidwa amawonjezeredwa. Pambuyo pake, chitsanzocho chimapangidwanso (mwachitsanzo, ndi kubowola kwamagetsi ndi cholumikizira cha pulasitiki choyenera microtube) ndikuchepetsa masekondi 25 pa centrifuge pa 13000 rpm. 8 μl kuchokera pa microtube iliyonse. supernatant imasamutsidwa ku mbale yogwiritsa ntchito.
Selulo ya acetate ya gelisi imachotsedwa mu chidebe chomenyera, imapukutidwa pakati pa mapepala awiri a fyuluta ndikuiyika ndi gawo logwirira ntchito pamwamba papulasitiki. Njira yothetsera mbaleyo imasunthidwa ndi ogwiritsa ntchitoyo mpaka gel osakaniza kangapo. Gelayi imasamutsidwa kuchipinda cha electrophoresis,
Gulu 2. Kapangidwe ka yankho lomwe limagwiritsidwa ntchito pothana ndi mapadi a acetate gel posanthula peptidase isoenzymes, dontho la utoto (bromophenol buluu) limayikidwa m'mphepete mwa gel.
TRIS HCl, 0,05M, Ph 8,0 2 ml
Peroxidase, 1000 U / ml madontho 5
o-dianisidine, 4 mg / ml madontho 8
MgCl2, 20 mg / ml madontho awiri
Gly-Leu, 15 mg / ml madontho 10
L-amino-acid oxidase, 20 u / ml madontho awiri
Electrophoresis imachitika kwa mphindi 20. pa 200 V. Pambuyo pa electrophoresis, gelisi imasamutsidwa patebulo lojambula ndikudetsedwa ndi yankho lapadera la penti (Gulu 2). 10 ml ya 1,6% DIFCO agar imasungunuka koyambirira mu uvuni wa microwave, utakhazikika mpaka 60 ° C, pambuyo pake 2 ml ya agar imasakanizidwa ndi kaphatikizidwe ka utoto ndikutsanulira gel osakaniza. Mikwingwirima imawonekera mkati mwa mphindi 15-20. L-amino-acid oxidase reagent imawonjezedwa nthawi yomweyo asanasakanize yankho ndi agar osungunuka.
M'madera aku Russia, malo a Pep 1 amaimiridwa ndi genotypes 100/100 ndi 92/100. Homozygote 92/92 ndi yosowa kwambiri (pafupifupi 0,1%). Locus Rehr 2 imayimilidwa ndi mitundu itatu ya genotypes 100/100, 100/112, ndi 112/112, ndipo mitundu yonse itatu ndiyofala (Elanky ndi Smirnov, 3, mkuyu. 2003).
Kafukufuku wa Genome
Cholepheretsa kutalika kwa chidutswa cha polymorphism ndi zotsatira zosakanizidwa (RFLP-RG 57)
DNA yonse imathandizidwa ndi enzyme yoletsa Eco R1, zidutswa za DNA zimasiyanitsidwa ndi electrophoresis mu agarose gel. Nyukiliya DNA ndi yayikulu kwambiri ndipo imakhala ndi zochitika zingapo zobwerezabwereza, zomwe zimapangitsa kuti zikhale zovuta kusanthula mwachindunji zidutswa zingapo zomwe zimapezeka chifukwa cha michere yoletsa. Chifukwa chake, zidutswa za DNA zomwe zidasiyanitsidwa ndi gel osakaniza zimasamutsidwa ku nembanemba yapadera ndikugwiritsidwa ntchito kuphatikizira ndi kafukufuku wa RG 57, womwenso ma nucleotide olembedwa ndi zilembo za radioactive kapena fluorescent. Kafukufukuyu amaphatikizika ndimabuku obwereza obwereza (Goodwin et al., 1992, Forbes et al., 1998). Pambuyo pakuwona zotsatira zakusakanikirana ndi zinthu zowunikira kapena zowulutsa ma radiation, mawonekedwe amitundu yambiri ophatikizika (zolemba zala) amapezeka, oyimiriridwa ndi zidutswa za 25-29 (Forbes et al., 1998). Asexual (clonal) ana adzakhala ndi mbiri zofananira. Mwa makonzedwe amtundu wa electrophoretogram, munthu amatha kuweruza kufanana ndi kusiyana kwa zinthu zofananazo.
DNA ya Mitochondrial haplotypes
M'maselo ambiri a eukaryotic, mtDNA imawoneka ngati ma molekyulu ozungulira a DNA, omwe, mosiyana ndi ma chromosomes a nyukiliya amamaselo a eukaryotic, amafanananso mosasamala komanso sagwirizana ndi ma molekyulu a protein.
Matupi a mitochondrial a P. infestans adatsatiridwa, ndipo ntchito zingapo zidaperekedwa pakuwunika kwa zoletsa zazing'ono (Carter Et al, 1990, Goodwin, 1991, Gavino, Fry, 2002). Griffith ndi Shaw (1998) atapanga njira yosavuta komanso yofulumira yodziwira mtDNA haplotypes, chikhomochi chidakhala chimodzi mwazodziwika kwambiri m'maphunziro a P. Infestans. F2-R2 (Gulu 4) ndi zoletsa zawo zotsatila ndi michere yoletsa MspI (chidutswa cha 4) ndi EcoR3 (chidutswa chachiwiri). Njirayi imakupatsani mwayi wodziwa ma haplotypes 1: Ia, IIa, Ib, IIb. Mtundu Wachiwiri umasiyana ndi mtundu I pakupezeka kwa 1 bp komanso malo osiyana siyana oletsa madera P2 ndi P4 (mkuyu 1881).
Kuyambira 1996, pakati pamavuto omwe amasonkhanitsidwa ku Russia, ma haplotypes Ia ndi IIa okha ndi omwe adadziwika (Elansky et al., 2001, 2015). Amatha kudziwika atapatukana ndi zoletsa ndi F2-R2 choyambira pamunda wamagetsi (mkuyu. 4, 5). Mitundu ya mtDNA imagwiritsidwa ntchito pakuwunika kofananira kwa mitundu ndi anthu. Mu ntchito zingapo, mitundu ya DNA ya mitochondrial idagwiritsidwa ntchito kupatula mizere yama clonal ndikudutsa ma P. infestans odzilekanitsa (Botez et al., 2007; Shein et al., 2009). Pogwiritsa ntchito njira ya PCR-RFLP, zidatsimikizika kuti mtDNA ndiyosiyana kwambiri mu zovuta za P. infestans (Elansky ndi Milyutina, 2007). Zinthu zokukweza: 1x (mphindi 500. 94 ° C), 40x (mphindi 30. 90 ° C, mphindi 30. 52 ° C, mphindi 90. 72 ° C); 1x (5 min. 72 ° C). Kusakaniza kosakanikirana: (20 μl): 0,2 U Taq DNA polymerase, 1x 2,5 mM MgCl2-Taq buffer, 0,2 mM dNTP iliyonse, 30 pM choyambira ndi 5 ng ya DNA yosanthula, madzi osakanikirana - mpaka 20 μl.
Kuletsa kwa mankhwala a PCR kumachitika kwa maola 4-6 kutentha kwa 37 ° C. Kuletsa kusakaniza (20 μl): 10x MspI (2 μl), 10x choletsa buffer (2 μl), madzi osungunuka (6 μl), PCR product (10 μl).
Gulu 3. Oyambira omwe amagwiritsidwa ntchito kukulitsa zigawo za mtDNA polymorphic
Malo | Malangizo oyambira | Kutalika koyambirira ndi kusungidwa | Kutalika kwa mankhwala a PCR | Kuchepetsa |
---|---|---|---|---|
P2 | F2: 5'- TTCCCTTTGTCCTCTACCGAT | 21; 13619-13639 | 1070 | MspI |
R2: 5'- TTACGGCGGTTTAGCACATACA | 22; 14688-14667 | |||
P4 | F4: 5'- TGGTCATCCAGAGGTTTATGTT | 22; 9329-9350 | 964 | EcoRI |
R4:5 - CCGATACCGATACCAGCACCAA | 22; 10292-10271 |
Kukulitsa mwachisawawa (RAPD)
Pochita RAPD, choyambira chimodzi chimagwiritsidwa ntchito (nthawi zina choyambira zingapo nthawi imodzi) ndimayendedwe osakanikirana a nucleotide, nthawi zambiri ma 10 a nucleotide m'litali, okhala ndi (50%) a GC nucleotides komanso kutentha kotsalira kotsalira (pafupifupi 35 ° C). Zoterezi "zimakhazikika" m'malo ambiri othandizira mu genome. Pambuyo pa kukulitsa, pamapezeka zida zazikulu zambiri. Chiwerengero chawo chimadalira zoyambira zomwe zagwiritsidwa ntchito komanso momwe zimachitikira (ndende ya MgCl2 ndi kutentha kwowonjezera).
Kuwonetseratu ma amplicons kumachitika chifukwa cha distillation mu polyacrylamide kapena agarose gel. Pochita kusanthula kwa RAPD, ndikofunikira kuwunika mosamala kuyera kwa zomwe zafufuzidwazo, popeza kuipitsidwa ndi zinthu zina zamoyo kungayambitse kuchuluka kwakukulu kwa zinthu zakale, zomwe ndizochulukirapo ngakhale pakupenda zinthu zoyera (Perez et al, 1998). Kugwiritsa ntchito njirayi pophunzira za P. infestans genome kumawonekera m'mabuku ambiri (Judelson, Roberts, 1999, Ghimire et al., 2002, Carlisle et al., 2001). Kusankhidwa kwa zoyankha ndi zoyambira (51 10-nucleotide primers adawerengedwa) zimaperekedwa m'nkhaniyi ndi Abu-El Samen et al., (2003).
Kusanthula Kubwereza kwa Microsatellite (SSR)
Kubwereza kwa Microsatellite (kubwereza kosavuta, SSR) kumabwereza mwachidule magawo a 1-3 (nthawi zina mpaka 6) a ma nucleotide omwe amapezeka mu ma genomes anyukiliya a ma eukaryote onse. Chiwerengero chobwereza motsatizana chimatha kuyambira 10 mpaka 100. Microsatellite loci imachitika pafupipafupi kwambiri ndipo imagawidwa mofananamo mu genome (Lagercrantz et al., 1993). Polymorphism yotsatizana kwa microsatellite imagwirizanitsidwa ndi kusiyanasiyana kwa kuchuluka kwa kubwereza kwa cholinga choyambirira. Zolemba za Microsatellite ndizodziwika bwino, zomwe zimapangitsa kuti zizigwiritsidwa ntchito pofufuza kuchuluka kwa anthu, kudziwa ubale, njira zosamukira, ndi zina zambiri. Zina mwazabwino za zolembazi, munthu ayenera kuzindikira kuchuluka kwawo kwa ma polymorphism, kubereka bwino, kusalowerera ndale, komanso kuthekera kochita kuwunika ndikuwunika. Kuwunika kwa polymorphism ya microsatellite kubwereza kumachitika ndi kukweza kwa PCR pogwiritsa ntchito zoyambilira zothandizirana ndi zochitika zapadera za microsatellite loci. Poyamba, kuwunikaku kunachitika ndikulekanitsa zomwe zimachitika pa gel ya polyacrylamide. Pambuyo pake, ogwira ntchito ku kampani ya Applied Biosystems adaganiza zogwiritsa ntchito zopangira zolembedwa bwino pozindikira zomwe zimachitika pogwiritsa ntchito chowunikira cha laser (Diehl et al., 1990), kenako ma sequencers wamba a DNA (Ziegle et al., 1992). Kulemba ma primers okhala ndi utoto wosiyanasiyana wa fulorosenti kumatha kuwunikira zolembera zingapo nthawi imodzi pamsewu umodzi ndipo, moyenera, zimawonjezera zokolola za njirayi ndikuwonjezera kulondola kwa kusanthula.
Zolemba zoyambirira zomwe zidagwiritsidwa ntchito pakuwunika kwa SSR pakuphunzira kwa P. infestans zidapezeka koyambirira kwa 2000s. (Knapova, Gisi, 2002). Sikuti zilembo zonse zomwe olembawo adalemba zidawonetsa kuchuluka kwa ma polymorphism, komabe, awiriwo (4B ndi G11) adaphatikizidwa m'gulu la zilembo za 12 SSR zoperekedwa ndi Lees et al. (2006) ndipo pambuyo pake adatengera ukonde wofufuza wa Eucablight (www.eucablight) .org) ngati muyezo wa P. infestans. Zaka zingapo pambuyo pake, kafukufuku adasindikizidwa pakupanga njira yowunikira ma multiplex a P. infestans DNA yozikidwa ndi zilembo zisanu ndi zitatu za SSR (Li et al., 2010). Pomaliza, atasanthula zolembera zonse zomwe zidafotokozedwapo kale ndikusankha zomwe zitha kuphunzitsidwa bwino, komanso kukonza ma primers, zolemba za fluorescent, ndi zokulitsa, gulu lomwelo la olemba lidapereka njira yosanthula masitepe angapo, kuphatikiza zolembera za 12 (Gulu 4; Li et al. , 2013a). Zoyambira zomwe zidagwiritsidwa ntchito m'dongosolo lino zidasankhidwa ndikulembedwa ndi imodzi mwazoyala zinayi za fulorosenti (FAM, VIC, NED, PET) kuti magulu azithunzi zoyambira omwe ali ndi zilembo zomwezo asadutse.
Olembawo adasanthula pa PTC200 amplifier (MJ Research, USA) pogwiritsa ntchito zida za QIAGEN multiplex PCR kapena QIAGEN Typeit Microsatellite PCR. Voliyumu ya zosakanikirazo inali 12.5 μL. Zinthu zokukweza zinali motere: kwa QIAGEN multiplex PCR: 95 ° C (15 min), 30x (95 ° C (20 s), 58 ° C (90 s), 72 ° C (60 s), 72 ° C (20 min); ya QIAGEN Type-it Microsatellite PCR: 95 ° C (5 min), 28x (95 ° C (30 sec), 58 ° C (90 sec), 72 ° C (20 sec), 60 ° C (30 min).
Kupatukana ndi kuwonera zinthu za PCR kunachitika pogwiritsa ntchito ABI3730 yodziyimira yokha ya capillary DNA analyzer (Applied Biosystems).
Gulu 4. Makhalidwe azizindikiro za 12 zovomerezeka za SSR zomwe zimagwiritsidwa ntchito popanga P. Infestans (Li et al., 2013a)
Mutu | Chiwerengero cha ma alleles | Kukula kwake Mitsinje (bp) | Oyambirira |
Onetsani: 11 | 13 | 130-180 | F: NED-TGCTATTTATCAAGCGTGGG R: GTTTCAATCTGCAGCCGTAAGA |
Kondanani | 4 | 255-275 | F: NED-ACTTGCAGAACTACCGCCC R: GTTTGACCACTTTCCTCGGTTC |
Sakanizani: | 4 | 325-360 | Gawo #: NED-TTAAGCCACGACATGAGCTG R: GTTTAGACAATTGTTTTGTGGTCGC |
D13 | 16 | 100-185 | F: FAM-TGCCCCCTCTGCTCACTC R: GCTCGAATTCATTTTACAGACTTG |
Sakanizani: | 4 | 250-275 | FAM-AATCTGATCGCAACTGAGGG R: GTTTACAAGATACACACGTCGCTCC |
Sakanizani: | 7 | 280-305 | Kufotokozera: FAM-TCTTGTTCGAGTATGCGACG R: GTTTCACTTCGGGAAAGGCTTC |
Kondanani | 4 | 160-175 | F: VIC-AGCGGCTTACCGATGG R: GTTTCAGCGGCTGTTTCGAC Chidziwitso: |
Kondanani | 3 | 185-205 | F: VIC-ATGAAAATACGTCAATGCTCG R: CGTTGGATATTTCTATTTCTTCG |
Sakanizani: | 3 | 230-250 | F: GTTTTGGTGGGTWGTTTT R: VIC-TCGCCACAAGATTTATTCCG |
Kondanani | 3 | 265-280 | F: VIC-ATGACGAAGATGAAAGTGAGG R: CGTATTTTCCTGTTATATAACACC |
Sakanizani: | 3 | 165-180 | F: Dongosolo la PET-CGACTTCTACATCAACCGGC R: GTTTGCTTGGACTGCGTCTTTAGC |
Pi4B | 5 | 200-295 | F: PET-AAAATAAAGCCTTTGGTTCA R: GCAAGCGAGGTTTGTAGATT |
Chitsanzo chowonera zotsatira zakusanthula chikuwonetsedwa mkuyu. 6. Zotsatirazo zidasanthulidwa pogwiritsa ntchito pulogalamu ya GeneMapper 3.7 poyerekeza zomwe zapezeka ndi zomwe zimadziwika zodzipatula. Kuwongolera kutanthauzira kwa zotsatira zakusanthula, ndikofunikira kuphatikiza omwe akutenga nawo mbali 1-2 omwe ali ndi genotype wodziwika paphunziro lililonse.
Njira yofufuzira yomwe adayeserayi idayesedwa pamitundu ingapo yamasamba, pambuyo pake olembawo adakhazikitsa ndondomeko pakati pama laboratories amabungwe awiri, James Hutton Institute (UK) ndi Wageningen University & Research (Netherlands), yomwe, kuphatikiza mwayi wogwiritsa ntchito makhadi a FTA osavuta Kutolera ndi kutumiza kwa P. infestans zitsanzo za DNA, zidatilola kuti tikambirane za kuthekera kogulitsa zamtunduwu. Kuphatikiza apo, njira yachangu komanso yolondola ya genotyping P. infestans imadzipatula pogwiritsa ntchito kusanthula kwa multiplex SSR kunapangitsa kuti athe kuchita kafukufuku wovomerezeka wa anthu omwe ali ndi tizilomboto padziko lonse lapansi, ndikupanga nkhokwe yapadziko lonse lapansi mochedwa chifukwa cha ntchito ya Eucablight (www.eucablight.org), kuphatikiza , kuphatikiza zotsatira zakusanthula kwa microsatellite, zidatheketsa kutsata kufalikira ndi kufalikira kwa mitundu yatsopano yamitundu padziko lonse lapansi.
Amplified choletsa chidutswa cha kutalika kwa polymorphism (AFLP). AFLP (kutalika kwa chidutswa cha polymorphism) ndiukadaulo wopanga ma molekyulu osagwiritsa ntchito mawonekedwe oyambira. Mu AFLP, DNA imathandizidwa ndikuphatikiza ma michere iwiri yoletsa. Ma adapter apadera amalumikizidwa kumapeto kwa zidutswa zoletsa.
Zidutswazi zimakwezedwa ndikugwiritsa ntchito zopangira zowonjezerapo kutsata kwa ma adapter ndi zoletsa, ndikuwonjeza chimodzi kapena zingapo zosakwanira kumapeto kwawo atatu. Zidutswa zomwe zidapezedwa zimatengera ma enzyme oletsa komanso ma nucleotide osankhidwa mwachisawawa kumapeto kwa 3'-end of the primers (Vos et al., 3). AFLP - genotyping imagwiritsidwa ntchito kuti iphunzire mwachangu masinthidwe amtundu wazinthu zosiyanasiyana.
Malongosoledwe atsatanetsatane a njirayi amaperekedwa mu ntchito za Mueller, Wolfenbarger, 1999, Savelkoul et al., 1999. Ntchito yambiri yoyerekeza kuthana ndi njira za AFLP ndi SSR yachitika ndi ofufuza aku China. Ma phenotypic ndi genotypic a 48 P. infestans amadzipatula omwe amasonkhanitsidwa kumadera asanu aku North China adaphunziridwa. Masewera a AFLP adawulula mitundu isanu ndi itatu ya DNA, mosiyana ndi ma SSR genotypes, omwe sanapezeke mosiyanasiyana (Guo et al., 2008).
Kukulitsa ndi zoyambira zoyambira kutengera momwe zinthu zimayendera
Zolemba zomwe zimachokera ku ma retrotransposons ndizosavuta kupanga mapangidwe amtundu, kafukufuku wamitundu yosiyanasiyana ndi njira zosinthira (Schulman, 2006). Zolemba zoyambilira zikapangidwira kuti zizigwirizana ndimayendedwe amtundu wina wama foni, ndizotheka kukulitsa zigawo za genome zomwe zili pakati pawo. M'maphunziro a causative agent wa vuto lakumapeto kwa njira, njira zokulitsira mbali za genome pogwiritsa ntchito choyambira chothandizirana ndi SINE (Short Interspersed Nuclear Elements) retropazone idagwiritsidwa ntchito bwino (Lavrova ndi Elansky, 2003). Pogwiritsa ntchito njirayi, kusiyana kudawululidwa ngakhale mwa ana oberekera omwe amakhala m'modzi. Pankhaniyi, zidatsimikizika kuti njira yapakati pa SINE - PCR ndiyodziwika bwino ndipo kuchuluka kwa mayendedwe a SINE mu mtundu wa Phytophthora ndikokwera.
Mu genome ya P. infestans, mabanja 12 a ma short retrotransposons (SINEs) adadziwika; magawidwe amitundu yama retrotransposons amafupipafupi anafufuzidwa, ma element (SINEs) adadziwika omwe amapezeka mu genome la P. infestans okha (Lavrova, 2004).
Makhalidwe ogwiritsa ntchito njira zowerengera zowerengera zamavuto m'maphunziro aanthu
Pokonzekera kuphunzira, ndikofunikira kumvetsetsa bwino zolinga zomwe zimatsata ndikugwiritsa ntchito njira zoyenera. Chifukwa chake, njira zina zimaloleza kupanga zizindikilo zambiri zodziyimira pawokha, koma nthawi yomweyo zimakhala ndi kubereka kocheperako ndipo zimadalira kwambiri ma reagents omwe amagwiritsidwa ntchito, momwe zimachitikira, komanso kuipitsidwa kwa zomwe zikuphunziridwa. Chifukwa chake, pakuwunika konse kwa gulu la mitundu, pamafunika kugwiritsa ntchito miyezo ingapo (maumboni) olekanitsa, koma ngakhale pano, zotsatira zoyesera zingapo ndizovuta kwambiri kuphatikiza.
Gulu la njirazi limaphatikizapo RAPD, AFLP, InterSSR, InterSINE PCR. Pambuyo pakukweza, zidutswa zambiri za DNA zamitundu yosiyanasiyana zimapezeka. Ndikofunika kugwiritsa ntchito njira ngati izi pakufunika kukhazikitsa kusiyana pakati pa mitundu yokhudzana kwambiri (kholo-kholo, zosintha zamtchire, ndi zina zambiri), kapena ngati pangafunike kusanthula pang'ono zazitsanzo zazing'ono. Chifukwa chake, njira ya AFLP imagwiritsidwa ntchito kwambiri pakupanga mapangidwe a P. infestans (van der Lee et al., 1997) komanso m'maphunziro a intrapopulation (Knapova, Gisi, 2002, Cooke et al, 2003, Flier et al, 2003). Njira zotere sizingagwiritsidwe ntchito popanga nkhokwe zamtundu, kuyambira Ndizosatheka kuphatikizira kuwerengera kwa zotsatira mukamasanthula muma laboratories osiyanasiyana.
Ngakhale zikuwoneka ngati kuphweka komanso kuthamanga kwakupha (kudzipatula kwa DNA kopanda kuyeretsa bwino, kukulitsa, kuwonera zotsatira), gulu ili la njira limafunikira kugwiritsa ntchito njira yapadera yolembera zotsatirazi: distillation mu polyacrylamide gel yokhala ndi zolembedwa (radioactively kapena luminescent) zoyambira ndikuwunikiridwa ndi zinthu zowunikira kapena zowunikira. Kulingalira moyenera kwa ethidium bromide agarose kujambula kwa gel sikuli koyenera njirazi chifukwa zidutswa zingapo za DNA zamitundu yosiyanasiyana zitha kusakanikirana.
Njira zina, m'malo mwake, zimaloleza kupanga zida zochepa ndi kubereka kwawo kwakukulu. Gululi limaphatikizapo kuphunzira za mitochondrial DNA haplotypes (ma haplotypes awiri okha a Ia ndi IIa amadziwika ku Russia), mitundu yosakanikirana (ambiri omwe amakhala okhaokha amagawika m'magulu awiri: A2 ndi A1, SF yodziberekera siyipezeka) ndi peptidase isozyme spectra (awiri a loci Pep2 ndi Pep1 , wopangidwa ndi ma isozymes awiri aliyense) ndi glucose-2-phosphate isomerase (ku Russia palibe kusiyanasiyana kwa khalidweli, ngakhale kuti ma polymorphism ofunikira amadziwika m'maiko ena adziko lapansi). Ndibwino kuti mugwiritse ntchito izi pofufuza zopereka, polemba madera akumidzi ndi apadziko lonse lapansi. Pakufufuza kwa isozymes ndi haplotypes ya mitochondrial DNA, ndizotheka kuchita popanda zovuta zilizonse, pomwe pakuwunika mitundu ya mating, mayeso awiri omwe amakhala ndi mitundu yodziwika bwino amafunika.
Zomwe zimachitika ndi ma reagents zimatha kukhudza kusiyanasiyana kwa malonda pa electrophoretogram; kuwonekera kwa zinthu zakale m'mitundu yamaphunziro iyi ndizokayikitsa.
Pakadali pano, anthu ambiri mdera la Europe ku Russia akuyimiridwa ndi mitundu ya mating (Table 6), pakati pawo pali mitundu yodzipatula yokhala ndi mitundu Ia ndi IIa ya mitochondrial DNA (mitundu ina ya mtDNA yomwe idapezeka padziko lapansi sinapezeke ku Russia pambuyo pa 1993). Mawonekedwe a peptidase isozymes amaimiridwa ndi ma genotypes awiri ku Pep1 locus (100/100, 92/92 ndi heterozygote 92/100, ndipo 92/92 genotype ndiyosowa kwambiri (<0,3%)) ndi ma genotypes awiri ku Pep 2 locus (100/100 , 112/112 ndi heterozygote 100/112, yokhala ndi genotype 112/112 yomwe imachitika pafupipafupi kuposa 100/100, komanso nthawi zambiri).
Panalibe kusiyanasiyana kwamagulu a isoenzymes a glucose-6-phosphate isomerase pambuyo pa 1993 (kutha kwa clonal line US-1); onse omwe anaphunzitsidwa kuti ali okhaokha anali ndi 100/100 genotype (Elansky ndi Smirnov, 2002).
Gulu lachitatu la njira limathandizira kupeza gulu lokwanira la zilembo zodziyimira palokha zomwe zimatha kuberekanso kwambiri. Lero, gululi likuphatikiza kafukufuku wa RFLP-RG57, womwe umatulutsa zidutswa za 25-29 za DNA zamitundu yosiyanasiyana. RFLP-RG57 itha kugwiritsidwa ntchito pofufuza zitsanzo ndikupanga nkhokwe. Komabe, njirayi ndi yokwera mtengo kwambiri kuposa yam'mbuyomu, ndiyotenga nthawi, ndipo imafuna kuchuluka kwakukulu kwa DNA yoyeretsedwa kwambiri. Chifukwa chake, wofufuzirayo akukakamizidwa kuchepetsa kuchuluka kwa zomwe zayesedwa.
Kukula kwa RFLP-RG57 koyambirira kwa zaka za m'ma 90 zam'zaka zapitazi kudakulitsanso maphunziro a anthu omwe amayambitsa vuto lakumapeto kwa ngozi. Inakhala maziko a njirayi potengera kusankha ndi kusanthula "Clonal lines" (onani pansipa). Pamodzi ndi RFLP-RG57, mtundu wa mating, zolemba zala za DNA (njira ya RFLP-RG57), mawonekedwe a peptidase ndi glucose-6-phosphate isomerase isoenzymes, ndi mtundu wa DNA wa mitochondrial amagwiritsidwa ntchito pozindikira mizere. Chifukwa cha iye, adawonetsedwa al., 1994), m'malo mwa anthu akale ndi atsopano (Drenth Et al, 1993, Sujkowski Et al, 1994, Goodwin Et al, 1995a), ndi mibadwo yama clonal yomwe ikupezeka m'maiko ambiri adziko lapansi idadziwika. Kafukufuku wazovuta zaku Russia pogwiritsa ntchito njirayi adawonetsa kuphulika kwamitundu yayikulu yamagawo aku Europe ndikuwonjezeka kwa anthu aku Asia ndi Far East ku Russia (Elansky et al, 2001). Ndipo njira iyi ikadali yofunikira kwambiri pamaphunziro a anthu a P. infestans. Komabe, kufalikira kwake kumalephereka chifukwa chokwera mtengo komanso kuchuluka kwa ogwira ntchito pakuphedwa.
Kufufuza kwa Microsatellite repeat (SSR) ndi njira ina yolonjeza yomwe imagwiritsidwa ntchito kawirikawiri pamaphunziro a P. infestans. Pakadali pano, njirayi imagwiritsidwa ntchito kupatula mizere yama clonal. Pofufuza zamavuto, mawonekedwe a phenotypic marker monga kupezeka kwa majeremusi amtundu wa mbatata (Avdey, 1995; Ivanyuk et al., 2002; Ulanova et al., 2003) ndi phwetekere amagwiritsidwa ntchito kwambiri (ndipo akupitilizabe kugwiritsidwa ntchito). Pakadali pano, mitundu yabwinobwino ya mitundu ya mbatata yataya mtengo wake monga zizindikilo zamaphunziro aanthu chifukwa chowoneka (kapena pafupi) kuchuluka kwa majeremusi achifwamba m'malo ambiri. Nthawi yomweyo, mtundu wa T1 virulence wa ma phwetekere omwe amakhala ndi mtundu wofanana wa Ph1 umagwiritsidwabe ntchito bwino ngati chikhomo (Lavrova et al., 2003; Ulanova et al., 2003).
M'maphunziro ambiri, kulimbana ndi fungicide kumagwiritsidwa ntchito ngati chikhomo. Khalidwe ili ndilofunika kugwiritsa ntchito m'maphunziro aanthu chifukwa chowoneka kosavuta kosintha kwa kukana mizere yama clonal mutagwiritsa ntchito metalaxyl- (kapena mefenoxam-) yomwe ili ndi fungicides m'munda. Mwachitsanzo, kusiyana kwakukulu pamlingo wotsutsa kunawonetsedwa mkati mwa mzere wa Sib1 clonal (Elansky et al., 2001).
Chifukwa chake, mtundu wa mating, peptidase isoenzyme sipekitiramu, mtundu wa DNA wa mitochondrial, RFLP-RG57, SSR ndizoyimira zomwe zimapangidwa popanga mabanki azidziwitso komanso kuyika zovuta m'magulu. Poyerekeza zitsanzo zochepa, ngati kuli koyenera kuyika mawonekedwe azolemba, mutha kugwiritsa ntchito AFLP, RAPD, InterSSR, Inter-SINE PCR (Gulu 5). Komabe, ziyenera kukumbukiridwa kuti njirazi sizingathe kuberekana, ndipo pakuyesa kwa aliyense payekha (amplification electrophoresis cycle), pamafunika zingapo zodzipatula.
Gulu 5. Kuyerekeza njira zosiyanasiyana zofufuzira zamavuto P. Ma infestans
muyezo | TS | Apolisi a Isofer | MtDNA | Gawo #: RFLP-RG57 | RAPD | ISSR | SSR | Mtengo wa AFLP | Rev |
---|---|---|---|---|---|---|---|---|---|
Zambiri zazidziwitso | Н | Н | Н | С | В | В | С | В | В |
Kubereka | В | В | В | В | Н | Н | С | С | С |
Kutheka kwa zinthu zakale | Н | Н | Н | Н | В | С | Н | С | В |
mtengo | Н | С | Н | В | Н | Н | Н | С | Н |
Ntchito mwamphamvu | Н | Н | Н | В | NS * | NS * | Н | С | NS * |
Liwiro lowunika ** | В | Н | Н | С | Н | Н | Н | Н | Н |
Chidziwitso: H - otsika, C - sing'anga, B - wokwera; НС * - mphamvu yogwira ntchito ndiyotsika mukamagwiritsa ntchito gel agarose kapena zokha
genotyper, sing'anga - ndi distillation mu polyacrylamide gel osakaniza zolembedwa zoyambira,
** - osawerengera nthawi yomwe mumagwiritsa ntchito kukulitsa mycelium yodzipatula kwa DNA.
Kapangidwe ka anthu
Mizere ya Clonal
Pakapanda kukumbukiranso kapena kuchepa kwake pakapangidwe ka anthu, kuchuluka kwa anthu kumakhala ndi ma clones angapo, kusinthana kwa majini pakati pawo ndikosowa kwambiri.
M'madera oterewa, ndizothandiza kwambiri kuti tisaphunzire kuchuluka kwa majini, koma mafupipafupi amitundu omwe amachokera chimodzimodzi (mizere yolumikizana kapena mizere yolumikizana) ndipo amasiyana kokha ndikusintha kwa mfundo. Kafukufuku wa kuchuluka kwa anthu omwe ali ndi kachilombo koyambitsa matendawa komanso kuwunika kwa mizere ya clonal kwachulukirachulukira kuyambira pomwe njira ya RFLP-RG57 kumayambiriro kwa zaka za m'ma 90 zapitazo. Pamodzi ndi RFLP-RG57, mtundu wa mating, mtundu wa peptidase ndi glucose-6-phosphate isomerase isoenzymes, ndi mtundu wa DNA wa mitochondrial amagwiritsidwa ntchito kuzindikira mizere yolumikizana. Makhalidwe azingwe zazitali kwambiri zowonetsedwa mu Gulu 6.
Clone US-1 inkalamulira anthu kulikonse mpaka kumapeto kwa ma 80, pambuyo pake idayamba kusinthidwa ndi ma clones ena ndikusowa ku Europe ndi North America. Tsopano ikupezeka ku Far East (Philippines, Taiwan, China, Japan, Korea, Koh et al., 1994, Mosa et al, 1993), ku Africa (Uganda, Kenya, Rwanda, Goodwin et al, 1994, Vega-Sanchez et al., 2000; Ochwo et al., 2002) ndi ku South America (Ecuador, Brazil, Peru, Forbes et al., 1997, Goodwin et al., 1994). Palibe zovuta za mzere wa US-1 zomwe zadziwika ku Australia kokha. Mwachiwonekere, P. infestans odzipatula adabwera ku Australia ndikusinthanso kwina (Goodwin, 1997).
Clone US-6 adasamukira kumpoto kwa Mexico kupita ku California kumapeto kwa ma 70 ndipo adayambitsa mliri mu mbatata ndi tomato pambuyo pazaka 32 zopanda matenda. Chifukwa chankhanza zake, idasamutsa chiwonetsero cha US-1 ndikuyamba kulamulira gombe lakumadzulo kwa United States (Goodwin et al., 1995a).
Ma genotypes US-7 ndi US-8 adapezeka ku United States mu 1992 ndipo kale ku 1994 adafalitsidwa ku United States ndi Canada. Pa nyengo imodzi yam'munda, choyerekeza cha US-8 chimatha kuthamangitsa kwathunthu US-1 m'minda ya mbatata yomwe idayambitsidwa ndimatombo onsewa mofanana (Miller ndi Johnson, 2000).
Clones BC-1 mpaka BC-4 adadziwika ku Britain Columbia mwa ochepa omwe adadzipatula ku Goodwin et al., 1995b). Clone US-11 imafalikira kwambiri ku United States ndikulanda US-1 ku Taiwan. Ma Clones JP-1 ndi EC-1, limodzi ndi choyerekeza cha US-1, ndizofala ku Japan ndi Ecuador, motsatana (Koh et al., 1994; Forbes et al., 1997).
SIB-1 ndi choyerekeza chomwe chidalipo ku Russia kudera lalikulu kuchokera kudera la Moscow kupita ku Sakhalin. M'chigawo cha Moscow, zidapezeka mu 1993, ndipo anthu ena m'minda amakhala makamaka ndi mitunduyi, yolimbana kwambiri ndi metalaxyl. Pambuyo pa 1993, kufalikira kwamtunduwu kunachepa kwambiri. Kunja kwa Urals mu 1997-1998, SIB-1 idapezeka paliponse, kupatula Khabarovsk Territory (choyerekeza SIB-2 chafalikira kumeneko). Kupatukana kwa ma clones okhala ndi mitundu yosiyanasiyana yokwatirana sikupatula njira zogonana ku Siberia ndi Far East. M'dera la Moscow, mosiyana ndi Siberia, anthu amaimiridwa ndi matanthwe ambiri; pafupifupi kudzipatula kulikonse kumakhala ndi genotype yapadera (Elansky et al., 2001, 2015). Zosiyanazi sizingafotokozedwe pongotengera kuitanitsa mitundu ya mafangasi ochokera kumadera osiyanasiyana padziko lapansi ndi mbewu zotumizidwa kunja. Popeza mitundu yonse iwiri ya kukwatirana imachitika mwaanthu, ndizotheka kuti kusiyanasiyana kwake kumapangidwanso chifukwa chokumbutsanso. Chifukwa chake, ku British Columbia, kutuluka kwa genotypes BC-2, BC-3, ndi BC-4 kumaganiziridwa chifukwa cha kusakanizidwa kwa ma clones BC-1 ndi US-6 (Goodwin et al., 1995b). N'zotheka kuti mitundu ya haibridi imapezeka mwa anthu aku Moscow. Mwachitsanzo, mavuto a MO-4, MO-8 ndi MO-11 heterozygous a malo a PEP atha kukhala hybrids pakati pamavuto MO-12, MO-21, MO-22, wokhala ndi mtundu wa A2 wophatikizika komanso wosakanikirana kamodzi kokha ka PEP locus ndi mavuto MO-8, yokhala ndi mtundu wokhathamira wa A1 komanso wosakanikirana ndi ena mwanjira ina. Ndipo ngati ndi choncho, ndipo mwa anthu amakono a P. infestans pali chizolowezi chowonjezeka pantchito yogonana, ndiye kuti phindu lazidziwitso pakuwunika kwamitundu yambiri lidzachepa (Elansky et al., 2001, 2015).
Kusiyanasiyana kwa mizere yama clonal
Mpaka zaka za m'ma 90 za m'zaka za zana la 20, mzere wachitsulo US-1 unali wofala padziko lapansi. Ambiri mwa magawo ndi zigawo zimakhala ndi mitundu yokhayokha ndi mtundu wa US-1. Komabe, kusiyana pakati pa odzipatula kunawonekeranso, makamaka chifukwa cha kusintha kwa zinthu. Masinthidwe adachitika mu nyukiliya komanso mitochondrial DNA ndipo adakhudza, mwa zina, mulingo wokana mankhwala a phenylamide komanso kuchuluka kwa majeremusi oyipa. Mizere yomwe imasiyana ndi ma genotypes oyambilira mwa kusintha kwa masinthidwe amawonetsedwa ndi manambala owonjezera pambuyo pa kadontho kamene kamatsatira dzina la genotype yoyambirira (mwachitsanzo, mzere wa US-1.1 wosinthika wa mzere wa US-1). Mizere yosindikiza zala ya DNA US-1.5 ndi US-1.6 ili ndi mizere yolumikizira yamitundu yosiyanasiyana (Goodwin et al., 1995a, 1995b); mzere wa clonal US-6.3 umasiyananso ndi US-6 ndi chingwe chimodzi chowonjezera (Goodwin, 1997, Gulu 7).
Pofufuza za mitochondrial DNA, zidapezeka kuti mu clonal mzere US-1 kokha mtundu 1b wa mitochondrial DNA umapezeka (Carter et al., 1990). Komabe, pofufuza zamagulu amtunduwu ochokera ku Peru ndi Philippines, odzipatula adapezeka omwe mitundu ya DNA ya mitochondrial inali yosiyana ndi 1b pakupezeka ndikulowetsa (Goodwin, 1991, Koh et al., 1994).
Tebulo 6. Mitundu yambiri yamagulu angapo ya P. infestans clonal lines
Mutu | Mtundu wa mating | Maselo | Zolemba za DNA | Mtundu wa MtDNA | |
GPI | PEP | ||||
US-1 | A1 | 86/100 | 92/100 | 1.0111010110011E + 24 | Ib |
US-2 | A1 | 86/100 | 92/100 | 1.0111010010011E + 24 | - |
US-3 | A1 | 86/100 | 92/100 | 1.0111000000011E + 24 | - |
US-4 | A1 | 100/100 | 92/92 | 1.0111010010011E + 24 | - |
US-5 | A1 | 100/100 | 92/100 | 1.0111010010011E + 24 | - |
US-6 | A1 | 100/100 | 92/100 | 1.0111110010011E + 24 | IIb |
US-7 | A2 | 100/111 | 100/100 | 1.0011000010011E + 24 | Ia |
US-8 | A2 | 100/111/122 | 100/100 | 1.0011000010011E + 24 | Ia |
US-9 | A1 | 100/100 | 83/100 | * | - |
US-10 | A2 | 111/122 | 100/100 | - | - |
US-11 | A1 | 100/111 | 92/100 | 1.0101110010011E + 24 | IIb |
US-12 | A1 | 100/111 | 92/100 | 1.0001000010011E + 24 | - |
US-14 | A2 | 100/122 | 100/100 | 1.0000000000011E + 24 | - |
US-15 | A2 | 100/100 | 92/100 | 1.0001000010011E + 24 | Ia |
US-16 | A1 | 100/111 | 100/100 | 1.0001100010011E + 24 | - |
US-17 | A1 | 100/122 | 100/100 | 1.0100010000011E + 24 | - |
US-18 | A2 | 100/100 | 92/100 | 1.0001000010011E + 24 | Ia |
US-19 | A2 | 100/100 | 92/100 | 1.0101010000011E + 24 | Ia |
Mtengo wa EC-1 | A1 | 90/100 | 96/100 | 1.1111010010011E + 24 | IIa |
SIB-1 | A1 | 100/100 | 100/100 | 1.0001000110011E + 24 | IIa |
SIB-2 | A2 | 100/100 | 100/100 | 1.0001000010011E + 24 | IIa |
SIB-3 | A1 | 100/100 | 100/100 | 1.1001010100011E + 24 | IIa |
MO-1 | A2 | 100/100 | 100/100 | 1.0001000110011E + 24 | IIa |
MO-2 | A2 | 100/100 | 100/100 | 1.0001000010011E + 24 | Ia |
MO-3 | A1 | 100/100 | 100/100 | 1.0101000010011E + 24 | IIa |
MO-4 | A1 | 100/100 | 92/100 | 1.0101110110011E + 24 | IIa |
MO-5 | A1 | 100/100 | 100/100 | 1.0001010010011E + 24 | IIa |
MO-6 | A1 | 100/100 | 100/100 | 1.0101010010011E + 24 | Ia |
MO-7 | A1 | 100/100 | 92/100 | 1.0001000110011E + 24 | IIa |
MO-8 | A1 | 100/100 | 92/92 | 1.0101100010011E + 24 | IIa |
MO-9 | A1 | 100/100 | 92/100 | 1.0001000010011E + 24 | IIa |
MO-10 | A1 | 100/100 | 100/100 | 1.0101100000011E + 24 | Ia |
MO-11 | A1 | 100/100 | 92/100 | 1.0101010010011E + 24 | Ia |
MO-12 | A2 | 100/100 | 100/100 | 1.0101010010011E + 24 | Ia |
MO-13 | A1 | 100/100 | 100/100 | 1.0101010000011E + 24 | Ia |
MO-14 | A1 | 100/100 | 100/100 | 1.01010010011E + 22 | Ia |
MO-15 | A1 | 100/100 | 100/100 | 1.101110010011E + 23 | Ia |
MO-16 | A1 | 100/100 | 100/100 | 1.0001000000011E + 24 | IIa |
MO-17 | A1 | 86/100 | 100/100 | 1.0101010110011E + 24 | Ib |
MO-18 | A1 | 100/100 | 100/100 | 1.0101110010011E + 24 | IIa |
MO-19 | A1 | 100/100 | 100/100 | 1.0101010000011E + 24 | IIa |
MO-20 | A2 | 100/100 | 100/100 | 1.0101010000011E + 24 | IIa |
MO-21 | A2 | 100/100 | 100/100 | 1.0101010000011E + 24 | IIa |
Chidziwitso: * - palibe data.
Gulu 7. Multilocus genotypes ndi mizere yawo yosinthika
Mutu | Mtundu wa mating | | Zolemba za DNA (RG57) | Mfundo | |
GPI | PEP-1 | ||||
US-1 | A1 | 86/100 | 92/100 | 1011101011001101000110011 | Chibadwa choyambirira 1 |
US-1.1 | A1 | 86/100 | 100/100 | 1011101011001101000110011 | Kusintha kwa PEP |
US-1.2 | A1 | 86/100 | 92/100 | 1011101010001101000110011 | Kusintha kwa RG57 |
US-1.3 | A1 | 86/100 | 92/100 | 1011101001001101000110011 | Kusintha kwa RG57 |
US-1.4 | A1 | 86/100 | 100/100 | 1011101010001101000110011 | Kusintha kwa RG57 ndi PEP |
US-1.5 | A1 | 86/100 | 92/100 | 1011101011001101010110011 | Kusintha kwa RG57 |
US-6 | A1 | 100/100 | 92/100 | 1011111001001100010110011 | Chibadwa choyambirira 2 |
US-6.1 | A1 | 100/100 | 92 /92 | 1011111001001100010110011 | Kusintha kwa PEP |
US-6.2 | A1 | 100/100 | 92/100 | 1011101001001100010110011 | Kusintha kwa RG57 |
US-6.3 | A1 | 100/100 | 92/100 | 1011111001011100010110011 | Kusintha kwa RG57 |
US-6.4 | A1 | 100/100 | 100/100 | 1011011001001100010110011 | Kusintha kwa RG57 ndi PEP |
US-6.5 | A1 | 100/100 | 92/100 | 1011111001001100010010011 | Kusintha kwa RG57 |
BR-1 | A2 | 100/100 | 100/100 | 1011101000001100001111011 | Chibadwa choyambirira 3 |
BR-1.1 | A2 | 100/100 | 100/100 | 1010101000001100001110011 | Kusintha kwa RG57 |
Palinso kusintha kwamitundu isozymes. Monga lamulo, zimayambitsidwa ndi kuwonongeka kwa chamoyo choyambirira heterozygous cha enzyme iyi kukhala yofanana. Mu 1993, pa zipatso za phwetekere, tidazindikira zovuta zomwe zimakhala ndi US-1: RG57 zolemba zala, mtundu wa mitochondrial DNA, ndi 86/100 genotype ya glucose-6-phosphati-isomerase, koma inali homozygous (100/100) ya peptidase locus yoyamba m'malo mwa heterozygote ya 92/100 yofanana ndi mzerewu. Tidatchula mtundu wamtunduwu MO-17 (Gulu 6). Mizere yosinthika US-1.1 ndi US-1.4 imasiyananso ndi US-1 pakusintha koyamba kwa peptidase locus (Gulu 7).
Masinthidwe obweretsa kusintha kwa kuchuluka kwa majeremusi a virulence a mitundu ya mbatata ndi tomato ndiofala. Adadziwika pakati paokhaokha a clonal line US-1 mwa anthu ochokera ku Netherlands (Drenth et al., 1994), Peru (Goodwin et al., 1995a), Poland (Sujkowski et al., 1991), kumpoto kwa North America (Goodwin et al., ., 1995b). Kusiyana kwa kuchuluka kwa majeremusi amtundu wa mbatata kunadziwikanso pakati pamagawo ena a US-7 ndi US-8 ku Canada ndi United States (Goodwin et al., 1995a), pakati pa mzere wokhawo wa SIB-1 ku Asia gawo la Russia (Elansky Et al, 2001 ).
Madera akutali omwe ali ndi magawano amtundu wa kukana kwa mankhwala a phenylamide adadziwika m'magulu am'modzi, onse omwe anali a cl-line Sib-1 (Elansky et al, 2001, Gulu 1). Pafupifupi mitundu yonse ya clonal line US-1 imatha kutengeka ndi metalaxyl; komabe, osagwirizana kwambiri pamzerewu adadzipatula ku Philippines (Koh et al., 1994) ndi ku Ireland (Goodwin et al., 1996).
Anthu amakono a P. infestans
Central America (Mexico)
Chiwerengero cha P. infestans ku Mexico chimasiyana mosiyana ndi anthu ena padziko lapansi, makamaka chifukwa cha mbiri yake. Kafukufuku wambiri wa anthuwa ndi mitundu yokhudzana ndi P. infestans ya Clade Phytophthora, komanso mitundu yakomweko ya mtundu wa Solanum, zidapangitsa kuti pakhale lingaliro loti kusintha kwa tizilombo toyambitsa matenda m'chigawo chapakati cha Mexico kudachitika limodzi ndi kusinthika kwa mbewu zomwe zimasungidwa ndipo zimalumikizidwa ndikupanganso kugonana (Grünwald, Flier , 2005). Mitundu yonse iwiri ya mating imayimilidwa mwa anthu, mofanana, komanso kupezeka kwa ma oospores m'nthaka, pazomera ndi ma tubers a mbatata ndi mitundu yokhudzana ndi kukula kwamtchire Solanum imatsimikizira kupezeka kwa njira yogonana pakati pa anthu (Fernández-Pavía et al., 2002). Kafukufuku waposachedwa wa Chigwa cha Toluca ndi madera ake (malo oyambilira a tizilombo toyambitsa matenda) adatsimikizira kusiyanasiyana kwamitundu ya P. infestans (134 multilocus genotypes mu zitsanzo za 176) komanso kupezeka kwa magulu angapo osiyana siyana m'derali (Wang et al., 2017). Zomwe zimapangitsa kuti pakhale kusiyana kumeneku ndi kugawidwa kwa madera omwe ali kumtunda kwa Mexico, kusiyanasiyana kwamomwe kulimidwa ndi mitundu ya mbatata yomwe imagwiritsidwa ntchito zigwa ndi mapiri, komanso kupezeka kwa mitundu yamtundu wa Solanum yamtchire yomwe ingakhale ngati magulu ena (Fry et al ., 2009).
Komabe, ziyenera kudziwika kuti anthu a P. infestans kumpoto kwa Mexico ndiwosokonekera ndipo amafanana kwambiri ndi anthu aku North America, omwe atha kuwonetsa kuti awa ndi majeremusi atsopano (Fry et al., 2009).
North America
Anthu aku North America a P. infestans nthawi zonse amakhala ndi mawonekedwe osavuta ndipo mawonekedwe awo anali atakhazikitsidwa kale kusanachitike kusanthula kwa microsatellite. Mpaka 1987, mzere wa clonal US-1 unkalamulira ku United States ndi Canada (Goodwin et al., 1995). Pakatikati mwa ma 70, pomwe fungicides yokhudzana ndi metalaxyl idawonekera, mawonekedwe awa adayamba kusinthidwa ndi mitundu ina, yolimbana nayo yomwe idasamukira ku Mexico (Goodwin et al., 1998). Pakutha kwa zaka za m'ma 90. genotype ya US-8 idasinthiratu mtundu wa US-1 ku United States ndipo idakhala mzere wolimba kwambiri wa mbatata (Fry et al., 2009; Fry et al., 2015). Zinthu zinali zosiyana ndi tomato, omwe amakhala ndi mizere ingapo yama clonal, ndipo kapangidwe kake kamasintha chaka ndi chaka (Fry et al., 2009).
Mu 2009, mliri waukulu wa matenda oopsa mochedwa udabuka ku United States pa tomato. Chowopsa cha mliriwu chidayamba pafupifupi nthawi imodzi m'malo ambiri kumpoto chakum'mawa kwa United States, ndipo zidapezeka kuti zimagwirizana ndi kugulitsa kwakukulu kwa mbande za phwetekere m'minda yayikulu yaminda (Fry et al., 2013). Kuwonongeka kwa mbewu kunali kwakukulu. Kusanthula kwa microsatellite kwamitundu yomwe yakhudzidwa kunawonetsa kuti vuto la mliriwo linali la clonal line US-22 A2-mtundu wa mating. Mu 2009, gawo la genotype iyi mwa anthu aku America a P. infestans lidafika 80% (Fry et al., 2013). M'zaka zotsatira, kuchuluka kwa ma genotypes ankhanza US-23 (makamaka tomato) ndi US-24 (pa mbatata) kudakulirakulira m'chiwerengero cha anthu, komabe, pambuyo pa 2011, kuchuluka kwa US-24 kunachepa kwambiri, ndipo mpaka pano, pafupifupi 90% ya anthu omwe ali ndi tizilomboto United States imayimilidwa ndi genotype ya US-23 (Fry et al., 2015).
Ku Canada, monga ku United States, kumapeto kwa zaka za m'ma 90. mtundu waukulu wa US-1 udalowedwa m'malo ndi US-8, udindo womwe sunasinthe mpaka 2008. Ku Canada, kudali miliri yoopsa yoyambitsidwa ndi kugulitsa mbande za phwetekere, koma zidayambitsidwa ndi genotypes US-2009 ndi US-2010 (Kalischuk et al., 23). Kusiyanitsa komveka bwino kwa mitundu iyi kunali kodabwitsa: US-8 idalamulira zigawo zakumadzulo za Canada (2012%), pomwe US-23 idalamulira zigawo zakummawa (68%). M'zaka zotsatira, US-8 idafalikira kumadera akum'mawa; komabe, kwakukulu, gawo lake mwa anthu lidatsika pang'ono motsutsana ndi mawonekedwe amitundu ya US-83 ndi US-23 mdziko muno (Peters et al., 22). Mpaka pano, US-24 ili ndiudindo waukulu ku Canada; US-2014 ilipo ku British Columbia, pomwe US-23 ndi US-8 amapezeka ku Ontario (Peters, 23).
Chifukwa chake, anthu aku North America a P. infestans amakhala mizere yambiri. Pazaka 40 zapitazi, kuchuluka kwa ma genotypes omwe amapezeka ndi clonal afikira 24. Ngakhale kuti mitundu yonse ya mating ikupezeka mwa anthu, mwayi wopezeka ndi majeremusi atsopano chifukwa chofunanso kugonana udakali wotsika kwambiri. Komabe, mzaka 20 zapitazi, milandu ingapo yakuwonekera kwa anthu omwe amapezeka mobwerezabwereza yalembedwa (Gavino et al., 2000; Danies et al., 2014; Peters et al., 2014), ndipo nthawi ina, zotsatira zakuwoloka zinali genotype US-11 , yomwe idakhazikitsidwa ku North America kwazaka zambiri (Gavino et al., 2000). Mpaka 2009, kusintha kwa kapangidwe ka anthu kumalumikizidwa ndi kutuluka kwamitundu yatsopano, yowopsa kwambiri ndikusamuka kwawo ndikusunthidwa kwa omwe adalipo kale. Zomwe zidachitika mu 2009-2010 ku USA ndi Canada, ma epiphytotic kwa nthawi yoyamba adawonetsa kuti munthawi ya kudalirana, kufalikira kwa matendawa kumatha kuphatikizidwa ndi kufalikira kwa mitundu yatsopano ya genotypes pogulitsa zinthu zobzala zomwe zili ndi kachilombo.
South America
Mpaka posachedwa, kafukufuku wa anthu ku South America a P. infestans sanali wamba kapena wokulirapo. Amadziwika kuti kapangidwe ka anthuwa ndiosavuta ndipo amaphatikizapo mizere 1-5 ya clonal m'dziko lililonse (Forbes et al., 1998). Chifukwa chake, pofika 1998, ma genotypes US-1 (Brazil, Chile) BR-1 (Brazil, Bolivia, Uruguay, Paraguay), EC-1 (Ecuador, Colombia, Peru ndi Venezuela), AR-1, AR -2, AR-3, AR-4 ndi AR-5 (Argentina), PE-3 ndi PE-7 (kumwera kwa Peru). Mtundu wokwatirana wa A2 udalipo ku Brazil, Bolivia ndi Argentina ndipo sunapezeke kupitirira malire a Bolivia ndi Peruvia mdera la Lake Titicaca, kumbuyo komwe mtundu wa EC-1 A1 unkalamulira ku Andes. Pa tomato, US-1 idakhalabe yotchuka ku South America.
Zinthu zidapitilirabe m'ma 2000. Chofunikira chinali kupezeka kwa mzere watsopano wa clonal EC-2 wamtundu wa A2 pamtundu wachibale wa mbatata (S. brevifolium ndi S. tetrapetalum) ku Andes Kumpoto (Oliva et al., 2010). Kafukufuku wa Phylogenetic awonetsa kuti mzerewu siwofanana kwathunthu ndi P. infestans, ngakhale ndiwofanana kwambiri, mokhudzana ndi zomwe adapatsidwa kuti aganizirepo, komanso mzere wina, EC-3, womwe umasiyana ndi mtengo wa phwetekere S. betaceum wokula ku Andes, mtundu watsopano wotchedwa P. andina; komabe, udindo wa mtundu uwu (mtundu wodziyimira pawokha kapena wosakanizidwa wa P. infestans wokhala ndi mzere wina wosadziwika) sunadziwikebe (Delgado et al., 2013).
Pakadali pano, anthu onse aku South America a P. infestans ndi clonal. Ngakhale kupezeka kwamitundu yonse iwiri ya mating, palibe anthu ophatikizananso omwe adadziwika. Pa tomato, mtundu wa US-1 umapezeka paliponse, mwachiwonekere wachoka mbatata ndi mitundu yakomweko, komwe kwenikweni sikudziwika. Ku Brazil, Bolivia ndi Uruguay, mtundu wa BR-1 ulipo; ku Peru, pamodzi ndi US-1 ndi EC-1, pali mitundu ina yambiri yakomweko. Ku Andes, udindo waukulu umasungidwa ndi clonal line EC-1, ubale womwe ulipo ndi P. andina wodziwika posachedwa sunadziwike. Malo okhawo "osakhazikika" komwe kwa nthawi ya 2003-2013. panali kusintha kwakukulu pakati pa anthu, adakhala Chile (Acuña et al., 2012), komwe mu 2004-2005. kuchuluka kwa tizilombo toyambitsa matenda kunadziwika chifukwa chokana metalaxyl ndi mitochondrial DNA haplotype (Ia m'malo mwa Ib). 2006 mpaka 2011 Pakati pa anthu, genotype 21 (malinga ndi SSR) idalamulira, gawo lomwe lidafika 90%, kenako kanjedza chidadutsa genotype 20, zomwe zimachitika zaka ziwiri zotsatira zidasungidwa pafupifupi 67% (Acuña, 2015).
Europe
M'mbiri ya Europe, pakhala pali mafunde osachepera awiri osamuka a P. infestans ochokera ku North America: m'zaka za zana la 1. (HERB-1) komanso koyambirira kwa zaka za 70th (US-1). Kufalikira kulikonse kwa mankhwala a fungus a metalaxyl m'ma XNUMXs. zidatsogolera kusamutsidwa kwa mtundu wapamwamba wa US-XNUMX ndikusinthidwa ndi ma genotypes atsopano. Zotsatira zake, m'maiko ambiri akumadzulo kwa Europe, kuchuluka kwa tizilomboti tidayimilidwa makamaka ndi mizere ingapo yama clonal.
Kugwiritsa ntchito microsatellite kusanthula kwa kuchuluka kwa tizilomboto kunapangitsa kuti kuwulule zosintha zazikulu zomwe zidachitika ku Western Europe mu 2005-2008. Mu 2005, chingwe chatsopano chinapezeka ku UK, chotchedwa 13_A2 (kapena "Blue 13") ndipo chodziwika ndi mtundu wa A2 wokhwima , kukwiya kwambiri komanso kukana kwa phenylamides (Shaw et al., 2007). Genotype yomweyi idapezeka m'mitundu yomwe idasonkhanitsidwa ku 2004 ku Netherlands ndi kumpoto kwa France, yomwe idati idasamukira ku UK kuchokera ku Continental Europe, mwina ndi mbatata za mbewu (Cooke et al., 2007). Kafukufuku wamtundu wa oimira mzerewu adawonetsa kuchuluka kwa ma polymorphism motsatizana kwake (pofika 2016, kuchuluka kwa kusiyanasiyana kwake kudafika 340) komanso kusiyanasiyana kwakukulu pamlingo wamitundu, kuphatikiza. zamoyo zomwe zimayambitsa matendawa (Cooke et al., 2012; Cooke, 2017). Izi, kuphatikizapo kuchuluka kwa gawo la biotrophic gawo, zitha kuyambitsa kukwiya kwa 13_A2 ndikutha kwake kupatsira mitundu ya mbatata yolimbana ndi vuto lakumapeto.
M'zaka zingapo zotsatira, genotype idafalikira mwachangu m'maiko aku Northwestern Europe (Great Britain, Ireland, France, Belgium, Netherlands, Germany) ndikuchotsa munthawi yomweyo mitundu yayikulu kwambiri 1_A1, 2_A1, 8_A1 (Montarry et al., 2010; Gisi et al. , 2011; Van den Bosch et al., 2011; Cooke, 2015; Cooke, 2017). Malinga ndi tsamba la webusayiti la www.euroblight.net, gawo la 13_A2 mwa anthu m'mayikowa lidafika 60-80% ndi ena; kupezeka kwa genotype iyi kwalembedwanso m'maiko ena akum'mawa ndi kumwera kwa Europe. Komabe, mu 2009-2012. 13_A2 idataya maudindo akuluakulu ku Great Britain ndi France, kutsatira 6_A1 mzere (8_A1 ku Ireland), ndipo ku Netherlands ndi Belgium idasinthidwa pang'ono ndi mitundu 1_A1, 6_A1, ndi 33_A2 (Cooke et al., 2012; Cooke, 2017; Stellingwerf, 2017).
Pakadali pano, pafupifupi 70% ya anthu aku Western Europe a P. infestans ndi amodzi. Malinga ndi tsamba la webusayiti la www.euroblight.net, mitundu yayikulu kwambiri m'maiko aku North-West Europe (UK, France,
Netherlands, Belgium) zidatsalira, pafupifupi mofanana, 13_A2 ndi 6_A1, ndipo zomalizirazi sizimachitika kunja kwa dera lotchulidwa (kupatula Ireland), koma zili ndi ma subclone osachepera 58 (Cooke, 2017). Kusiyanasiyana kwa 13_A2 kumapezeka mu Germany, ndipo kumaonekanso mwa apo ndi apo m'maiko a Central ndi Southern Europe. Genotype 1_A1 ndi gawo lalikulu la anthu ku Belgium ndipo pang'ono ndi pang'ono Netherlands ndi France. Genotype 8_A1 yakhazikika pakati pa anthu aku Europe pamlingo wa 3-6%, kupatula Ireland, komwe imapitilizabe kutsogola ndipo imagawidwa m'magulu awiri (Stellingwerf, 2017). Pomaliza, mu 2016, panali kuwonjezeka kwakuchuluka kwakanthawi kwamitundu yatsopano 36_A2 ndi 37_A2, yoyamba kulembedwa mu 2013-2014; mpaka pano, mitundu iyi imapezeka ku Netherlands ndi Belgium ndipo mwina ku France ndi Germany, komanso kumwera kwa Great Britain (Cooke, 2017). Pafupifupi 20-30% ya anthu aku Western Europe amayimilidwa ndi mitundu yosiyanasiyana yapachibale chaka chilichonse.
Mosiyana ndi Western Europe, pofika nthawi yomwe 13_A2 genotype idawonekera, anthu aku Northern Europe (Sweden, Norway, Denmark, Finland) sanayimilidwe ndi mizere, koma ndi mitundu yambiri yapadera (Brurberg et al.,
2011). Munthawi ya kufalikira kwa 13_A2 ku Western Europe, kupezeka kwa mtundu uwu ku Scandinavia sikunadziwike mpaka 2011, pomwe udapezeka koyamba ku North Jutland (Denmark), komwe mitundu yambiri ya mbatata imabzalidwa pogwiritsa ntchito metalaxyl fungicides (Nielsen et al., 2014). Malinga ndi www.euroblight.net, genotype 13_A2 idapezekanso m'mitundu ingapo kuchokera ku Norway ndi Denmark mu 2014 komanso m'mitundu ingapo yaku Norway ku 2016; Kuphatikiza apo, mu 2013, kupezeka kwa genotype 6_A1 pang'ono kunadziwika ku Finland. Chifukwa chachikulu cholephera kwa 13_A2 ndi mizere ina yolanda ku Scandinavia akuti ndiko kusiyana kwanyengo mdera lino kuchokera kumayiko a Western Europe.
Kuphatikiza pa kuti nyengo yotentha ndi nyengo yozizira imathandizira kupulumuka kwa ma oospores m'malo mwa vegetative mycelium (Sjöholm et al., 2013), nthaka yozizira nthawi yachisanu (yomwe nthawi zambiri sichimachitika m'maiko otentha a Western Europe) imathandizira kulumikizana kwa ma oospores kumera ndi kubzala. mbatata, yomwe imathandizira gawo lawo ngati gwero la matenda oyamba (Brurberg et al., 2011). Tiyeneranso kukumbukira kuti, kumpoto, kukula kwa matenda opitilira muyeso kukupitilira kukula kwa matenda opatsirana, omwe pamapeto pake amalepheretsa kulamulira kwamphamvu kwambiri, koma pambuyo pake adapanga mizere yama clonal (Yuen, 2012). Kapangidwe ka anthu omwe amaphunzira kwambiri a P. infestans ku Eastern Europe (Poland, Baltic States) ndi ofanana kwambiri ndi aku Scandinavia.
Mitundu yonse iwiri ya mating ilipo pano, ndipo mitundu yambiri yamitundu yomwe imatsimikiziridwa ndi kusanthula kwa SSR ndiyapadera (Chmielarz et al., 2014; Runno-Paurson et al., 2016). Monga ku Northern Europe, kufalikira kwa mizere yama clonal (makamaka 13_A2 genotype) sikunakhudze kuchuluka kwa tizilombo toyambitsa matenda, komwe kumakhalabe kosiyanasiyana mosiyanasiyana pakalibe mizere yayikulu.
Kukhalapo kwa 13_A2 nthawi zina kumawonedwa m'minda yokhala ndi mitundu ya mbatata. Ku Russia, izi zikuchitikanso chimodzimodzi. Kusanthula kwama Microsatellite a P. infestans amadzipatula omwe adasonkhanitsidwa mu 2008-2011 M'magawo 10 osiyanasiyana mdera la Europe ku Russia, adawonetsa kusiyanasiyana kwamitundu yosiyanasiyana komanso kusowa kwazinthu zofanana ndi mizere yaku Europe (Statsyuk et al., 2014). Zaka zingapo pambuyo pake, kafukufuku wazitsanzo za P. infestans omwe adatoleredwa mdera la Leningrad mu 2013-2014 adawonetsa kusiyana kwakukulu pakati pawo ndi ma genotypes ochokera mdera lino omwe atchulidwa m'mbuyomu. M'maphunziro onsewa, ma genotypes aku Western Europe sanapezeke (Beketova et al., 2014; Kuznetsova et al., 2016).
Mitundu yambiri yamitundu yakum'mawa kwa Europe ya P. infestans komanso kusowa kwa mizere yolimba mwa iwo kungakhale chifukwa cha zifukwa zingapo. Choyamba, monga Kumpoto kwa Europe, nyengo yam'mayiko omwe akuwerengedwa imathandizira pakupanga ma oospores ngati gwero lalikulu la matenda (Ulanova et al., 2010; Chmielarz et al., 2014). Chachiwiri, mbatata zochuluka zomwe zimapangidwa m'maiko amenewa zimalimidwa m'minda ing'onoing'ono yabizinesi, yomwe nthawi zambiri imazunguliridwa ndi nkhalango kapena zina zopinga kuyenda kwaulere kwa zinthu zopatsirana (Chmielarz et al., 2014). Monga lamulo, mbatata zomwe zimakula munthawi zotere sizimachiritsidwa ndi mankhwala, ndipo kusankha mitundu kumakhazikitsidwa chifukwa cha kukana kwawo mochedwa, i.e. palibe kukakamizidwa kosankha kukwiya komanso kukana metalaxyl, yomwe imachotsa ma genotypes osagwirizana, monga 13_A2, maubwino kuposa ma genotypes ena (Chmielarz et al., 2014). Pomaliza, chifukwa chakuchepa kwa malo, eni ake nthawi zambiri samazungulira mbewuzo, kulima mbatata pamalo omwewo kwa zaka zambiri, zomwe zimapangitsa kuti pakhale kuchuluka kwa inoculum yamitundu yosiyanasiyana (Runno-Paurson et al., 2016; Elansky, 2015; Elansky et al. ., 2015).
Asia
Mpaka posachedwa, kapangidwe ka P. infestans ku Asia sikamamvetsetseka bwino. Zimadziwika kuti imayimilidwa makamaka ndi mizere yama clonal, ndipo zomwe zimachitika pakukonzanso zachiwerewere kutuluka kwa mitundu yatsopano ndizochepa kwambiri. Mwachitsanzo, mu 1997-1998. M'chigawo cha Asia ku Russia (Siberia ndi Far East), tizilombo toyambitsa matenda tidayimiridwa ndi mitundu itatu yokha yomwe ili ndi mbiri ya SIB-1 genotype (Elansky et al., 2001). Kupezeka kwa mizere yama clonal pathogen kwawonetsedwa m'maiko monga China, Japan, Korea, Philippines, ndi Taiwan (Koh et al., 1994; Chen et al., 2009). Mzere wachitsulo US-1, womwe unkalamulira gawo lalikulu la Asia, kumapeto kwa zaka za m'ma 90 - koyambirira kwa 2000. pafupifupi kulikonse kunayamba kusinthidwa ndi ma genotypes ena, omwe nawonso, adalowa m'malo mwa atsopano. Nthawi zambiri, kusintha kwamapangidwe ndi kapangidwe ka anthu m'maiko aku Asia kumalumikizidwa ndi kusamuka kwamitundu yatsopano kuchokera kunja. Chifukwa chake, ku Japan, kupatula JP-3 genotype, mitundu ina yonse yaku Japan yomwe idabwera pambuyo pa US-1 (JP-1, JP-2, JP-3) ili ndi mbiri yakunja yotsimikizika (Akino et al., 2011) ... Pakadali pano pali mitundu itatu yayikulu ya tizilomboto ku China, yogawika bwino; Palibe kapena kufooka kwa jini pakati pa anthuwa (Guo et al., 2010; Li et al., 2013b). Genotype 13_A2 idawonekera kudera la China m'maiko ake akumwera (Yunnan ndi Sichuan) mu 2005-2007, ndipo mu 2012-1014. idawonekanso kumpoto chakum'mawa kwa dzikolo (Li et al., 2013b). Ku India, 13_A2 imawoneka nthawi imodzi ndi ku China, makamaka ndi mbatata zambewu (Chowdappa et al., 2015), komanso mu 2009-2010. idadzetsa matenda oopsa a epiphytotic oopsa mochedwa pa phwetekere kumwera kwa dzikolo, pambuyo pake idafalikira ku mbatata ndipo mu 2014 idayambitsa kuphulika kwa ngozi mochedwa ku West Bengal, zomwe zidapangitsa kuti alimi ambiri akumaloko awonongeke (Fry, 2016).
Africa
Mpaka 2008-2010 kafukufuku wokhazikika wa P. infestans m'maiko aku Africa sanachitike. Pakadali pano, anthu aku Africa a P. infestans atha kugawidwa m'magulu awiri, ndipo gawoli limalumikizidwa bwino ndikuti kubwera kwa mbatata kuchokera ku Europe.
Kumpoto kwa Africa, komwe kumatumiza mbatata kuchokera ku Europe, mtundu wa A2 wokhwima umayimiriridwa pafupifupi zigawo zonse, zomwe zimapereka lingaliro la kuthekera kwa kutuluka kwa mitundu yatsopano yamtunduwu chifukwa chokhazikitsanso kugonana (Corbière et al., 2010; Rekad et al., 2017). Kuphatikiza apo, ku Algeria, kupezeka kwa genotypes 13_A2, 2_A1, ndi 23_A1 kumadziwika ndikulamulira koyambirira kwa iwo, komanso kuchepa pang'ono pang'onopang'ono kwamitundu yapadera yothetsera kutha (Rekad et al., 2017). Mosiyana ndi dera lonselo, ku Tunisia (kupatula kumpoto chakum'mawa kwa dzikolo), tizilombo toyambitsa matenda timayimilidwa makamaka ndi mtundu wa mating A1 (Harbaoui et al., 2014).
Mzere wama clonal NA-01 ndiwowonekera apa. Mwambiri, kuchuluka kwa mizere yama clonal mwa anthu ndi 43% yokha. Kum'mawa ndi Kummwera kwa Africa, komwe kuchuluka kwa mbewu zomwe zimagulitsidwa kunja ndizochepa kwambiri (Fry et al., 2009), P. infestans imayimilidwa ndi mizere iwiri yokha yamtundu wa A1, US-1 ndi KE-1, ndipo yomalizirayi imachotsapo mbatata ( Pule et al., 2012; Njoroge et al., 2016). Mpaka pano, mitundu iwiri yonseyi ili ndi mitundu yambiri yazosiyanasiyana.
Australia
Lipoti loyamba lakuchepa kwa mbatata ku Australia lidayamba mchaka cha 1907, ndipo epiphytotia yoyamba, yomwe mwina idayamba chifukwa cha mvula yambiri m'nyengo yachilimwe, idachitika mu 1909-1911. (Drenth et al., 2002). Mwambiri, komabe, kuwonongeka mochedwa kulibe tanthauzo lililonse pachuma. Kuphulika kwakanthawi kwakanthawi kochepa, komwe kumayambitsidwa ndi nyengo yomwe imapereka chinyezi chambiri, kumachitika kangapo kamodzi pazaka 5-7 zilizonse ndipo zimapezeka makamaka kumpoto kwa Tasmania ndi central Victoria. Pokhudzana ndi zomwe zatchulidwazi, zofalitsa zomwe zidafufuza momwe anthu aku Australia a P. infestans zilibe. Zomwe zilipo posachedwa ndikuchokera 1998-2000. (Drenth et al., 2002). Malinga ndi olembawo, kuchuluka kwa anthu aku Victoria kudali mzere waku US-1.3, womwe umatsimikizira mosapita m'mbali kusamuka kwa mtunduwu kuchokera ku United States. Zoyimira za Tasmanian zidasankhidwa kukhala AU-3, zosiyana ndi ma genotypes omwe analipo nthawi imeneyo kumadera ena adziko lapansi.
Zida zakukula kwa ngozi yaku Russia
Ku Europe, matenda omwe amabwera ndi matenda a tubers a mbewu, ma oospores omwe adadutsa munthaka, komanso zoosporangia zomwe zimabwera ndi mphepo yochokera ku mbewu zomwe zimamera kuchokera ku masamba omwe adasandulika m'minda yatha chaka chatha ("zodzipereka"), kapena milu yazomera Bookmark posungira tubers. Mwa izi, mbewu zomwe zimakula pamulu wa tubers zomwe zatayidwa zimawerengedwa kuti ndi gwero loopsa kwambiri la matenda. pamenepo, kuchuluka kwa ma tubers ophuka nthawi zambiri kumakhala kofunikira, ndipo zoosporangia itha kunyamulidwa kuchokera ku iwo pamtunda wautali. Zina zonse (oospores, "odzipereka" zomera) sizowopsa, chifukwa sizolowera kubzala mbewu m'minda imodzimodzi kangapo kamodzi pazaka 3-4 zilizonse. Kutenga kachilombo koyambitsa matenda a mbeu sikokwanira chifukwa cha kayendedwe kabwino ka mbeu.
Mwambiri, kuchuluka kwa inoculum mwa anthu aku Europe kumakhala kochepa, chifukwa chake kuwonjezeka kwa mliriwu kuli pang'onopang'ono ndipo kumatha kuyendetsedwa bwino pogwiritsa ntchito mankhwala a fungicidal kukonzekera. Ntchito yayikulu mdziko la Europe ndikulimbana ndi matenda mu gawo pomwe kufalikira kwa zoosporangia kuchokera kuzomera zomwe zakhudzidwa kumayamba.
Ku Russia, zinthu ndizosiyana kwambiri. Mbewu zambiri za mbatata ndi phwetekere zimalimidwa m'minda ing'onoing'ono; zodzitetezera mwina sizimachitidwa pa iwo konse, kapena mankhwala ophera fungic amachitika mosakwanira ndipo amayamba pambuyo poti chiwopsezo chachedwa pamwamba pake. Zotsatira zake, minda yamasamba yabwinobwino imakhala gwero lalikulu la matenda, komwe zoosporangia amanyamulidwa ndi mphepo kupita kuzomera zamalonda. Izi zikutsimikiziridwa ndi zomwe tawona ku Moscow, Bryansk, Kostroma, zigawo za Ryazan: kuwonongeka kwa mbewu m'minda yabwinobwino kumawonedwa ngakhale mankhwala a fungicide asanayambe. Pambuyo pake, mliri m'minda yayikulu umaletsedwa kugwiritsa ntchito fungicidal kukonzekera, pomwe m'minda yabwinobwino pali chitukuko chofulumira cha vuto lakumapeto.
Potengera njira zolakwika kapena "bajeti" yodzala m'minda yamalonda, chidwi cha zoipitsa mochedwa chimapezeka m'minda; Pambuyo pake akukula, akutenga madera akuluakulu (Elansky, 2015). Kutenga matenda m'minda yamwini kumakhudza kwambiri miliri m'misika yamalonda. M'madera onse olima mbatata ku Russia, dera lomwe mumakhala mbatata m'minda yabwinobwino limakulirapo kangapo kuposa malo onse olima. M'malo otere, minda yamasamba yabwinobwino imatha kuwonedwa ngati chida cha inoculum padziko lonse lapansi chamalonda. Tiyeni tiyesere kuzindikira zinthu zomwe zimafanana ndi mitundu ya zovuta m'minda yabwinobwino.
Kubzala mbewu zopanda mbewu ndi kuika kwaokha mbatata, mbewu za phwetekere zomwe zimapezeka kwaopanga akunja odabwitsa, kulima mbatata ndi tomato kwa nthawi yayitali m'malo omwewo, mankhwala osayenera a fungicide kapena kupezeka kwawo kumabweretsa ma epiphytoties ovuta m'magulu aboma, zomwe zotsatira zake ndi zaulere kuwoloka, kusakanizidwa ndikupanga ma oospores m'minda yabwinobwino. Zotsatira zake, mitundu yayikulu kwambiri ya tizilomboto timawonedwa, pomwe pafupifupi mtundu uliwonse wa matenda umakhala wapadera mu mtundu wake (Elansky et al., 2001, 2015). Kubzala mbatata za mbewu zosiyanasiyana zimapangitsa kuti kukhale kovuta kuti mizere yolimbana ndi mitundu ina ipangike. Mitundu yomwe yasankhidwa pankhaniyi imasiyanitsidwa ndi kusunthika kwawo poyerekeza ndi mitundu yomwe yakhudzidwa, ambiri aiwo amakhala ndi mitundu yambiri yamtundu wa virulence. Izi ndizosiyana kwambiri ndi machitidwe a "clonal lines" omwe amapezeka m'minda ikuluikulu yamabizinesi azolimo omwe ali ndi dongosolo loyenera lodzitchinjiriza ku ngozi yochedwa. "Clonal lines" (pomwe mitundu yonse yamatenda oyenda mochedwa m'munda ikuyimiridwa ndi mtundu umodzi kapena zingapo) imapezeka kulikonse m'maiko momwe kulima mbatata kumachitika kokha ndi minda yayikulu: USA, Netherlands, Denmark, ndi zina zambiri ku England, Ireland, Poland, komwe magawo amnyumba amakhalanso ofala Kukula kwa mbatata, palinso mitundu ina yayikulu kwambiri m'minda yabwinobwino. Kumapeto kwa zaka za zana la 20, "mizere ya clonal" inali ponseponse kumadera aku Asia ndi Far East ku Russia (Elansky et al., 2001), zomwe zikuwoneka chifukwa chogwiritsa ntchito mitundu yofananira ya mbatata pokhapokha pobzala. Posachedwa, zomwe zidachitika mzigawozi zidayambanso kusintha kuwonjezeka kwa mitundu yosiyanasiyana ya anthu.
Kuperewera kwa chithandizo champhamvu ndi fungicidal kukonzekera kuli ndi zotsatira zina, zachindunji - palibe kudzikundikira kwa zovuta zosagwira m'minda. Zowonadi, zotsatira zathu zikuwonetsa kuti mitundu yothana ndi metalaxyl imapezeka kawirikawiri m'minda yabwinoko kuposa m'minda yamalonda.
Kuyandikira kwambiri kwa malo obzala mbatata ndi phwetekere, omwe amapezeka m'minda yabwinobwino, kumathandizira kusunthika kwa mitundu pakati pa mbewuzo, zomwe chifukwa chake, mzaka khumi zapitazi, pakati pa mitundu yopanda mbatata, kuchuluka kwa mitundu yomwe imanyamula jini yokana mitundu yamatcheri yamatcheri (T1), omwe kale anali "" phwetekere "zovuta. Matenda omwe ali ndi jini la T1 nthawi zambiri amakhala ovuta kwambiri pa mbatata ndi tomato.
M'zaka zaposachedwa, vuto lochedwa pa phwetekere lidayamba kuonekera nthawi zambiri kuposa mbatata. Mbande za phwetekere zimatha kudzazidwa ndi ma oospores m'nthaka, kapena ma oospores omwe amapezeka mu nthangala za phwetekere kapena amamamatira (Rubin et al., 2001). M'zaka 15 zapitazi, mbewu zambiri zotsika mtengo, makamaka zotumizidwa kunja, zawonekera m'masitolo, ndipo ambiri opanga ang'onoang'ono asintha kuti azigwiritse ntchito. Mbeu zimatha kukhala ndimitundumitundu ndi ma genotypes ofanana ndi zigawo zomwe amalima. M'tsogolomu, ma genotypes awa amaphatikizidwa munjira zogonana m'minda yabwinobwino, zomwe zimapangitsa kuti pakhale mitundu yatsopano yatsopano.
Chifukwa chake, titha kunena kuti minda yabwinobwino ndi "poto wosungunuka" wapadziko lonse lapansi, momwe, chifukwa cha kusinthana kwa majini, mitundu yomwe ilipo imasinthidwa ndikuwonekeranso yatsopano. Kuphatikiza apo, kusankha kwawo kumachitika pansi pazikhalidwe zomwe ndizosiyana kwambiri ndi zomwe zidapangidwira mbatata m'minda yayikuru: kusapezeka kwa fungicidal press, mitundu yosiyanasiyana yazomera, kutsogola kwa mbewu zomwe zimakhudzidwa ndimatenda osiyanasiyana ndi mabakiteriya, pafupi ndi tomato ndi nightshades zakutchire, kuwoloka mwachangu ndi mapangidwe a oospore, kuthekera kuti oospores akhale ngati gwero la matenda kwa chaka chamawa.
Zonsezi zimabweretsa kusiyanasiyana kwakukulu kwa mabanja. Momwe zimakhalira ndi epiphytotic m'minda yamasamba, vuto lochedwa mochedwa limafalikira mwachangu kwambiri ndipo spores wambiri amatulutsidwa, akuwuluka kupita kumalo obzala pafupi. Komabe, atalowa m'minda yamalonda ndi njira yolondola yaukadaulo waulimi ndi chitetezo chamankhwala, ma spores omwe abwera alibe mwayi woyambitsa ma epiphytotic m'munda, chifukwa cha kusowa kwa mizere yolimbana ndi fungicides komanso yotchuka ndi mitundu yolimidwa.
Gwero lina la main inoculum lingakhale matenda a tubers omwe atsekeredwa mu mbande zamalonda. Izi tubers zidakula, monga lamulo, m'minda yokhala ndi ukadaulo wabwino waulimi komanso chitetezo chamankhwala. Ma genotypes amitundu yokhayokha omwe amakhudza ma tubers amasinthidwa kuti akwaniritse mitundu yawo. Mitundu iyi ndi yoopsa kwambiri pobzala malonda kuposa inoculum yochokera kuminda yabwinobwino. Zotsatira zamaphunziro athu zimathandiziranso izi. Anthu omwe amakhala kutali ndi minda yayikulu yokhala ndi chitetezo choyenera cha mankhwala komanso ukadaulo wabwino waulimi samasiyana pamitundu yosiyanasiyana. Nthawi zambiri iyi ndi mizere ingapo yama clonal yomwe imakhala yolusa kwambiri.
Zoyenga kuchokera kuzinthu zogulitsa zimatha kulowa anthu m'minda yamasamba ndikuchita nawo zomwe zikuchitika. Komabe, m'munda wamasamba, mpikisano wawo umakhala wotsika kwambiri poyerekeza ndi wamalonda, ndipo posachedwa adzaleka kukhalapo ngati clonal line, koma majini awo atha kugwiritsidwa ntchito mwa anthu "m'munda".
Matenda omwe amapezeka pazomera "zongodzipereka" komanso pamulu wa zimbudzi panthawi yokolola sizothandiza kwenikweni ku Russia, chifukwa M'madera omwe amalima mbatata kwambiri ku Russia, nthaka yozizira kwambiri yozizira imawonedwa, ndipo zomera kuchokera ku tubers zomwe zakhala m'nthaka nthawi zambiri sizimera. Kuphatikiza apo, monga momwe kuyesa kwathu kukuwonetsera, tizilombo toyambitsa matenda tomwe timachedwa ndi matendawa sizimakhalabe pamafunde otentha ngakhale pa ma tubers omwe apitilizabe kugwira ntchito. Kudera louma, komwe kulimidwa mbatata zoyambirira kumachitika, vuto lochedwa mochedwa ndilosowa chifukwa cha nyengo yowuma komanso yotentha.
Chifukwa chake, pano tikuwona kugawidwa kwa P. infestans anthu kukhala "m'munda" komanso "m'munda". Komabe, m'zaka zaposachedwa, njira zakhala zikuwonetsedwa zomwe zimapangitsa kuti pakhale kulumikizana ndi kulowererapo kwa majeremusi ochokera kwa anthuwa.
Mwa iwo, titha kuzindikira kuwonjezeka kwakukulu kwa kuwerenga kwa opanga ang'onoang'ono, kutuluka kwa mapaketi ang'onoang'ono a mbatata za mbewu, kufalikira kwa mapangidwe a fungicidal m'maphukusi ang'onoang'ono, komanso kutaya mantha kwa "chemistry" ndi anthu.
Zinthu zimachitika pomwe, chifukwa chantchito yogwira ntchito ya m'modzi wogulitsa, midzi yonse imabzalidwa ndi mbewu zambewu zamtundu womwewo ndikupatsidwa maphukusi ang'onoang'ono a mankhwala omwewo. Titha kuyerekezera kuti mbatata za mitundu yofananayo zitha kupezeka m'malo obzala malonda pafupi.
Kumbali inayi, makampani ena ogulitsa malonda a mankhwala akupititsa patsogolo njira za "bajeti" yothandizira mankhwala. Poterepa, kuchuluka kwa chithandizo chamankhwala kumayesedwa mopepuka ndipo fungicides yotsika mtengo imaperekedwa, ndipo sikuti kutsindika kuti kuthana ndi vuto lakumapeto kwa mdulidwe ndikutchetcha nsonga, koma pakuchedwa kwa epiphytoty pofuna kuwonjezera zokolola. Ndondomeko zoterezi ndizovomerezeka pachuma polima mbatata kuchokera kuzinthu zotsika, pomwe kulibe lingaliro lopeza zokolola zambiri. Komabe, pakadali pano, mosiyana ndi kuchuluka kwa anthu m'munda, mbadwa zomwe zatulutsidwa za mbatata zimathandizira pakusankha mitundu yakuthupi, yomwe ndi yoopsa pamitundu iyi.
Mwambiri, zizolowezi zakuphatikizana kwa "munda" ndi "munda" njira zopangira mbatata zimawoneka ngati zowopsa kwa ife. Pofuna kupewa zotsatirapo zake zoyipa, mnyumba komanso zamalonda, zikufunika kuwongolera mitundu yonse ya mbatata zambewu ndi mitundu yambiri ya fungicides yoperekedwa kwa eni ake m'maphukusi ang'onoang'ono, komanso kutsatira njira zotetezera mbatata komanso kugwiritsa ntchito fungicides mgulu lazamalonda.
M'madera azinsinsi, pali chitukuko chambiri chongokhala zoopsa osati mochedwa kokha, komanso Alternaria. Ambiri omwe ali ndi minda yabwinobwino satenga njira zapadera zodzitetezera ku Alternaria, kulakwitsa kukonza kwa Alternaria pakufota kwamasamba kapena kukulitsa koipitsa mochedwa. Chifukwa chake, ndikukula kwakukula kwa Alternaria pamitundu yomwe ingatengeke mosavuta, ziwembu zanyumba zitha kukhala gwero la inoculum pakudzala malonda.
Njira zosinthira
Njira yosinthira
Popeza kusintha kwa masinthidwe ndi njira yokhazikika yopitilira pafupipafupi, kupezeka kwa masinthidwe pamalo aliwonse kumatengera kuchuluka kwa kusintha kwa malowa komanso kuchuluka kwa anthu. Mukamaphunzira kuchuluka kwa masinthidwe amtundu wa P. infestans, kuchuluka kwa madera omwe amakula pazosankha zamagetsi atalandira chithandizo chamankhwala kapena mankhwala mutagens nthawi zambiri amatsimikiziridwa. Monga tingawonere kuchokera pazambiri zomwe zafotokozedwa mu Gome 8, kuchuluka kwakusintha kwamtundu womwewo m'malo osiyanasiyana kumatha kusiyanasiyana ndi maulamuliro angapo. Kusinthasintha kwakanthawi kosagwirizana ndi metalaxyl kungakhale chimodzi mwazifukwa zopezera mitundu yolimbana nayo m'chilengedwe.
Pafupipafupi zosintha mwadzidzidzi kapena zoyambitsa, zowerengedwa pamaziko a zoyeserera za labotale, sizimagwirizana nthawi zonse ndi zomwe zimachitika mwachilengedwe, pazifukwa izi:
1. Pokhala ndi ma fungo a nyukiliya osakanikirana, ndizosatheka kuyerekezera kuti zingasinthe pafupipafupi bwanji m'badwo umodzi wa zida za nyukiliya. Chifukwa chake, zoyeserera zambiri zimapereka chidziwitso chokhudzana ndi kusintha kwakanthawi, osasiyanitsa zochitika ziwiri zosintha ndi chochitika chimodzi chotsatira mitosis.
2. Kusintha kwa gawo limodzi nthawi zambiri kumachepetsa kuchepa kwa matupi athu, chifukwa chake, ndikupeza malo atsopano, kulimba kwa thupi kumachepa. Zambiri mwazomwe zasinthidwa poyeserera zimachepetsa kukwiya ndipo sizinalembedwe mwa anthu achilengedwe. Chifukwa chake, kulumikizana kokwanira pakati pamlingo wokana kwa P. infestans mutants to phenylamide fungicides and the grow rate on a artificial medium was on average (-0,62), and the resistance of fungicides and aggressionness on the masamba masamba (-0,65) (Derevyagina et al. , 1993), zomwe zikuwonetsa kuchepa kwa zosintha. Masinthidwe olimbana ndi dimethomorph adathandizidwanso ndi kuchepa kwakukulu kwa mphamvu (Bagirova et al., 2001).
3. Zosintha mwadzidzidzi komanso zosinthika ndizochulukirapo ndipo sizidziwonetsera zokha poyesa, koma zimapanga kusungidwa kosiyanasiyana kwa mitundu yachilengedwe. Mitundu yama Mutant yokhayokha poyesa labotale imakhala ndi kusintha kwakukulu (Kulish ndi Dyakov, 1979). Mwachiwonekere, diploidy ya zida za nyukiliya imalongosola zoyesayesa zosapambana zopezera zosintha motsogozedwa ndi kuyatsa kwa UV zomwe zimakhala zoyipa pamitundu yomwe kale inali yolimba (McKee, 1969). Malinga ndi kuwerengera kwa wolemba, zosinthazi zitha kuchitika pafupipafupi zosakwana 1: 500000. Kusintha kwamasinthidwe achilengedwe kukhala mkhalidwe wofanana, wowoneka bwino wa phenotypically kumatha kuchitika chifukwa chokhudzana ndi kugonana kapena kukonzanso zina (onani pansipa). Komabe, ngakhale pakadali pano, kusinthaku kumatha kuphimbidwa ndi maulamuliro akuluakulu amtundu wamtchire mu cenotic (multinucleated) mycelium ndi phenotypically okhazikika pokhapokha mononuclear zoospores ipangidwe.
Gulu 8. Pafupipafupi pa P. infestans amasintha pazinthu zoletsa zomwe zimachitika poyambitsa nitrosomethylurea (Dolgova, Dyakov, 1986; Bagirova et al., 2001)
Chophatikizira | Kusintha kwafupipafupi |
Mpweya | 6,9 x XUMUM-8 |
Pezani nkhaniyi pa intaneti Blasticidin S | 7,2 × 10-8 |
Streptomycin | 8,3 x10-8 |
Chithradze | 1,8 x XUMUM-8 |
Mpweya wambiri | 2,1 x XUMUM-8 |
Daaconil | <4 x 10-8 |
Zamatsenga | 6,3 x XUMUM-7 |
Metalaxil | 6,9 x XUMUM-6 |
Kukula kwa kuchuluka kwa anthu kumathandizanso kwambiri pakusintha kwadzidzidzi. M'magulu akulu kwambiri, momwe kuchuluka kwa maselo N> 1 / a, komwe kuli kuchuluka kwakusintha, kusintha kumatha kukhala chinthu chachilendo (Kvitko, 1974).
Kuwerengera kumawonetsa kuti pakakhala kufalikira kwa munda wa mbatata (mawanga 35 pachomera chilichonse), 8x1012 spores zimapangidwa tsiku lililonse pa hekitala imodzi (Dyakov ndi Suprun, 1984). Zikuwoneka kuti, anthu oterewa amakhala ndi kusintha konse kololedwa ndi mtundu wosinthana pamalo aliwonse. Ngakhale kusintha kosowa, komwe kumachitika pafupipafupi 10-9, kudzapezedwa ndi anthu chikwi mwa mamiliyoni omwe amakhala pa hekitala imodzi yamunda wa mbatata. Zosintha zomwe zimachitika pafupipafupi (mwachitsanzo, 10-6), mwa anthu otere, kusintha kosiyanasiyana kwamagulu kumatha kuchitika tsiku lililonse (nthawi imodzi ma loci awiri), i.e. njira zosinthira m'malo mwa zokonzanso.
Kusamuka
Kwa P. infestans, mitundu iwiri ikuluikulu yakusamukira imadziwika: kutseka mtunda (mkati mwa munda wa mbatata kapena madera oyandikana nawo) pofalitsa zoosporangia ndi mafunde am'mlengalenga kapena kutsitsi kwamvula, komanso mtunda wautali - ndikubzala ma tubers kapena kunyamula zipatso za phwetekere. Njira yoyamba imathandizira kukulitsa chidwi cha matendawa, chachiwiri - kukhazikitsidwa kwa malo atsopano m'malo akutali kwambiri.
Kufalikira kwa matenda opatsirana ndi tomato tubers ndi zipatso sizimangothandiza kuti matendawa ayambe kupezeka m'malo atsopano, komanso ndiye gwero lalikulu la kusiyanasiyana kwamitundu mwa anthu. M'dera la Moscow, mbatata zimabzalidwa, zochokera kumadera osiyanasiyana a Russia ndi Western Europe. Zipatso za phwetekere zimabwera kuchokera kumadera akumwera a Russia (dera la Astrakhan, dera la Krasnodar, North Caucasus). Mbeu za phwetekere, zomwe zitha kukhalanso ngati magwero a matenda (Rubin et al., 2001), amatumizidwanso kuchokera kumadera akumwera a Russia, China, mayiko aku Europe ndi mayiko ena.
Malinga ndi kuwerengera kwa E. Mayr (1974), kusintha kwa majeremusi m'deralo komwe kumachitika chifukwa cha kusintha kwa thupi sikunapitirire 10-5 pamalopo, pomwe kuli anthu otseguka, kusinthana chifukwa cha kutuluka kwa majini kuli osachepera 10-3 - 10-4.
Kusamuka kwa ma tubers omwe ali ndi kachilombo koyambitsa matendawa ndi komwe kumayambitsa kulowa kwa P. infestans ku Europe, kufalikira kumadera onse padziko lapansi komwe mbatata zimabzalidwa; adayambitsa kusintha kwakukulu kwa anthu. Chochedwa chakumapeto kwa mbatata chinawonekera m'dera la Ufumu wa Russia pafupifupi nthawi yomweyo ndikuwonekera ku Western Europe.
Popeza matendawa adadziwika koyamba mu 1846-1847 ku Baltic States ndipo kokha m'zaka zotsatira adafalikira ku Belarus ndi zigawo zakumpoto chakumadzulo kwa Russia, chiyambi chake chakumadzulo kwa Europe ndichodziwikiratu. Gwero loyamba la zoyipitsa mochedwa mu Dziko Lakale silodziwika kwenikweni. Lingaliro lopangidwa ndi Fry et al. (Fry et al., 1992; Fry, Goodwin, 1995, Goodwin et al., 1994) akuwonetsa kuti tizilomboti tidachokera koyamba ku Mexico kupita ku North America, komwe kudafalikira kudzera mu mbewu, kenako nkupititsidwa ku Western Europe. (mkuyu 7).
Chifukwa cha kubowoleza mobwerezabwereza (zotsatira ziwiri za "botolo"), matanthwe amodzi adafika ku Europe, ana omwe adayambitsa mliri kudera lonse la Old World, komwe mbatata zimabzalidwa. Monga umboni wa malingaliro awa, olembawo amatchula, choyamba, kupezeka kwa mtundu umodzi wokha wa mating (A1) ndipo, chachiwiri, kusakanikirana kwa mitundu ya mitundu yophunziridwa yochokera kumadera osiyanasiyana (yonseyi idakhazikitsidwa ndi ma molekyulu, kuphatikiza 2 isozyme loci, zolemba zala za DNA, ndi Kapangidwe ka DNA ya mitochondrial ndi ofanana, ndipo amafanana ndi choyerekeza cha US-1 chofotokozedwa ku USA). Komabe, deta zina zimabweretsa kukayikira pazinthu zina zomwe zanenedwa. Kufufuza kwa P. infestans DNA ya mitochondrial yotalikirana ndi mitundu ya mbatata ya herbarium yomwe idatengera matendawa nthawi yoyamba ya epiphytotic m'ma 40 idawonetsa kuti amasiyana pamapangidwe a DNA ya mitochondrial kuchokera ku choyerekeza cha US-1, chomwe chinali osati gwero lokhalo lotenga matenda ku Europe (Ristaino et al, 2001).
Mavuto akuchedwa adayambiranso kukulira m'zaka za m'ma 80s. Zosintha izi zachitika:
1) Chiwawa cha anthu chikuwonjezeka, zomwe zadzetsa, makamaka, kufalikira kwa mitundu yoyipa kwambiri yamatsenga - kuwonongeka kwa petioles ndi zimayambira.
2) Panali kusintha munthawi yowawa kwa mbatata - kuyambira kumapeto kwa Julayi mpaka koyambirira kwa Julayi mpaka kumapeto kwa Juni.
3) Mtundu wokhathamira wa A2, womwe kale kudalibe ku Old World, wafika ponseponse.
Zosinthazi zidachitika ndi zochitika ziwiri: kugwiritsa ntchito kwakukulu kwa fungala metalaxyl (Schwinn ndi Staub, 1980) komanso kutuluka kwa Mexico ngati wogulitsa kunja mbatata (Niederhauser, 1993). Malinga ndi izi, zifukwa ziwiri zakusintha kwa kuchuluka kwa anthu zidaperekedwa: kusintha kwa mtundu wa mating motsogozedwa ndi metalaxyl (Ko, 1994) ndikukhazikitsa mitundu yatsopano yamatenda omwe ali ndi kachilombo kochokera ku Mexico (Fry ndi Goodwin, 1995). Ngakhale kutembenuka kwamitundu yosakanikirana motengera mphamvu ya metalaxyl sikunapezeke kokha ndi Ko, komanso ntchito zomwe zidachitika mu labotale ya Moscow State University (Savenkova, Chherepennicova-Anikina, 2002), lingaliro lachiwiri ndilabwino. Pamodzi ndikuwoneka kwamtundu wachiwiri wa mating, kusintha kwakukulu kunachitika mu genotypes ya Russian P. infestans strains, kuphatikiza majini osalowerera ndale (isozyme ndi RFLP loci), komanso kapangidwe ka DNA ya mitochondrial. Kusintha kwakusintha kumeneku sikungathe kufotokozedwa ndimachitidwe a metalaxyl; M'malo mwake, panali tanthauzo lalikulu la mitundu yatsopano yochokera ku Mexico, yomwe, pokhala yankhanza kwambiri (Kato et al., 1997), idasamutsa mavuto akale (US-1), kukhala olamulira mwa anthu. Kusintha kwa kuchuluka kwa anthu aku Europe kunachitika munthawi yochepa kwambiri - kuchokera 1980 mpaka 1985 (Fry et al., 1992). M'madera omwe kale anali USSR, "mitundu yatsopano" idapezeka m'magulu ochokera ku Estonia mu 1985, ndiye kuti, koyambirira kuposa ku Poland ndi Germany (Goodwin et al., 1994). Nthawi yotsiriza "mavuto akale US-1" ku Russia adasiyanitsidwa ndi phwetekere wokhala ndi kachilomboka m'chigawo cha Moscow mu 1993 (Dolgova et al., 1997). Komanso ku France, mitundu "yakale" idapezeka m'mabzala a phwetekere mpaka koyambirira kwama 90, ndiye kuti, atatha kale mbatata (Leberton ndi Andrivon, 1998). Kusintha kwa zovuta za P. infestans kunakhudza mikhalidwe yambiri, kuphatikiza yofunikira kwambiri, ndikuwonjezera kuwonongeka kwa vuto lakumapeto.
Kubwezeretsanso kugonana
Pofuna kuti kukumbukiraninso pakuthandizira kusiyanasiyana, ndikofunikira, choyamba, kupezeka kwa mitundu iwiri ya mating m'chiwerengero cha 1: 1, ndipo kachiwiri, kupezeka kwa kuchuluka kwa anthu.
Chiwerengero cha mitundu yokwatirana chimasiyanasiyana kwambiri m'magulu osiyanasiyana ngakhale zaka zosiyana pagulu limodzi (Gulu 9,10, 90). Zifukwa zosinthira motere pakuchulukitsa mitundu yamatenda mwa anthu (monga, mwachitsanzo ku Russia kapena ku Israeli koyambirira kwa zaka za m'ma 2002 za mzaka zapitazi) sizikudziwika, koma akukhulupirira kuti izi ndichifukwa chokhazikitsa mitundu yolimbirana (Cohen, XNUMX).
Zina zosadziwika zikuwonetsa kachitidwe kogonana mzaka zina komanso zigawo zina:
1) Kafukufuku wa anthu ochokera mdera la Moscow adawonetsa kuti mwa anthu 13 momwe kuchuluka kwa mtundu wa mating A2 kunali kochepera 10%, kusiyanasiyana konse komwe kumawerengedwa atatu isozyme loci anali 0,08, ndipo mwa anthu 14 momwe gawo la A2 lidapitilira 30%, kusiyanasiyana kwa majini kunali kokwera kawiri (0,15) (Elansky et al., 1999). Chifukwa chake, kuthekera kwakutali kwakugonana, kumachulukitsa mitundu ya anthu.
2) Chiyanjano pakati pa chiŵerengero cha mitundu yokwatirana mwa anthu komanso kuchuluka kwa mapangidwe a oospore chidawonedwa ku Israeli (Cohen et al., 1997) ndi ku Holland
(Flier et al., 2004). Kafukufuku wathu adawonetsa kuti m'mitundu yomwe imadzipatula ndi mtundu wa matupi a A2 imakhala 62, 17, 9, ndi 6%, ma oospores amapezeka m'masamba a mbatata a 78, 50, 30, ndi 15% (okhala ndi mawanga awiri kapena kupitilira apo), motsatana.
Zitsanzo zokhala ndi mawanga awiri kapena kupitilira apo nthawi zambiri zimakhala ndi oospores kuposa zitsanzo zokhala ndi malo amodzi (2 ndi 1% yazitsanzo, motsatana) (Apryshko et al., 32).
Oospores anali ofala kwambiri m'masamba apakati komanso otsika a mbewu ya mbatata (Mytsa et al., 2015; Elansky et al., 2016).
3) M'madera ena, mitundu yodziwika bwino yodziwika bwino yapezeka, zomwe zimachitika chifukwa chokhudzana ndi kugonana. Chifukwa chake, ku Poland mu 1989 komanso ku France mu 1990, mavuto a homozygous a glucose-6-
mankwala isomerase (GPI 90/90). Popeza kale ma heterozygotes 10/90 okha ndi omwe adakumana nawo kwa zaka 100, homozygosity akuti imayambitsanso kugonana (Sujkowski et al., 1994). Ku Colombia (USA), kupatula kuphatikiza A2 ndi GPI 100/110 ndi A1 ndi GPI 100/100 ndizofala, komabe, kumapeto kwa nyengo ya 1994 (Ogasiti 16 ndi Seputembara 9), mavuto okhala ndi zophatikizanso zophatikizika (A1 GPI 100/110 ndi A2 GPI 100/100) (Miller et al., 1997).
4) Mwa anthu ena ochokera ku Poland (Sujkowski et al., 1994) ndi North Caucasus (Amatkhanova et al., 2004), kugawa kwa zala za DNA loci ndi allozyme protein loci kukufanana ndi kugawa kwa Hardy-Weinberg, komwe kukuwonetsa
za gawo lalikulu lazopereka zofananira pakugonana pakusiyanasiyana kwa anthu. M'madera ena a Russia, palibe makalata omwe amafotokozedwa ndi Hardy-Weinberg mwa anthu omwe adapezeka, koma kupezeka kwa mgwirizano wolumikizana kunawonetsedwa, kuwonetsa kuchuluka kwa kubereka kwa clonal (Elansky et al., 1999).
5) Kusiyanasiyana kwa majini (GST) pakati pamavuto amitundu yosiyanasiyana (A1 ndi A2) inali yotsika poyerekeza pakati pa anthu osiyanasiyana (Sujkowski et al., 1994), yomwe imawonetsa misewu yogonana.
Nthawi yomweyo, zopereka zakukumbutsa zakugonana pakusiyanasiyana kwa anthu sizingakhale zazikulu kwambiri. Choperekachi chinawerengedwa kwa anthu okhala m'chigawo cha Moscow (Elansky et al., 1999). Malinga ndi kuwerengera kwa Lewontin (1979), "kukonzanso, komwe kumatha kupanga mitundu yatsopano kuchokera ku ma loci awiri pafupipafupi osapitilira zomwe zimachokera ku heterozygosities awo, kumakhala kothandiza pokhapokha ngati kuchuluka kwa heterozygosity kwa ma alleles onse kwakhala kale kale."
Ndi kuchuluka kwa mitundu iwiri ya mapangidwe, omwe ali m'chigawo cha Moscow, wofanana ndi 4: 1, mafupipafupi okhala 0,25. Kuthekera koti mitanda yodutsa idzakhala heterozygous kwa awiri mwa atatu mwa omwe adaphunzira isozyme loci mwa anthu omwe anaphunzira anali 0,01 (2 mitundu kuchokera ku 177). Chifukwa chake, kuthekera kwakupezeka kwa ma heterozygotes ophatikizika chifukwa chobwezeretsanso sikuyenera kupitilira zomwe zidachulukitsidwa ndikuwoloka (0,25x0,02x0,02) = 10-4, i.e. zophatikizanso zogonana nthawi zambiri sizigwera muzitsanzo zophunziridwa za mitundu. Kuwerengera uku kunapangidwira anthu ochokera kudera la Moscow omwe amadziwika ndi kusiyanasiyana kwakukulu. M'madera a monomorphic monga a ku Siberia, njira zogonana, ngakhale zitachitika mwa anthu amodzi, sizingayambitse mitundu yawo.
Kuphatikiza apo, P. infestans amadziwika ndi kusokonekera kwa chromosome pafupipafupi mu meiosis, komwe kumabweretsa aneuploidy (Carter et al., 1999). Zophwanya izi zimachepetsa kubereka kwa ma hybrids.
Kuphatikizanso kwa parasexual, kusintha kwa majini a mitotic
Poyesera kusakanikirana kwa P. infestans mitundu ndi kusintha kwa kukana kwa zoletsa zosiyanasiyana zakukula, kutuluka kwa misolates yolimbana ndi zoletsa zonse ziwiri kunapezeka (Shattock ndi Shaw, 1975; Dyakov, Kuzovnikova, 1974; Kulish, Dyakov,
1979). Zovuta zomwe sizimagwirizana ndi zoletsa kukula ziwiri zidayamba chifukwa cha heterokaryotization ya mycelium, ndipo potero adalumikizana panthawi yobereka ndi mononuclear zoospores (Judelson, Ge Yang, 1998), kapena sanadziphatikize mu ana a monozoosporous, chifukwa anali ndi tetraploid (popeza opatula koyamba ndi diploid) nuclei (K (1979). Ma Heterozygous diploids omwe amapatulidwa pafupipafupi chifukwa cha haploidization, chromosome nondisjunction, komanso mitotic yodutsa (Poedinok et al., 1982). Pafupipafupi pa njirazi zitha kuwonjezeka mothandizidwa ndi zovuta zina pa heterozygous diploids (mwachitsanzo, UV walitsa wa zikumera).
Ngakhale mapangidwe azomera zam'magulu osagwiritsidwa ntchito kawiri samapezeka mu vitro kokha, komanso muzitsamba za mbatata zomwe zimasakanikirana ndi zosintha (Kulish et al., 1978), ndizovuta kudziwa momwe parasexual recombination ingapangire mibadwo yatsopano ya genotypes mwa anthu. Kuchuluka kwa magawo opangidwa chifukwa cha haploidization, kusadukiza ma chromosomes ndi mitotic kuwoloka popanda zovuta zina ndizochepa (zosakwana 10-3).
Kutuluka kwa magawo amadzimadzi amtundu wa heterozygous kumatha kutengera kutembenuka kwa mitotic kudutsa ndikusintha kwa majini a mitotic, komwe ku P. sojae kumachitika pafupipafupi 3 x 10-2 mpaka 5 x 10-5 pamalopo, kutengera kupsinjika (Chamnanpunt et al. , 2001).
Ngakhale pafupipafupi ma heterokaryons ndi heterozygous diploids amakhala osayembekezereka okwera (kufika makumi khumi peresenti), njirayi imachitika kokha ngati zikhalidwe zosintha zomwe zimapezeka munthawi yomweyo zimadulidwa. Mukamagwiritsa ntchito mitundu yosiyanasiyana yolekanitsidwa ndi chilengedwe, heterokaryotization sichimachitika (kapena kumachitika pafupipafupi) chifukwa chakupezeka kosagwirizana kwa masamba (Poedinok ndi Dyakov, 1981; Anikina et al., 1997b; Cherepennikova-Anikina et al., 2002). Zotsatira zake, gawo lakukonzanso parasexual kumatha kuchepetsedwa kungokhala kukonzanso kophatikizana kwamkati mwa heterozygous nuclei ndikusintha kwa majini amtundu wina kukhala mkhalidwe wosagonana. Izi zitha kukhala zofunikira kutengera matenda am'magazi omwe ali ndi kusintha kosintha kwa fungicide. Kusintha kwake kukhala mkhalidwe wosagwirizana chifukwa cha zochitika zapadera kumakulitsa kukana kwa omwe asintha kusintha (Dolgova, Dyakov, 1986).
Kufotokozera kwa majini
Mitundu ya Heterothallic Phytophthora imatha kuswana ndikupanga ma haibridi osakanizidwa (onani Vorob'eva ndi Gridnev, 1983; Sansome et al., 1991; Veld et al., 1998). Mtundu wosakanizidwa wachilengedwe wa mitundu iwiri ya Phytophthora udali wankhanza kwambiri mwakuti udapha anthu masauzande ambiri ku UK (Brasier et al., 1999). P. infestans imatha kuchitika ndi mitundu ina yamtunduwu (P. erythroseptica, P. nicotianae, P. Cactorum, etc.) pazomera wamba komanso m'nthaka, koma palibe zambiri m'mabukuwa za kuthekera kwa mitundu ingapo ya hybridi. Pansi pama labotale, hybrids adapezeka pakati pa P. infestans ndi P. Mirabilis (Goodwin and Fry, 1994).
Tebulo 9. Gawo la P. infestans limakumana ndi mtundu wa mating a A2 m'maiko osiyanasiyana padziko lapansi kuyambira 1990 mpaka 2000 (malinga ndi chidziwitso chopezeka m'mabuku ndi masamba a www.euroblight.net, www.eucablight.org)
dziko | 1990 | 1991 | 1992 | 1993 | 1994 | 1995 | 1996 | 1997 | 1998 | 1999 | 2000 |
---|---|---|---|---|---|---|---|---|---|---|---|
Belarus | 33 (12) | 34 (29) | |||||||||
Belgium | 15 (49 *) | 6 (66) | 20 (86) | ||||||||
Ecuador | 0 (13) | 0 (12) | 0 (19) | 0 (21) | 12 (41) | 25 (39) | 15 (75) | 22 (73) | 25 (68) | 0 (35) | |
Estonia | 8 (12) | ||||||||||
England | 4 (26) | 3 (630) | 9 (336) | ||||||||
Finland | 0 (15) | 19 (117) | 12 (16) | 21 (447) | 6 (509) | 9 (432) | 43 (550) | ||||
France | 0 (35) | 0 (56) | 0 (83) | 0 (67) | 0 (86) | 2 (135) | 7 (156) | 6 (123) | 0 (73) | 0 (285) | 0 (135) |
Hungary | 72 (32) | ||||||||||
Ireland | 4 (145) | ||||||||||
Kumpoto. 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) | ||||||||
Amuna | 0 (16) | 7 (97) | 0 (48) | 0 (25) | |||||||
Korea | 36 (42) | 10 (130) | 15 (98) | ||||||||
China | 20 (142, 1995-98) | 0 (6) | 0 (8) | 0 (35) | |||||||
Colombia | 0 (40, 1994-2000) | ||||||||||
Uruguay | 100 (25, 1998-99) | ||||||||||
Morocco | 60 (108, 1997-2000) | 52 (25) | 42 (40) | ||||||||
Serbia | 76 (37) | ||||||||||
Mexico (Toluca) | 28 (292, 1988-89) | 50 (389, 1997-98) |
Tebulo 10. Gawo la P. infestans limakanika ndi mtundu wa mating A2 m'maiko osiyanasiyana padziko lapansi kuyambira 2000 mpaka 2011.
dziko | 2001 | 2002 | 2003 | 2004 | 2005 | 2006 | 2007 | 2008 | 2009 | 2010 | 2011 |
---|---|---|---|---|---|---|---|---|---|---|---|
Austria | 65 (83) | ||||||||||
Belarus | 42 (78) | ||||||||||
Belgium | 20 (102 *) | 4 (32) | 50 (14) | 25 (16) | 62 (13) | 54 (26) | 70 (54) | 30 (23) | 29 (35) | 62 (71) | 45 (49) |
Switzerland | 89 (19) | ||||||||||
Czech Republic | 35 (31) | 54 (64) | 38 (174) | 12 (80) | |||||||
Germany | 95 (53) | ||||||||||
Denmark | 48 (52) | ||||||||||
Ecuador | 5 (178) | 6 (108) | 9 (121) | 18 (94) | 2 (44) | 0 (66) | 5 (47) | ||||
Estonia | 54 (25) | 0 (24) | 33 (62) | 45 (140) | 25 (100) | 12 (103) | |||||
England | 4 (47) | 10 (96) | 31 (55) | 55 (790) | 68 (862) | 70 (552) | 68 (299) | ||||
Finland | 47 (162) | 12 (218) | 42 | ||||||||
France | 0 (186) | 4 (108) | 8 (61) | 22 (103) | 33 (303) | 65 (378) | 74 (331) | 75 (125) | 75 (12) | ||
Hungary | 48 (27) | 48 (90) | 9 | 7 | |||||||
Kumpoto. 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) | |||||||
Amuna | 25 (12) | 68 (106) | 80 (88) | 92 (143) | 75 (45) | ||||||
Korea | 46 (26) | ||||||||||
Brazil | 0 (49) | 0 (30) | |||||||||
China | 10 (30) | 0 (6) | 0 (6) | ||||||||
Вьетнам | 0 (294, 2003-04) | ||||||||||
Uganda | 0 (8) |
Mphamvu zamtundu wa genotypic wa anthu
Kusintha kwa mtundu wa genotypic wa P. infestans kuchuluka kumatha kuchitika chifukwa cha kusamuka kwa miyala yatsopano kuchokera kumadera ena, machitidwe azaulimi (kusintha mitundu, kugwiritsa ntchito mafangasi), komanso nyengo. Zokopa zakunja zimakhudza mawonekedwe am'magulu osiyanasiyana munthawi zosiyanasiyana za moyo; Chifukwa chake, chaka chilichonse anthu amakumana ndi kusintha kwamitundumitundu ya majini omwe amasankhidwa, chifukwa chosintha gawo lalikulu la kusunthika kwa majini ndikusankhidwa.
Mphamvu zakusiyanasiyana
Mitundu yatsopano yomwe ili ndi majini othandiza kutsutsana molunjika (R-genes) ndi chinthu champhamvu chosankha, kusankha ma clones okhala ndi majeremusi owonjezera a P. infestans. Pakakhala kusakanikirana kwamitundu yosiyanasiyana ya mbatata, komwe kumalepheretsa kukula kwa tizilomboti, njira yosinthira mitundu yayikulu kwambiri ya anthu imachitika mwachangu kwambiri. Chifukwa chake, atafalikira kudera la Moscow la Domodedovsky zosiyanasiyana, lomwe lili ndi R3 jini yolimbana, mafupipafupi amtundu woyipa wazosiyanazi adakula kuchoka pa 0,2 mpaka 0,82 mchaka chimodzi (Dyakov ndi Derevjagina, 2000).
Komabe, kusintha kwa kuchuluka kwa majeremusi a virulence (pathotypes) mwa anthu kumachitika osati kokha chifukwa cha mitundu yolimidwa ya mbatata. Mwachitsanzo, ku Belarus mpaka 1977, ma clones omwe ali ndi majeremusi oyambitsa 1 ndi 4 adalamulira, zomwe zidachitika chifukwa cha kulima mitundu ya mbatata yolimbana ndi majini a R1 ndi R4 (Dorozhkin, Belskaya, 1979). Komabe, kumapeto kwa ma 70s a 2002th century, ma clones adawoneka ndi mitundu yosiyanasiyana ya ma virulence ndi kuphatikiza kwawo, ndipo mitundu yothandizirana yolimbana nayo sinagwiritsidwepo ntchito pobzala mbatata (majeremusi owonjezera) (Ivanyuk et al., XNUMX). Chifukwa cha mawonekedwe amenewa, mwachiwonekere, ndi chifukwa cha kusamukira ku Europe kwa zinthu zopatsirana zochokera ku Mexico ndi tubers za mbatata. Kunyumba, matanthwewa amayamba osati pa mbatata zokha, komanso mitundu yamtchire yomwe imanyamula mitundu yambiri yamavuto; Chifukwa chake, kuphatikiza mitundu yambiri yamavuto mu genome kunali kofunikira kuti mukhale ndi moyo munthawiyo.
Ponena za mitundu yosagwirizana kwenikweni, iwo, pochepetsa kuchepa kwa tizilombo toyambitsa matenda, amachepetsa kusinthika kwa anthu ake, omwe, monga tanenera kale, ndi ntchito yochulukirapo. Popeza kupsa mtima kumakhala kosiyanasiyana, matanthwe okhala ndi chibadwa chochuluka cha "kupsa mtima" amadziunjikira mwachangu kukula kwa anthu. Chifukwa chake, mafuko aukali kwambiri samapangidwa chifukwa cha kusintha kwa mitundu yolimidwa mosagwirizana kwenikweni, koma, m'malo mwake, imapezeka kwambiri mukamabzala mitundu yovuta kwambiri yomwe imapezekanso ndi tiziromboti.
Chifukwa chake, ku Russia, anthu ankhanza kwambiri a P. Infestans amapezeka kumadera a epiphytoties apachaka (anthu ochokera ku Sakhalin, Leningrad, ndi madera a Bryansk). Kukwiya kwa anthuwa kunakhala kwakukulu kuposa aku Mexico (Filippov et al., 2004).
Kuphatikiza apo, ma oospores ochepa amapangidwa m'masamba amitundu yosagonjetsedwa kuposa omwe ali pachiwopsezo (Hanson ndi Shattock, 1998), ndiye kuti, kulimbikira kopanda tanthauzo kwa mitunduyi kumathandizanso kuti tizilomboto tisathenso kukumbatirana komanso kuthekera kwa njira zina zachisanu.
Mphamvu ya fungicides
Mafungicides samachepetsa kuchuluka kwa bowa wa phytopathogenic, i.e. zimakhudza mawonekedwe owerengeka a anthu awo, koma amathanso kusintha mafupipafupi amitundu yamtundu uliwonse, i.e. zimakhudza kapangidwe kabwino ka anthu. Zina mwazizindikiro zofunika kwambiri za kuchuluka kwa anthu omwe akusintha chifukwa cha fungicides ndi izi: kusintha kwa kulimbana ndi fungicides, kusintha kwaukali ndi nkhanza, komanso kusintha kwa njira zoswana.
Mphamvu ya fungicides pakukaniza komanso kupsa mtima kwa anthu
Kukula kwa chikoka kumatsimikizika, choyambirira, ndi mtundu wa fungicide yomwe imagwiritsidwa ntchito, yomwe imatha kugawidwa polysite, oligosite komanso monosite.
Zakale zimaphatikizapo fungicides yolumikizana kwambiri. Kukaniza kwa iwo (ngati kuli kotheka konse) kumayendetsedwa ndi majini ambiri ofooka kwambiri. Katunduyu amadziwika kuti kulibe kusintha kosawoneka bwino kwa anthu atalandira chithandizo ndi fungicides (ngakhale pakuyesa kwina, kuwonjezeka kwina kwa kukana kunapezeka). Chiwerengero cha mafangasi omwe amasungidwa atapopera mankhwala ophera fungicides omwe ali ndi magulu awiri amitundu:
1) Zovuta zomwe zimasungidwa m'malo omwe simunalandire mankhwalawa. Popeza kulibe kulumikizana ndi fungicide, kukwiya komanso kukana kwa mitundu iyi sikusintha.
2) Mitsinje yolumikizana ndi fungicide, yomwe ndende zake zinali zochepa poyerekeza ndi zowopsa. Monga tafotokozera pamwambapa, kulimbikira kwa gawo ili la anthu sikusintha, komabe, chifukwa cha kuwonongeka pang'ono kwa fungicide ngakhale mu mitsempha yochepetsetsa ya kagayidwe kake ka fungal cell, kulimbitsa thupi komanso gawo lake lazinyama, kukwiya, kuchepa (Derevyagina ndi Dyakov, 1990).
Chifukwa chake, ngakhale gawo la anthu omwe sanamwalire, atakumana ndi fungicide, ali ndi nkhanza zochepa ndipo sangakhale gwero la ma epiphytotic. Chifukwa chake, chithandizo chamankhwala mosamala chomwe chimachepetsa kuchuluka kwa kuchuluka kwa anthu osakhudzana ndi fungicide ndichikhalidwe chothandizira kuteteza. Kukaniza ma fungicides oligosite kumayang'aniridwa ndi majini angapo owonjezera.
Kusintha kwa jini lirilonse kumabweretsa kuwonjezeka kwa kukana, ndipo kuchuluka kwake kwakukaniza kumachitika chifukwa cha kusintha kwa kusintha koteroko. Chifukwa chake, kuwonjezeka kwa kukana kumachitika pang'onopang'ono. Chitsanzo cha kuwonjezeka kwapang'onopang'ono kwa kukana ndi kusintha kwa mphamvu polimbana ndi fungicide dimethomorph, yomwe imagwiritsidwa ntchito kwambiri kuteteza mbatata ku vuto lakumapeto. Kukaniza kwa Dimethomorph ndi polygenic komanso zowonjezera. Kusintha kamodzi kumawonjezera kukana.
Kusintha kulikonse komwe kumatsatira kumachepetsa kukula kwake, motero, kuchuluka kwa masinthidwe otsatira (Bagirova et al., 2001). Kuchuluka kwa kukana kwapakati pa anthu atalandira chithandizo mobwerezabwereza ndi oligosite fungicide kumachitika pang'onopang'ono komanso pang'onopang'ono. Kufulumira kwa njirayi kumatsimikiziridwa ndi zinthu zitatu: kuchuluka kwa kusintha kwa majini otsutsana, koyefishienti yotsutsana (chiŵerengero cha mlingo woopsa wa mankhwala osagonjetseka poyerekeza ndi yovuta) komanso momwe masinthidwe amtundu wa majini olimbikira amakhala olimba.
Zomwe zimachitika pakasinthidwe kalikonse ndizotsika poyerekeza ndi zam'mbuyomu, chifukwa chake njirayi ili ndi vuto lochepa (Bagirova et al., 2001). Komabe, ngati njira zopangidwanso (zogonana kapena zogonana) zimachitika mwa anthu, ndiye kuti ndizotheka kuphatikiza kusintha kosiyanasiyana kwa makolo mumtundu wosakanizidwa ndikufulumizitsa ntchitoyi. Chifukwa chake, anthu a panmix amayamba kukana mwachangu kuposa agamic, ndipo kumapeto kwake, anthu omwe alibe zovuta zosagwirizana pazomera mwachangu kuposa anthu omwe amalekanitsidwa ndi zotchinga zotere. Pankhaniyi, kupezeka kwa mitundu ya anthu yomwe imasiyana pamitundu ikukhathamiritsa kumathandizira njira yolimbana ndi fungicides ya oligosite.
Zinthu zachiwiri ndi zachitatu sizimathandizira kukulira kwachangu kwa mitundu yolimbana ndi dimethomorph mwa anthu. Kusintha kulikonse komwe kumatsatira kumachulukitsa kukana, komwe kulibe tanthauzo, komanso nthawi yomweyo kumachepetsa kukula komwe kumapangidwe komanso kukwiya (Bagirova et al., 2001; Tsinde, Kirk, 2004). Ichi ndi chifukwa chake mulibe mitundu yolimbana pakati pa masoka achilengedwe a P. infestans, ngakhale omwe amatengedwa kuchokera kubzala kwa mbatata yothandizidwa ndi dimethomorph.
Chiwerengero cha anthu omwe amachiritsidwa ndi fungicide ya oligosite adzaphatikizanso magulu awiri amtundu: omwe sanalumikizane ndi fungicide, chifukwa chake sanasinthe mawonekedwe oyambilira (ngati mitundu yolimbana ikupezeka mgululi, sichidzadzikundikira chifukwa chaukali komanso mpikisano wamagulu azovuta), ndipo imakumana ndi zovuta zowopsa za fungicide. Ndipakati pazomwezi kuti kupezeka kwa mitundu yolimbana ndi zotheka ndi kotheka, chifukwa apa ali ndi zabwino kuposa zovuta.
Chifukwa chake, mukamagwiritsa ntchito fungicides ya oligosite, sichithandizo chokwanira kwambiri chomwe chimafunikira ngati kuchuluka kwa mankhwala, kangapo kuposa kuchuluka kwakupha, chifukwa ndi stepwise mutagenesis, kukana koyamba kwa mitundu yosinthidwa ndikotsika.
Pomaliza, kusintha kosagwirizana ndi fungicides yayikulu ndikofotokozera, ndiye kuti, kusintha kamodzi kumatha kunena zakukaniza kwakukulu mpaka kumaliza kutaya mtima. Chifukwa chake, kuchuluka kwa kukana kwa anthu kumachitika mwachangu kwambiri.
Chitsanzo cha fungicides zotere ndi phenylamides, kuphatikiza mankhwala a fungala metalaxyl. Kusintha kwa kukana izi kumachitika pafupipafupi, ndipo kuchuluka kwakusintha kwa zosintha kumakhala kwakukulu kwambiri - kumadutsa kupsinjika kovuta ndi chinthu chikwi kapena kupitilira apo (Derevyagina et al., 1993). Ngakhale kukula ndi kukwiya kwa zosintha zosagonjetsedwa kumachepa poyambira chifukwa chakufa kwa tizilombo toyambitsa matenda kuchokera ku fungicic systemic, kuchuluka kwa anthu osagwirizana kukukula mwachangu ndipo kukwiya kwawo kukukulira mofananamo. Chifukwa chake, patadutsa zaka zingapo ndikugwiritsa ntchito fungicide, kulimbana kwa mitundu yosagonjetsedwa sikungangofanana ndi nkhanza za omwe ali ndi vuto, komanso kulilambalala (Derevyagina, Dyakov, 1992).
Zovuta zakumbukirananso zogonana
Popeza kupezeka kwapafupipafupi kwa mtundu wa A2 wokwatirana mu P. infestans kuchuluka kudagwirizana ndikugwiritsa ntchito kwambiri metalaxyl motsutsana ndi vuto lakumapeto, akuti amati metalaxyl imathandizira kutembenuka kwamtundu wamating. Mu P. parasitica, kutembenuka koteroko mothandizidwa ndi chloroneb ndi metalaxyl kunatsimikiziridwa poyesa (Ko, 1994). Vuto limodzi lokhala ndi sing'anga lokhala ndi metalaxyl locheperako lidapangitsa kuti patuluke mitundu yodzipatula kuchokera ku P. infestans yomwe imazindikira metalaxyl yokhala ndi mtundu wa A1 (Savenkova ndi Cherepnikova-Anikina, 2002). M'magawo otsatila atolankhani okhala ndi metalaxyl yochulukirapo, palibe mtundu umodzi wokha wa mtundu wa A2 womwe unapezeka, komabe, ambiri amadzipatula, akawoloka ndi A2, m'malo mwa oospores, adapanga zovuta za mycelium ndipo anali osabala. Magawo amtundu wina wosakanikirana wokhala ndi mtundu wa mating A2 pa media wokhala ndi metalaxyl yochuluka amatilola kuti tipeze mitundu itatu yamitundu yosinthira: 1) kusabereka kwathunthu tikadutsa ndi kudzipatula kwa A1 ndi A2; 2) homotallism (mapangidwe a oospores mu monoculture); 3) kutembenuka kwa mtundu wa mating A2 kukhala A1. Chifukwa chake, metalaxyl imatha kuyambitsa kusintha kwamitundu yamatenda mwa anthu a P. infestans ndipo, chifukwa chake, kupezeka kwamalingaliro okhudzana ndi kugonana mwa iwo.
Zotsatira zakukumbukiranso kwamasamba
Mitundu ina ya maantibayotiki yolimbana ndi maantibayotiki idachulukitsa kuchuluka kwa hephokaryotization ya hyphal ndi diploidization ya nyukiliya (Poedinok ndi Dyakov, 1981). Monga tanena kale, heterokaryotization ya hyphae panthawi yosakanikirana kwamatenda osiyanasiyana a P. infestans imachitika kawirikawiri chifukwa chazovuta zakusagwirizana kwamankhwala mu bowa. Komabe, majini olimbana ndi maantibayotiki ena amatha kukhala ndi zotsatirapo, zimawonetsedwa pothana ndi kusagwirizana pazomera. Katunduyu anali ndi 1S-1 mutant streptomycin resistance gene. Kukhalapo kwa zosintha zotere m'minda ya phytophthora kumatha kukulitsa kuchuluka kwa majini pakati pamavuto ndikuthandizira kusintha kwa anthu onse kukhala mitundu yatsopano kapena fungicides.
Ma fungicides ndi maantibayotiki amatha kusokoneza kukonzanso kwa mitotic, komwe kumasinthanso kuchuluka kwa mitundu ya anthu. Fungicide benomyl yomwe imagwiritsidwa ntchito kwambiri imagwirizana ndi beta-tubulin, puloteni yomwe ma microtubules a cytoskeleton amamangidwa, motero imasokoneza njira zopatulira kwa chromosome mu anaphase ya mitosis, ndikuwonjezera kuchuluka kwa mitotic recombination (Hastie, 1970).
Fungicide para-fluorophenylalanine, yomwe imagwiritsidwa ntchito pochizira matenda achi Dutch mu elms, ili ndi malo omwewo. Para-fluorophenylalanine idakulitsa kuchuluka kwakubwezeretsanso kwa heterozygous diploids P. infestans (Poedinok et al., 1982).
Kusintha kwamitundu yambiri pakupanga kwamankhwala m'thupi la P. infestans
Makulidwe akale a P. infestans mdera lotentha amakhala ndi magawo anayi.
1) Gawo lokulirapo la kuchuluka kwa anthu (polycyclic phase) ndimibadwo yayifupi. Gawoli nthawi zambiri limayamba mu Julayi ndipo limatha miyezi 1,5-2.
2) Gawo lolepheretsa kuchuluka kwa anthu chifukwa chakuchepa kwakanthawi kwamitundu yosakhudzidwa kapena kuyambika kwanyengo. Gawo ili m'minda yomwe imachita masamba asanakolole masamba atachotsedwa kumapeto kwa chaka.
3) Gawo la nyengo yozizira mu ma tubers, limodzi ndi kuchepa kwakukulu kwa kuchuluka kwa anthu chifukwa cha matenda mwangozi a tubers, kukula pang'ono kwa matenda mwa iwo, kusapezeka kwa matenda a tubers, kuwola ndikuchotsa ma tubers okhudzidwa munthawi yosungira.
4) Gawo lakukula pang'onopang'ono m'nthaka ndi mmera (monocyclic phase), momwe nthawi yoberekera imatha kufikira mwezi umodzi kapena kupitilira apo (kumapeto kwa Meyi - koyambirira kwa Julayi). Kawirikawiri panthawiyi, masamba odwala amakhala ovuta kuwazindikira, ngakhale atawona mwapadera.
Gawo lakukula kwa chiwonetsero cha anthu (gawo la polycyclic)
Zowonera zambiri (Pshedetskaya, Kozubova, 1969; Borisenok, 1969; Osh, 1969; Dyakov, Suprun, 1984; Rybakova, Dyakov, 1990) adawonetsa kuti koyambirira kwa epiphytoty, miyala yamphamvu kwambiri komanso yolusa pang'ono imakhalapo, yomwe imasinthidwa ndikuwopsa. Kukula kwa kuchuluka kwa anthu kumakhala kwakukulu, zocheperako zotsalira zimatsika.
Pamene chiwerengero cha anthu chikukula, kuchuluka kwa majini onse osankhidwa omwe amayambitsidwa mumitundu yamalonda (R1-R4) ndikusankhapo mbali (R5-R11) kumawonjezeka. Chifukwa chake, mwa anthu omwe ali pafupi ndi Moscow mu 1993, kuchuluka kwa nkhanza kuyambira kumapeto kwa Julayi mpaka pakati pa Ogasiti kudakwera kuchoka pa 8,2 mpaka 9,4, ndipo chiwonjezeko chachikulu chidawonedwa chifukwa cha jini ya virulence R5 (kuyambira 31 mpaka 86% yamiyala yoyipa) (Smirnov, 1996 ).
Kuchepa kwa kuchuluka kwa kuchuluka kwa anthu kumayendera limodzi ndi kuchepa kwa ziwalo za anthu. Chifukwa chake, mzaka zokhumudwitsa, mitundu yonse yamitundu ndi kuchuluka kwa mitundu yamphamvu kwambiri ndiyotsika poyerekeza ndi ya epiphytotic (Borisenok, 1969). Ngati nyengo ya epiphytotic ikasinthira kukhala yovuta chifukwa chamatenda akuchedwa ndipo kufalikira kwa mbatata kumachepa, kuchuluka kwa miyala yoopsa komanso yamwano kumachepa (Rybakova et al., 1987).
Kuwonjezeka kwa mafupipafupi a majini omwe amakhudza kusokonekera komanso kukwiya kwa anthu atha kukhala chifukwa chosankha mitundu yowopsa kwambiri komanso yosokoneza mwa anthu osakanikirana. Kuwonetsa kusankha, njira yowunikira kusintha kosalowerera ndale idapangidwa, yomwe idagwiritsidwa bwino ntchito popanga yisiti (Adams et al., 1985) ndi Fusarium graminearum (Wiebe et al., 1995).
Kuchuluka kwa zosintha zosagwirizana ndi blasticidin S m'magawo a P. infestans kunachepa chimodzimodzi ndikukula kwankhanza kwa anthu, zomwe zikuwonetsa kusintha kwamitundu yayikulu pakukula kwa anthu (Rybakova et al., 1987).
Gawo lozizira mu tubers
Pakati pa nyengo yozizira mu tubers ya mbatata, chiwopsezo ndi nkhanza za P. infestans zimachepa, ndipo kuchepa kwa virulence kumachitika pang'onopang'ono kuposa kukwiya (Rybakova ndi Dyakov, 1990). Mwachiwonekere, pansi pazikhalidwe zomwe zimapangitsa kukula kwakukula kwa kuchuluka kwa anthu (r-kusankha), majini "owonjezera" a virulence ndiukali kwambiri ndi othandiza, chifukwa chake kukula kwa ma epiphytotic kumatsagana ndi kusankha kwamiyala yoyipa kwambiri komanso yamakani. M'mikhalidwe yodzaza chilengedwe, pomwe sikuchulukana, koma kulimbikira kukhalapo m'malo osavomerezeka (K-kusankha) kumachita gawo lofunikira, majini "owonjezera" aukazitape ndiukali amachepetsa kulimba, ndipo ma clones okhala ndi majini awa ndi omwe amayamba kufa, kotero kuti kukwiya kwapakati komanso nkhanza za anthu zikuchepa.
Gawo lazomera m'nthaka
Gawo ili ndichinsinsi kwambiri m'moyo (Andrivon, 1995). Kukhalapo kwake kudatumizidwa mwakachetechete - chifukwa chakusowa chidziwitso chazomwe zimachitika ndi tizilombo toyambitsa matenda kwanthawi yayitali (nthawi zina kuposa mwezi) - kuchokera pomwe mbande za mbatata zidayamba kuwonekera pomwe matendawa adayamba. Pogwiritsa ntchito zowunika ndi zoyeserera, machitidwe a bowa munthawi yamoyo adamangidwanso (Hirst ndi Stedman, 1960; Boguslavskaya, Filippov, 1976).
Kutsekemera kwa bowa kumatha kupangidwa ndi ma tubers omwe ali ndi kachilombo m'nthaka. Zomwe zimatuluka zimamera ndi ma hyphae, omwe amatha kubzala nthawi yayitali m'nthaka. Zoyambira (zopangidwa ndi ma tubers) ndi zachiwiri (pa mycelium m'nthaka) zimamera m'nthaka ndi mafunde a capillary, koma zimatha kupatsira mbatata masamba ake otsika atatsika ndikumana ndi nthaka. Masamba otere (omwe ndi mawanga oyamba a matendawa amapezeka pa iwo) samapanga nthawi yomweyo, koma atakula kwakanthawi ndikukula kwa nsonga za mbatata.
Chifukwa chake, gawo lazomera za saprotrophic amathanso kukhalanso m'moyo wa P. infestans. Ngati gawo la parasitic la moyo waukali ndi gawo lofunikira kwambiri pakulimbitsa thupi, ndiye kuti pakusankha gawo la saprotrophic cholinga chake ndikuchepetsa zida zamatenda, monga zikuwonetsedwa poyesa bowa wina wa phytopathogenic (onani Carson, 1993). Chifukwa chake, mgawo lino lazungulira, katundu waukali ayenera kutayika kwambiri. Koma pakadali pano palibe zoyeserera zachindunji zomwe zachitika kuti zitsimikizire malingaliro omwe ali pamwambapa.
Kusintha kwa nyengo kumakhudza osati tizilombo toyambitsa matenda a P. infestans, komanso kukana kwa fungicides, komwe kumakula mu gawo la polycyclic (nthawi ya epiphytoties), ndipo kumachepa nthawi yosungira nthawi yachisanu (Derevyagina et al., 1991; Kadish ndi Cohen, 1992). Kutsika kwakukulu kwa kukana kwa metalaxyl kunawonedwa pakati pa kubzala kwa ma tubers okhudzidwa ndikuwoneka kwa mawanga oyamba a matenda m'munda.
Intraspecific specialization ndi kusintha kwake
P. infestans akuyambitsa miliri mu mbewu ziwiri zofunika kuchita, mbatata ndi phwetekere. Epiphytoties pa mbatata idayamba bowa atangolowa m'malo atsopano. Kugonjetsedwa kwa phwetekere kunadziwikanso patangopita nthawi yochepa kuchokera ku matenda a mbatata, koma ma epiphytoties a phwetekere adadziwika patatha zaka zana limodzi - pakati pa zaka za XNUMXth. Izi ndi zomwe Hallegli ndi Niederhauser alemba zakugonjetsedwa kwa tomato ku USA
(1962): "Pafupifupi zaka 100 pambuyo pa epiphytoty yoopsa ya 1845, zochepa kapena zoyesayesa zoyesayesa zomwe zidapangidwa kuti zitha kupeza mitundu yolimbana ndi phwetekere. Ngakhale kuwawalitsa mochedwa koyamba kulembedwa pa tomato koyambirira kwa 1848, sikunakhale chinthu chofunikira kwambiri kwa obzala pa chomeracho mpaka kubuka kwamatenda mu 1946. M'dera la Russia mochedwa choipitsa cha phwetekere adalembetsedwa m'zaka za zana la 60. “Kwa nthawi yayitali, ofufuza sanasamale za matendawa, chifukwa sanayambitse mavuto azachuma. Koma m'ma 70s ndi 1979s. Epiphytoties zakumapeto kwa zaka za m'ma XNUMX zakumapeto kwa vuto la tomato zimapezekanso ku Soviet Union, makamaka mdera la Lower Volga, ku Ukraine, North Caucasus, ku Moldova ... ”(Balashova, XNUMX).
Kuchokera nthawi imeneyo, kuwonongeka kwa tomato nthawi yayitali kumakhala chaka chilichonse, kufalikira kudera lonse lakulima kwamakampani ndi nyumba ndikuwononga chuma chambiri. Chinachitika ndi chiyani? Kodi nchifukwa ninji kuwonekera koyamba kwa tiziromboti pa mbatata ndi chotupa cha epiphytotic cha mbewuyi zidachitika pafupifupi nthawi imodzi, ndipo bwanji zidatenga zaka zana kuti epiphytotic iwoneke pa phwetekere? Kusiyanaku kumathandizira waku Mexico osati waku South America wopatsirana. Ngati mitundu ya Phytophthora infestans idapangidwa ngati tizirombo tomwe timakhala ndi tizilombo tating'onoting'ono tomwe timapezeka ku Mexico Solanum, ndizomveka chifukwa chake mbatata zolimidwa zomwe zili mgawo lomwelo monga mitundu ya Mexico zidakhudzidwa kwambiri, koma chifukwa chosowa kophatikizana ndi tiziromboti, zomwe sizinapangitse njira zosagwirizana ndi ena.
Phwetekere ndi gawo lina la mtunduwo, mtundu wake wosinthana umasiyana kwambiri ndi mitundu ya tuberous, chifukwa chake, ngakhale kuti phwetekere siliri kunja kwa chakudya cha P. infestans, kuchuluka kwa kuwonongeka kwake sikunali kokwanira kutayika kwakukulu kwachuma.
Kutuluka kwa epiphytoties pa phwetekere kumachitika chifukwa cha majeremusi omwe amasintha, omwe adakulitsa kusintha kwake (pathogenicity) atasakazidwa. Tikukhulupirira kuti mtundu watsopanowu womwe udzawononga phwetekere ndi mtundu wa T1 wofotokozedwa ndi M. Gallegly, womwe umakhudza mitundu ya phwetekere (Red Cherry, Ottawa), yolimbana ndi mpikisano wa T0 wofalikira pa mbatata (Gallegly, 1952). Mwachiwonekere, kusintha (kapena kusintha kosiyanasiyana) komwe kunasanduliza mpikisano wa T0 kukhala mpikisano wa T1 ndikuwatsogolera kuwoneka kwama clones omwe adasinthidwa kuti agonjetse phwetekere. Monga zimachitika kawirikawiri, kuwonjezeka kwa tizilombo toyambitsa matenda kwa munthu m'modzi kumayendera limodzi ndi kuchepa kwa wina, ndiye kuti, woyamba, wosakwanira ukadaulo wa intraspecific udayamba - mbatata (mtundu T0) ndi phwetekere (mtundu T1).
Kodi umboni wa lingaliro lotere ndi uti?
- Zomwe zimachitika mbatata ndi tomato. Pamasamba a phwetekere, mpikisano wa T1 umakhalapo, pomwe pamasamba a mbatata ndizochepa. Malinga ndi S.F.Bagirova ndi T.A. Oreshonkova (osasindikizidwa) m'chigawo cha Moscow mu 1991-1992, kupezeka kwa mpikisano wa T1 m'mabzala a mbatata kunali 0%, ndipo m'minda yobzala tomato - 100%; mu 1993-1995 - 33% ndi 90%, motsatana; mu 2001 - 0% ndi 67%. Zambiri zofananira zidapezeka ku Israel (Cohen, 2002). Kuyesedwa kwa matenda a tubers a mbatata omwe amakhala ndi mtundu wa T1 osakanikirana komanso osakanikirana ndi T0 ndi T1 adawonetsa kuti olekanitsidwa ndi mpikisano wa T1 samasungidwa bwino mumachubu ndipo amalowetsedwa m'malo ndi othamanga a T0 (Dyakov et al., 1975; Rybakova, 1988).
2) Mphamvu za mpikisano wa T1 m'mabzala a phwetekere. Matenda oyambilira a masamba a phwetekere amachitika ndi gulu lokhalokha la T0, lomwe limayang'anira kusanthula kwa matenda m'madontho oyamba opangidwa pamasamba. Izi zikutsimikizira chiwembu chovomerezeka cha kusamuka kwa tiziromboti: Matenda akulu a mbatata amapangidwa ndi mpikisano wa T0, komabe, miyala yochepa ya T1 yosungidwa mu mbatata, kamodzi pa phwetekere, imachotsa mpikisano wa T0 ndikudziunjikira kumapeto kwa nyengo ya epiphytotic. Ndikothekanso kuti pali njira ina yomwe ingayambitsire tsamba la phwetekere ndi mpikisano wa T1, womwe siwamphamvu ngati mizu ya mbatata ndi masamba, koma mosalekeza. Chifukwa chake, gwero ili limafooketsa mawonekedwe amtundu wa anthu omwe akupha phwetekere, koma pambuyo pake limatsimikizira kuchuluka kwa mpikisano wa T1 (Rybakova, 1988; Dyakov et al., 1994).
3) Kulusa mbatata ndi tomato. Matenda opangidwa ndi masamba a phwetekere ndi mbatata omwe amakhala ndi mitundu yokhayokha T0 ndi T1 adawonetsa kuti zoyambazo ndizovuta kwambiri ku mbatata kuposa phwetekere, ndipo zomalizazi ndizoyipa kwambiri kwa phwetekere kuposa mbatata. Kusiyanaku kumawonetsedwa pakusamuka kwa anthu osadzipangira okha omwe si "awo" ochokera pakati pa anthu osakanikirana panthawi yama masamba mumtengowo (D'yakov et al., 1975) komanso m'minda yolima (Leberton et al., 1999); kusiyanasiyana kwakuchepa kwa kachilombo koyambitsa matenda, kutalika kwa nthawi, kukula kwa mawanga opatsirana ndi kupanga ma spore (Rybakova, 1988; Dyakov et al., 1994; Legard et al., 1995; Forbes et al., 1997; Oyarzun et al., 1998; Leberton et al. al., 1999; Vega-Sanchez et al., 2000; Knapova, Gisi, 2002; Sussuna et al., 2004).
Kukhwima kwa omwe amadzipatula pamtundu wa T1 kupita ku mitundu ya phwetekere yomwe ilibe majini otsutsana ndiwokwera kwambiri kwakuti izi zimasiyanitsa spore pamasamba ngati cholumikizira michere popanda kupukusa minofu yomwe ili ndi kachilomboka (Dyakov et al., 1975; Vega-Sanchez et al., 2000).
4) Kusefukira kwa mbatata ndi tomato. Mpikisano wa T1 umakhudza mitundu ya phwetekere ndi mtundu wa Ph1, pomwe mpikisano wa T0 sungathe kupatsira mitundu iyi, i.e. ali ndi vutoli locheperako. Pokhudzana ndi kusiyanitsa
Mitundu ya mbatata imakhala yolumikizana, i.e. Mitundu yokhayokha yamasamba a phwetekere ndiyowopsa kwambiri kuposa mitundu ya "mbatata" (Gulu 11)
5) Zolemba zandale. Kufufuza kwa zolembera zosalowerera ndale mwa anthu a P. infestans omwe amadya mbatata ndi tomato kumatsimikiziranso kusankha kosiyanasiyana kwamitundu yambiri. Mwa anthu aku Brazil a P. infestans, masamba obalidwa ndi phwetekere anali amtundu wa US-1, ndipo ochokera masamba a mbatata anali a BR-1 mzere (Suassuna et al., 2004). Ku Florida (USA), kuyambira 1994, mtundu wa US-90 udayamba kulamulira mbatata (zomwe zimachitika kupitilira 8%), ndikuwombera US-11 ndi US-17 pa phwetekere, ndipo omwe amadzipatulawa amakhala akuvuta kwambiri phwetekere kuposa mbatata (Weingartner , Tombolato, 2004). Kusiyanitsa kwakukulu pamitundu yama genotype (zolemba zala za DNA) m'matumba ndi mbatata zidakhazikitsidwa pazaka 1200 P. infestans tizilombo ta ku United States kuyambira 1989 mpaka 1995 (Deahl et al., 1995).
Kugwiritsa ntchito njira ya AFLP kunapangitsa kuti kuthe kusiyanitsa mitundu 74 yomwe imasonkhanitsidwa kuchokera masamba a mbatata ndi tomato mu 1996-1997. ku France ndi Switzerland, m'magulu 7. Mitundu ya mbatata ndi phwetekere sinasiyane bwino, koma mitundu ya "mbatata" inali chibadwa chosiyanasiyana kuposa "phwetekere". Zoyambazo zidapezeka m'magulu asanu ndi awiriwo, ndipo omalizirayo, anayi okha, zomwe zikuwonetsa genome yapaderadera kwambiri (Knapova ndi Gisi, 2002).
6) Njira zodzipatula. Ngati anthu omwe ali ndi tiziromboti pamitundu iwiri yazomera asintha kuti akhale ochepa okha, ndiye kuti pali njira zingapo zomwe zimalepheretsa kusinthana kwamitundu (Dyakov ndi Lekomtseva, 1984).
Kafukufuku angapo adasanthula momwe gwero la zovuta za makolo limathandizira pakusakanikirana. Powoloka mitundu yotalikirana ndi mitundu yosiyanasiyana ya mtundu wa Solanum ku Ecuador (Oliva et al., 2002), zidapezeka kuti mitundu yokhudzana ndi mtundu wa A2 kuchokera kutchire Solanaceae (clonal line EC-2) ndi yomwe imadutsa kwambiri kuchokera ku phwetekere (mzere EC -3), ndikuwoloka bwino kwambiri ndi mavuto a mbatata (EC-1).
Zosakanizidwa zonse zimapezeka kuti sizomwe zimayambitsa matenda. Olembawo amakhulupirira kuti kuchuluka kwa kuperewera ndi kuchepa kwa tizilombo tating'onoting'ono chifukwa cha njira zoperekera zakubadwa za anthu.
Poyesa kwa Bagirova et al. (1998), mitundu yambiri ya mbatata ndi phwetekere idawoloka ndi mitundu ya T0 ndi T1. Mitengo yachonde kwambiri inali mitanda ya T1xT1 yokhayokha kuchokera ku phwetekere (ma oospores 36 m'mayikulosikopu momwe amawonera, 44% ya kumera kwa oospore), osagwira ntchito kwambiri inali mitanda yamipikisano ya T0xT1 yotalikirana ndi magulu osiyanasiyana (kuchuluka kocheperako kwa oospores omwe amatukuka komanso ophulika) ... Kuchita bwino kwa mitanda pakati pa olekanitsidwa ndi mpikisano wa T0 wopatula mbatata kunali kwapakatikati. Popeza kuti mitundu yayikulu yamtundu wa T0 imakhudza mbatata, ili ndi gwero lodalirika la nyengo yozizira - tubers ya mbatata, chifukwa chake kufunika kwa ma oospores monga magawo ozizira opatsirana a anthu ochokera ku mbatata ndikotsika. "Maonekedwe a phwetekere" amatha nyengo yachisanu pa phwetekere ngati oospores (onani m'munsimu) motero amakhala ndi zokolola zambiri pazakugonana. Chifukwa cha kubereka kwake kwakukulu, T1 imapeza mwayi wodziyimira payokha wodwala matendawa mu phwetekere. Zotsatira zomwe Knapova et al. (Knapova et al., 2002) atha kutanthauziridwa chimodzimodzi. Mitanda yamitundumitundu yotalikirana ndi mbatata yokhala ndi tizilombo ta phwetekere idapereka ma oospores ochuluka kwambiri - 13,8 pa sq. sing'anga (ndikufalikira kwa 5-19) ndi kuchuluka kwapakati pakumera kwa oospores (6,3 ndikufalikira kwa 0-24). Kuwoloka kwa mitundu yotalikirana ndi phwetekere kunapereka magawo otsika kwambiri a oospores (7,6 ndikufalikira kwa 4-12) ndi kuchuluka kwambiri kwakumera kwawo (10,8). Mitanda yapakati pa zovuta zomwe zimasiyanitsidwa ndi mbatata idapereka ma oospores angapo (8,6 okhala ndi kufalikira kwakukulu kwa data - 0-30) komanso gawo lotsikitsitsa la kumera kwa oospores (2,7). Chifukwa chake, zovuta za mbatata ndizocheperako poyerekeza ndi za phwetekere, koma mitanda yolowererapo sikunabweretse zotsatira zoyipa kuposa kuchuluka kwa anthu. Ndizotheka kuti kusiyana ndi zomwe zili pamwambazi ndi Bagirova et al. amafotokozedwa ndikuti ofufuza aku Russia adagwira ntchito ndi mitundu yokhayokha koyambirira kwa zaka za m'ma 90 za zana la makumi awiri, ndipo ofufuza aku Switzerland - omwe ali ndi mitundu yokhayokha kumapeto kwa ma 90s.
Maziko oberekera ochepa atha kukhala heteroploidy yamavuto. Ngati ndi anthu aku Mexico, komwe njira zogonana komanso matenda oyamba ndi ana oospore amakhala wamba, mitundu yambiri ya P. Infestans ndi diploid, ndiye kuti m'maiko a Old World kuchuluka kwa ma polymorphism of ploidy kumawonedwa (di-, tri- and tetraploid strains, komanso heterokaryotic strains with heteroploid nuclei) , ndi mitundu yokhala ndi mitundu yosiyanasiyana yokwatirana, i.e. Chomera chokhwima, chosiyana ndi zida za nyukiliya (Therrien et al., 1989, 1990; Whittaker et al., 1992; Ritch, Daggett, 1995). Kusiyanasiyana kwa ma nuclei mu antheridia ndi oogonia kumatha kukhala chifukwa chakubala pang'ono.
Ponena za kusinthana kwa zida za nyukiliya pakati pa hyphae panthawi ya anastomoses, izi zimapewedwa chifukwa chosagwirizana kwamasamba, komwe kumagawana asexual m'miyala yambiri yobadwa nayo (Poedinok ndi Dyakov, 1987; Gorbunova et al., 1989; Anikina et al., 1997b).
7) Kukhazikika kwa anthu. Zomwe zatchulidwazi zikuwonetsa kuti kuphatikiza kwa "mbatata" ndi "phwetekere" P. infestans strains ndikotheka. Kubwezeretsanso kachilombo koyambitsa magulu osiyanasiyana ndikotheka, ngakhale ndikuchepetsa kukwiya.
Kafukufuku wowerengera anthu omwe amakhala m'madera oyandikana ndi mbatata ndi phwetekere ku 1993 adawonetsa kuti pafupifupi kotala la magawo osiyanitsidwa ndi masamba a phwetekere adasamutsidwa kuchokera kumunda woyandikira wa mbatata (Dolgova et al., 1997). Mwachidziwitso, titha kuganiza kuti kusiyana kwa anthu m'magulu awiriwo kungakule ndikubweretsa kutuluka kwa mitundu yapadera ya intraspecific (f.sp. mbatata ndi f.sp. Phwetekere), makamaka popeza ma oospores amatha kupitilirabe pazinyalala za mbewu (Drenth et al., 1995 ; Bagirova, Dyakov, 1998) ndi nthanga za phwetekere (Rubin et al., 2001). Chifukwa chake, tomato pakadali pano ali ndi gwero la kusinthika kwamasiku osadalira zipatso za mbatata.
Komabe, zonse zinachitika mosiyana. Kudzadzidwa ndi ma oospores kudalola kuti tizilomboto tipewe gawo lochepetsetsa m'moyo wake - gawo lokhalokha lazomera m'nthaka, pomwe zimatha kuchepa, zomwe zimabwezeretsedwanso gawo la polycyclic chilimwe.
Tebulo 11. Mafupipafupi amtundu wa virulence kumitundu yosiyanitsa mbatata m'matenda a P. infestans
dziko | Год | Avereji ya majeremusi achiwawa m'matenda | wolemba | |
kuchokera ku mbatata | kuchokera phwetekere | |||
France | 1995 | 4.4 | 3.3 | Leberton et al., 1999 |
1996 | 4.8 | 3.6 | Leberton, Andrivon, 1998 | |
France, Switzerland | 1996-97 | 6.8 | 2.9 | Knapova, Gisi, 2002 |
United States | 1989-94 | 5 | 4.8 | Goodwin et al., 1995 |
USA, Zap. Washington | 1996 | 4.6 | 5 | Dorrance et al., 1999 |
1997 | 6.3 | 3.5 | " | |
Ecuador | 1993-95 | 7.1 | 1.3 | Oyarzun et al., 1998 |
Israel | 1998 | 7 | 4.8 | Cohen, 2002 |
1999 | 6 | 5.7 | " | |
2000 | 6.7 | 6.1 | " | |
Russia, Mosk. dera | 1993 | 8.9 | 6.7 | Smirnov, 1996 |
Russia, madera osiyanasiyana | 1995 | 9.4 | 8 | Kozlovskaya ndi ena. |
1997 | 9.2 | 9.2 | " | |
2000 | 8.7 | 4.8 | " |
Zoyambira zoosporangia ndi zoospores, zomwe zimamera ma oospores, zimakhala ndi zochitika zambiri zaparasiti, makamaka ngati ma oospores adapangidwa parthenogenetically mchikakamizo cha ma pheromones amtundu wina wotsutsana. Chifukwa chake, zomwe zimafalikira pa mbande za phwetekere zomwe zimakula kuchokera ku mbewu zomwe zili ndi ma oospores ndizoyambitsa matenda a phwetekere ndi mbatata.
Kusintha kumeneku kunadzetsa kukonzanso kwa anthu ena, kotchulidwa posintha kumeneku kuchokera pakuwunika kwa matenda:
- Matenda a phwetekere omwe ali ndi kachilombo akhala chinthu chofunikira kwambiri pakupatsira mbatata (Filippov, Ivanyuk, mauthenga ake).
- Epiphytoties pa mbatata idayamba kuwonetsedwa koyambirira kwa Juni, pafupifupi mwezi umodzi kale kuposa masiku onse.
- M'mabzala mbatata, kuchuluka kwa mpikisano wa T1 kudakulirakulira, komwe kumapezeka kale mumtengo wopanda pake (Ulanova et al., 2003).
- Zovuta zomwe zimasiyanitsidwa ndi masamba a phwetekere sizinasiyanenso ndimitundumitundu ya mbatata yolimbana ndi mbatata yosiyanitsa mitundu yabwinobwino ndipo idayamba kupitirira zovuta za "mbatata" mwamphamvu osati pa phwetekere kokha, komanso pa mbatata (Lavrova et al., 2003; Ulanova et al. , 2003).
Chifukwa chake, m'malo mosiyana, panali kuphatikiza kwa anthu, kutuluka kwa mtundu umodzi pazomera ziwiri zomwe zimakhala ndiukali komanso nkhanza kuzinthu zonse ziwiri.
Pomaliza
Chifukwa chake, ngakhale zaka zopitilira 150 zophunzira mozama za P. infestans, mu biology, kuphatikiza biology ya anthu ya nthumwi yofunikira kwambiri yamatenda ofunikira a mbewu zolimidwa ndi solanaceous, zambiri sizikudziwika. Sizikudziwika bwino momwe gawo la magawo azomwe zimakhudzira kuchuluka kwa kuchuluka kwa anthu, ndi njira ziti zamatenda zosiyanasiyanirana zankhanza ndi nkhanza, chiŵerengero cha njira zoberekera ndi kubereka mwa anthu achilengedwe, momwe kusagwirizana kwamasamba kumatengera cholowa, udindo wa mbatata ndi tomato pakuyambitsa matendawa ndi chiyani chimakhudza bwanji kuchuluka kwa tiziromboti. Mpaka pano, zinthu zofunikira monga majini osinthira kukhathamira kwa tiziromboti kapena kukokoloka kwa kukana kwa mbatata kopanda tanthauzo sikunathetsedwe. Ndikukula ndi kufalikira kwa kafukufuku wokhudzana ndi mbatata mochedwa, kachilomboka kamabweretsa mavuto atsopano kwa ofufuza. Komabe, kupititsa patsogolo kuyeserera koyeserera, kutuluka kwa njira zatsopano zogwiritsira ntchito majini ndi mapuloteni kumatilola kuyembekezera yankho labwino pamafunso omwe afunsidwa.
Nkhaniyi idasindikizidwa munyuzipepala yoteteza "mbatata" (No. 3, 2017)