ʻO S.N. Elansky, L.Yu. Kokaeva, N.V. Statsyuk, Yu.T. ʻO Dyakov
Hōʻike
ʻO Oomycete Phytophthora infestans (Mont.) ʻO De Bary, ka mea hana o ka hopena o ke ahi, ka maʻi ʻoi loa o ka uala a me nā tōmato, ua kau nui ka manaʻo o nā kānaka noiʻi mai nā ʻāina like ʻole no hoʻokahi kenekulia a me ka hapa. ʻIke koke ʻia i ʻEulopa i waenakonu o ke kenekulia XNUMX, ua kumu ia i kahi maʻi potato i hoʻomau ʻia i ka hoʻomanaʻo ʻana o nā hanauna he nui.
A hiki i kēia manawa, kapa pinepine ʻia ia "mushroom of the Irish pōloli". Ma kahi o hoʻokahi haneli mau makahiki ma hope o nā maʻi ʻōpala mua, ua ʻike ʻia nā ʻano ʻuala ʻĀina hihiu kūʻē i ka hopena o ke ahi, ua hoʻomohala ʻia nā ʻano hana o ke keʻa iā lākou me nā ʻuala i mahi ʻia (Muller, 1935), a ua loaʻa nā ʻano kūʻē mua kūpale mua (Pushkarev, 1937). Eia nō naʻe, ma hope koke iho o ka hoʻomaka ʻana o kā lākou mahiʻai kalepa ʻana, nā heihei o ka lohi o nā maʻi pathogen i virulent i nā ʻano kūʻē i hōʻiliʻili ʻia. a me ka hoʻokomo ʻana i nā genes kūʻē hou mai nā ʻuala ʻāhiu o Mekiko i nā ʻano i hoʻomaka i ka lilo wikiwiki ʻana i ka pono.
ʻO nā kūleʻa ʻole me ka hoʻohana ʻana i ke kūʻē ʻana o monotaga (pākū) i nā mea hoʻohuli i hoʻohuli ʻia e huli no nā ala ʻoi aku ka paʻakikī o ka hoʻohana ʻana i ke kūʻē polygenic (papamoe) kū ʻole. I nā makahiki i hala iho nei, ua hoʻomaka nā ʻāʻī huhū nui e hōʻiliʻili i kēlā me kēia heluna kanaka o ka parasite, e hoʻoneoneo ana i ke kūʻē ʻole nonspecific. ʻO ka hōʻea ʻana mai o nā ʻano kūpilikiʻi kūpale i mea hoʻopilikia i ka hoʻohana ʻana i nā kemika pale ʻuala.
Ma muli o nā ʻokoʻa nui ma waena o nā oomycetes a me nā fungi i ka hoʻohui kemika, ultrastructure, a me ka metabolism, fungicides, ʻo ia hoʻi nā ʻōnaehana i hoʻohana ʻia e pale i nā mea kanu mai nā maʻi fungal he nui ʻole e kūʻē i nā oomycetes.
No laila, i ka pale kemika e pili ana i ka hopena o ke ahi, ua hoʻohana ʻia ka nui (a i 12 mau manawa i kēlā me kēia kau a ʻoi paha) me ka hoʻomākaukau hoʻomākaukau o kahi ʻano ākea o ka hana. ʻO kahi hana kipi ka hoʻohana ʻana i nā phenylamides, he mea ʻino ia i nā oomycetes a hoʻolaha ʻōnaehana i nā mea kanu. Eia nō naʻe, ua alakaʻi wikiwiki ka hoʻohana ākea ʻana i ka hōʻiliʻili ʻana o nā ʻano kūʻē i nā lāhui fungal (Davidse et al., 1981), kahi mea hoʻopīhoihoi i ka pale ʻana o nā mea kanu. ʻO P. infestans ka mea hoʻokahi wale nō parasite o ka ciate temperate, ka ʻino mai hiki ʻole ke hoʻopau ʻia i ka mahiʻai meaola me ka hoʻohana ʻole ʻia o nā ʻano kemika o ka pale (Van Bruggen, 1995).
Hōʻike ka mea i luna i ka nānā nui i nā mea noiʻi mai nā ʻāina like ʻole e uku i ke aʻo ʻana i ka P. infestans heluna kanaka, ka ikaika o kā lākou nui a me ka haku mele ʻana, a me nā ʻano genetics o ka loli.
ʻO ke ola o R. INFESTANS
Hoʻokumu ʻo Oomycete Phytophthora infestans i kahi mycelium intercellular me ka haustoria i loko o nā lau ʻuala. ʻO ka hānai ʻana i nā mea lau, kumu ia ke kūkulu ʻana i nā kiko ʻeleʻele, e ʻeleʻele a pala hoʻi i ka wā pulu. Me ka eo ikaika, make ka lau holoʻokoʻa. Ma hope o kahi manawa o ka hānai ʻana, hana ʻia nā mea ulu i ka mycelium - sporangiophores - e ulu i waho ma o ka stomata. I ke aniau pulu, hana lākou i pua keʻokeʻo a puni nā kiko ma lalo o nā lau. Ma nā wēlau o ka sporangiophores, ua hoʻokumu ʻia ka zoosporangia o ka lemon, i haki a lawe ʻia e ka ʻehu o ka ua (Fig 1). Ke hāʻule nei i loko o nā kulu wai ma luna o kahi lau ʻuala, sporangia germination me 6-8 zoospores, a, ma hope o kahi manawa o ka neʻe ʻana, puni ʻia, uhi ʻia me ka pūpū a ulu me kahi ʻōpuʻu ʻōpuʻu. Hoʻopili ka ʻōpuʻu ma o ka stomata i loko o ka lau lau. Ma lalo o kekahi mau kūlana, hiki i ka sporangia ke ulu i loko o ka paipu ulu pololei i kaʻiʻo lau. Ma lalo o nā kūlana maikaʻi, ʻo ka manawa mai ka maʻi i ka hoʻokumu ʻana i ka sporulation hou he 3-4 mau lā wale nō.
I ka manawa ma ka honua a kānana ʻia ma o ka lepo, hiki i ka sporangia ke hoʻopili i nā tubers. Piʻi nā tubers i loli i ka wā e mālama ai; i ka nāwaliwali, e hoʻomau ka maʻi a hiki i ke kau aʻe. Hoʻohui ʻia, hiki i ka ʻākena causative o ka hopena blight ke hoʻomau i ka hoʻoilo i ke ʻano o nā oospores (nā paia moe kolohe e moe ana) i ka lepo ma nā ʻōpala a me nā hua kōmato. Hoʻokumu ʻia ʻo Oospores ma nā mea ulu o ke ola ke hui ʻia nā ʻano o nā ʻano moena like ʻole me ka momona. I ka puna, kūkulu ʻia ka sporulation asexual ma luna o nā tubers i kanu ʻia a me nā koena mea kanu me nā oospores; komo nā zoospores i ka lepo a hoʻoulu i ka maʻi o nā lau lalo o nā mea kanu. I kekahi mau hihia, hiki i ka mycelium ke ulu aʻe mai ka tuber maʻi ma ka ʻaoʻao ʻōmaʻomaʻo o ka mea kanu a ʻike pinepine ʻia ma ka ʻaoʻao luna o ke koʻokoʻo.
ʻO kahi ʻokoʻa nui ma waena o nā oomycetes a me ka hapanui o nā fungi e waiho ana i ka predominance o diplophase i ko lākou ola pōʻaiapili me ka meiosis gametic a me ka germination o zygotes (oospores) me ka ʻole o ka nukelika reductive. ʻO kēia hiʻohiʻona, a me ka dipolar heterotallism e pani nei i ka bisexuality, me he mea lā e hiki ke hoʻohana i nā oomycetes i nā ala i hoʻomohala ʻia no ke aʻo ʻana i nā lehulehu o nā eukaryotes kiʻekiʻe (ka anamanaʻo o ka panmixia a me ka hoʻokaʻawale o nā heluna kanaka, nā intra- a me nā interpopulation gene flow, a pēlā aku.). Eia naʻe, ʻekolu mau kumu i ʻae ʻole i ka lawe ʻana i kēia mau ala i ke aʻo ʻana i ka P. infestans heluna kanaka.
1. Me nā oospores hybrid, hana ʻia nā oospores momona a me nā parthenogenetic i nā heluna (Fife a me Shaw, 1992; Anikina et al., 1997a; Savenkova, Cherepnikoba-Anirina, 2002; Smirnov, 2003), a lawa paha ke alapine o kā lākou hoʻokumu ʻana i mea e hoʻohuli ai. ma nā hopena hōʻike.
2. ʻO ke kaʻina moekolohe ma P. infestans hāʻawi i kahi kōkua nui ʻole i ka ikaika o ka nui o ka heluna kanaka, no ka mea, hana nui ka fungus i nā spore vegetative, e hana ana ma mua o 90% o nā hopena o ke kālailai ʻana i ka ʻano male e ka hana kuʻuna ma kahi mea momona. ... ke kau e ulu nei i kekahi mau hanauna o ka sporulation asexual (hoʻomohala maʻi polycyclic). He mea nui ko Oospores i ka mālama ʻana i nā meaola i loko o ka wā ke ʻole nā mea kanu ʻōmaʻomaʻo (i ka hoʻoilo) a me ka maʻi nui o nā mea kanu. A laila, i ke kauwela, ka hana hou ʻana o clonal a me ka hoʻonui a i ʻole, ʻo ka hoʻoliʻiliʻi o ka helu o nā klone pākahi i kū aʻe ma muli o ka hopena o ka recombination moekolohe, ka mea i hoʻoholo nui ʻia e ke koho ʻana o nā mea i hoʻopili ʻia. No laila, hiki i ka lakio o kēlā me kēia clone i ka heluna kanaka i ka hoʻomaka a me ka hopena o nā epiphytotics hiki ke ʻokoʻa loa.
3. ʻO ke kaʻapuni i wehewehe ʻia ka hiʻona o nā lāhui maoli o P. infestans i ko lākou homeland, ʻAmelika Hui Pū ʻIa. I nā wahi ʻē aʻe o ka honua, ʻaʻole i ʻike ʻia ke kaʻina hana moekolohe ma mua o 100 mau makahiki; ʻo ka mycelium vegetative i nā tubers potato i hoʻopili ʻia ka wā hoʻoilo. Ua wikiwiki loa ke ola, a he kikowaena ka pālahalaha: ʻo ka maʻi mai nā tubers i kanu ʻia i hoʻopili ʻia i hele i nā lau, e hoʻokumu ana i ka foci mua o ka maʻi, i hiki ke hui pū me ka ulu nui ʻana o ka maʻi.
No laila, i kekahi mau wahi aia paha he kahuli o ka pōʻai kolohe a me ka moe kolohe, ʻoiai ma nā poʻe ʻē aʻe - pōʻaiapili asexual wale nō.
Ke kumu o P. INFESTANS
Ua ʻike ʻia ʻo P. infestans ma ʻEulopa i ka hopena o ka hapa mua o ke kenekulia 1991. ʻOiai ʻo ka ʻuala he ʻōiwi ia ma ka ʻākau hikina o ʻAmelika Hema, ua manaʻo ʻia ua lawe ʻia ka parasite mai ʻEulopa i ka wā o ka paʻakai paʻakai o Chile. Eia nō naʻe, ua hana ʻia nā noiʻi ma ke kahua kūʻuʻu ʻo Rockefeller Center ma ke awāwa ʻo Toluca, Mekiko i koi ʻia e kipa hou ʻia kēia manaʻo (Niederhauser, 1993, XNUMX).
1. Ma ke awāwa ʻo Toluca, he mau ʻāpana potato tuberous kūloko (Solanum demissum, S. bulbocastanum, a me nā mea like ʻole.) ʻO nā lāhui ʻAmelika Hema, e like me nā ʻuala, nele i nā genes kūʻē.
2. Ma ke awāwa ʻo Toluca, ʻike ʻia nā kaʻawale me nā ʻano pilina o A1 a me A2, a ʻo ka hopena e pālahalaha ana ka lehulehu o P. infestans ʻoiai i ka ʻāina ʻōiwi o nā mahi uala i mahi ʻia, ʻAmelika Hema, ua pālahalaha ka parasite.
3. Ma ke awāwa ʻo Toluca, aia kekahi mau maʻi ahulau koʻikoʻi o ka hopena o ke ahi. No laila, i waena o nā kānaka noiʻi ʻAkau ʻAkau ʻAkau (Ke Kulanui ʻo Cornell), ua hoʻokumu ʻia ka manaʻo e pili ana iā Mesoamerica (ʻAmelika Hui Pū ʻIa) ma ke ʻano kahi i hānau ʻia ai ka pottphthora ʻuala (Goodwin et al., 1994).
ʻAʻole ʻimi like nā mea noiʻi ʻAmelika Hema i kēia manaʻo. Manaʻoʻiʻo lākou i ka ʻuala i mahi ʻia a me kāna parasite P. infestans i ka homeland maʻamau - ka South American Andes. Kākoʻo lākou i ko lākou manaʻo e nā noiʻi molekini ma ke kālailai ʻana i nā polymorphism DNA o ka genito mitochondrial (mtDNA) a me nā genas nukle RAS a me β-tubulin (Gomez-Alpizar et al., 2007). Ua hōʻike lākou i ka hōʻiliʻili ʻia o nā ʻano ʻāpana o nā wahi like ʻole o ka honua mai nā laina kūpuna ʻekolu i loaʻa (aia ʻekolu) i loaʻa i nā ʻAmelika Hema ʻAmelika. ʻO nā haplotypes Andean he mau keiki ia o nā lālani ʻelua: hoʻokaʻawale ʻia ka moʻokūʻauhau mtDNA kahiko loa ma Solanaceae hihiu mai ka ʻāpana Anarrhicomenum ma Ecuador, ʻoiai he maʻa mau nā kaʻawale o ka lālani ʻelua ma kaʻuala, ʻōmato, a me nā nighthades hihiu. I Toluca, ʻo nā haplotypes kakaʻikahi e iho mai kahi moʻokūʻauhau hoʻokahi wale nō, me ka loli ʻana o nā ʻano Toluca (ke alapine haʻahaʻa pinepine o kekahi mau pae loli) e hōʻike nei i kahi hopena hoʻokumu ikaika ma muli o ka ʻauheʻe o kēia manawa.
Hoʻohui ʻia, loaʻa kahi ʻano hou P. andina i ka Andes, morphologically a me genetically like me P. infestans, kahi, e like me ka mea kākau, kuhikuhi i ka Andes ma ke ʻano he wahi wela o ke kikoʻī ma ka ʻano Phytophthora. ʻO ka mea hope loa, ma ʻEulopa a me ʻAmelike Hui Pū ʻIa, ʻo P. Infestans ka heluna kanaka e komo pū me nā kūpuna Andes ʻelua, ʻoiai ma Toluca hoʻokahi wale nō.
Ua hōʻeuʻeu kēia paʻi i ka pane mai kahi hui o nā kānaka noiʻi mai nā ʻāina like ʻole, nā mea i hana i nā hana hoʻokolohua he nui e hoʻoponopono hou i ka noi mua (Goss et al., 2014). I kēia hana, ʻo ka mea mua, ua hoʻohana ʻia nā kaʻina DNA microsatellite hou aku e hoʻopaʻa i ka DNA polymorphism; ʻo ka lua, no ke kālailai ʻana i nā puʻupuʻu, nā ala neʻe, ka manawa o ka loli o nā lehulehu, etc. ua hoʻohana ʻia nā hiʻohiʻona hou loa (F-statistics, Bayesian approximations, etc.) a, ke kolu, ua hoʻohana ʻia ka hoʻohālikelike ʻaʻole wale me ka Andean species P. andina, kahi i hoʻokumu ʻia ai kahi ʻano hybrid (P. infestans x Phytophthora sp.) , akā me nā ʻano endemiko Mekiko P. mirabilis, P. Ipomoeae, a me Phytophthora phaseoli - pili genetically P. infestans no ka papa like (Kroon et al., 2012). Ma ke ʻano he hopena o kēia mau anamanaʻo, hōʻikeʻike ʻole ʻia ka ʻāpana o ke kumu lāʻau filogetic o nā ʻano āpau o ka genus Phytophthora i lawe ʻia i loko o ke aʻo ʻana, koe wale no ka hybrid P. andina, no nā ʻano Mekiko, a ʻo ke kahe neʻe ka kuhikuhi ma Mexico - Andes, a ʻo ka ʻaoʻao ʻē, a ʻo ka hoʻomaka me ke ʻano like me ko ʻEulopa. ka noho ʻana o ka Honua Hou (300-600 makahiki i hala). No laila, ke kū ʻana o ka P. infestans species, hana loea no ka lilo ʻana o kaʻuala, i kū i ke kikowaena genetical lua o ka hoʻokumu ʻia ʻana o nā nighthad tuberous, ʻo ia hoʻi ma ʻAmelike Waena.
Genome o P. INFESTANS
I ka makahiki 2009, ua hoʻonohonoho ka hui ʻepekema kūwaho i ka genome P infestans (Haas et al, 2009), a ʻo ka nui o ia mea he 240 MB. ʻOi aku kēia mau manawa ma mua o nā ʻano pili pili like P. sojae (95 Mb), e hoʻoulu ana i nā aʻa aʻa a me P. Ramorum (65 Mb), e hoʻopili ana i nā ʻano kumulāʻau waiwai e like me ka ʻoka, beech, a me kekahi poʻe ʻē aʻe. Ua hōʻike ʻia ka ʻikepili i loaʻa i ka genome ka nui o nā kope o nā kaʻina hana hou - 74%. Aia i loko o ka genome nā genes 17797 protein-coding, ʻo ka hapa nui o lākou nā gen i pili i nā kaʻina hana pūnaewele, e like me ka DNA replication, transcription a me ka unuhi ʻana o nā protein.
Hoʻohālikelike kahi genome o ka genus Phytophthora i kahi hui maʻamau o ka genome, me nā poloka o nā kaʻina o nā genes mālama ʻia, kahi i kiʻekiʻe ai ka nui o ka gen a me nā ʻike o nā kaʻina hana pinepine ʻana, a me nā ʻāpana o kēlā me kēia me nā kaʻina gen ʻole mālama ʻole ʻia, me ka haʻahaʻa o ka ʻāpana haʻahaʻa a me kahi kiʻekiʻe o nā wahi i hana hou ʻia. ʻO nā palaka Conservative no 70% (12440) o nā ʻano āpau P. infestans protein-coding genes. Ma waena o nā poloka conservative, pili pinepine nā genes me kahi mamao waena waena o 604 bp. Ma nā wahi ma waena o nā poloka conservative, ʻoi aku ka mamao o ka intergenic (3700 bp) ma muli o ka hoʻonui ʻana i ka nui o nā mea hana hou. Aia nā genes secretory effector wikiwiki i nā wahi ilihune.
Ua hōʻike ʻia ka hōʻiliʻili ʻana o ka genome P. Infestans ma kahi o ka hapakolu o ka genome e pili ana i nā mea transposable. Loaʻa i ka genome o P. infestans nā ʻohana like ʻole o transposons ma mua o nā genome ʻē aʻe i ʻike ʻia. ʻO ka hapa nui o P. infestans transposons no ka ʻohana Gipo.
Ua ʻike ʻia kahi helu nui o nā ʻohana gen kikoʻī e pili ana i ka pathogenesis ma ka genome P. infestans. ʻO kahi ʻāpana nui o lākou e hoʻopili i nā protein effector e hoʻololi i ka physiology o ka mea kanu hoʻokipa a hāʻawi i kāna maʻi. Hāʻule lākou i ʻelua mau ākea ākea: nā hopena apoplastic, ka mea e hana i nā wahi intercellular (apoplasts), a me nā hopena cytoplasmic, i komo i nā pūnaewele ma o haustoria. Hoʻopili nā hopena Apoplastic i nā enzyme hydrolytic huna e like me proteases, lipase a me glycosylases e luku i nā hunaola mea kanu; nā mea kūpale o nā enzyme pale mea kanu hoʻomalu; a me nā mea ʻino nerotizing e like me Nep1-like proteins (NPLs) a me Pcf e like me nā protein cystine-rich liʻiliʻi (SCRs).
ʻO nā genes effector P. infestans he lehulehu a ʻoi aku ka nui ma mua o nā genes nonpathogenic. ʻO ka mea i ʻike nui ʻia ʻo nā hopena cytoplasmic RXLR a me Crinkler (CNR). ʻO nā hopena cytoplasmic maʻamau o oomycetes he protein RXLR. Loaʻa nā ʻaoʻao effector RXLR āpau i ka hui amino-terminal Arg-XLeu-Arg, kahi ʻo X kahi waikawa amino. Ma ke ʻano he hopena, ua manaʻo ʻia aia he 563 mau gen RXLR ma ka genome P. infestans, ʻo ia ka 60% ʻoi aku ma mua o P. sojae a me P. ramorum. Ma kahi o ka hapalua o nā gen RXLR ma ka genome P. infestans he kikoʻī kikoʻī. Loaʻa i nā hopena RXLR ka nui o nā kaʻina. I waena o lākou, hoʻokahi ʻohana nui a me 150 i ʻike ʻia. ʻAʻole like me ka proteome nui, aia nā genes effector RXLR ma nā ʻāpana ʻilihuna a waiwai hou o ka genome. ʻO nā mea kelepona e hoʻoholo i ka dynamism o kēia mau ʻāina e kōkua i ka recombination i kēia mau ʻāpana.
Ua ʻike ʻia nā hopena Cytoplasmic CRN ma P. infestans transcripts e hoʻopili ana i nā peptides nekrosisʻaʻaʻa. Mai ko lākou loaʻa ʻana, ʻike iki ʻia e pili ana i ka ʻohana o kēia mau hopena. Ua hōʻike ʻia ka ʻikepili o ka genome P. Infestans i kahi ʻohana nui o 196 gen CRN, ʻoi aku ka nui ma mua o P. sojae (100 CRN) a me P. ramorum (19 CRN). E like me RXLRs, CRNs he proteins modular a komo pū ʻia me kahi papa L-LAKAK mālama nui ʻia ʻo N-terminal (50 mau amino acid) a me kahi kikowaena DWL e pili ana i nā genes like ʻole. ʻO ka hapa nui o nā CRN (60%) he peptide hōʻailona.
Ua aʻo ʻia ka hiki o nā CRN like ʻole e hoʻopilikia i nā hana kelepona o ka mea kanu hoʻokipa. I ke kālailai ʻana i nā nekrosis mea kanu, ʻo ka hemo ʻana o nā protein CRN2 i hiki ke ʻike i ka moku ʻo C-terminal me 234 amino acid (nā kūlana 173-407, DXG domain) a me ka make ʻana o ka cell. Ua hōʻike ʻia ka ʻikepili o P. infestans CRN genes ʻehā mau ʻāpana C-terminal, a he mea ia e make ai i ka pūnaewele ma loko o ka mea kanu. Hoʻopili kēia i nā kāʻei DC hou (P. Infestans he 18 mau a me 49 pseudogenes), a ʻo D2 (14 a me 43) a me DBF (2 a me 1) mau ʻāpana e like me nā kinases protein. Mālama ʻia nā protein o nā kāʻei CRN i hōʻike ʻia i kahi mea kanu (i ka loaʻa ʻole o nā peptides hōʻailona) i loko o kahi huna mea kanu a hoʻoulu i ka make o ka cell e kahi ʻano intracellular. ʻO nā kaʻina ʻē aʻe he 255 i loaʻa ʻole nā ʻāpana CRN e like me nā genes.
ʻO ka piʻi ʻana o ka helu a me ka nui o ka RXLR a me CRN effector ʻohana ʻohana ma muli o ka recombination homologous non-allelic a me ka hana hou ʻana o ka ʻenekane. ʻOiai ka mea i loko o ka genome i kahi helu nui o nā ʻeleu paʻa paʻa, ʻaʻohe mea hōʻike pololei o ka hoʻololi o nā gen effector.
Nā ʻano hana i hoʻohana ʻia i ke aʻo ʻana i ka hanana lehulehu
Hoʻokumu ʻia ke aʻo ʻana i ka hanana genetika o nā heluna kanaka i kēia manawa ma ke kālailai ʻana i nā moʻomeheu maʻemaʻe o kāna mau ʻano constituent. Lawe pū ʻia ka anamanaʻo o nā lehulehu me ka hoʻokaʻawale ʻole ʻana i nā mea kanu pono no nā kumu kikoʻī, e laʻa me ke aʻo ʻana i ka huhū o kahi lehulehu a i ʻole ke kū ʻana o nā ʻano kūpilikiʻi e kūʻē i nā fungicides (Filippov et al., 2004; Derevyagina et al., 1999). Pili kēia ʻano noiʻi me ka hoʻohana ʻana i nā kiʻina kūikawā, ka wehewehe ʻana ma ʻō aku o ka palena o kēia loiloi. No ke kālailai hoʻohālikelike o nā ʻano, hoʻohana ʻia kekahi mau ʻano hana ma muli o ke kālailai ʻana i ka hanana DNA a ma ke aʻo ʻana i nā hōʻike phenotypic. Pono ka hoʻopili ʻana o ka heluna kanaka i kahi nui o nā kaʻawale, kahi e hoʻokau ai i kekahi mau koi ma nā ʻano hana i hoʻohana ʻia. Ma ke kūpono, pono lākou e hoʻokō i nā koi aʻe (Cooke, Lees, 2004, Mueller, Wolfenbarger, 1999):
- maʻalahi, maʻalahi e hoʻokō, ʻaʻole koi i nā lilo manawa nui, e hoʻokumu ʻia i nā ʻenehana loaʻa maʻamau (e laʻa me PCR);
- pono e hana i kahi helu lawa o nā hiʻohiʻona marker codominant kūʻokoʻa;
- i kiʻekiʻe reproducibility;
- hoʻohana i ka palena iki o nā kiko e nānā ai;
- e kikoʻī i ka substrate (ʻaʻole pono ka hopena i kēia hopena i ka hopena);
- mai koi aku i ka hoʻohana ʻana i nā kaʻina hana weliweli a me nā kemika ʻino loa.
Minamina, ʻaʻohe ʻano hana e kūlike ana i nā palena ma luna. No kahi noiʻi hoʻohālikelike i nā ʻano i ko mākou manawa, hoʻohana ʻia nā ʻano hana ma ke kālailai ʻana i nā ʻano phenotypic: virulence to potato and Tomato varieties (potato and Tomato lāhui), mating type, spectra of peptidase isoenzymes and glucose-6-phosphate isomerase, and on analysis of DNA DNA: lōʻihi polymorphism ʻāpana palena (RFLP), ka mea i hoʻonui pinepine ʻia me ka hybridization probe RG 57, ka hoʻopili ʻana o nā microsatellite repeats (SSR a me InterSSR), hoʻonui ʻia me nā primers (RAPD), hoʻonui i nā ʻāpana kapu (AFLP), hoʻonui me nā primers homologous i nā kaʻina o nā mea kelepona (ʻo kahi laʻana, Inter SINE PCR), hoʻoholo o nā haplotypes DNA mitochondrial.
Nā wehewehe pōkole o nā hana no ka hoʻohālikelike ʻana i nā ʻano i hoʻohana ʻia me P. Infestans
Nā hiʻohiʻona māka Phenotypic
Heihei "ʻUala"
ʻO nā lāhui "ʻUala" kahi māka noiʻi maʻamau a hoʻohana ʻia. ʻO nā lāhui "ʻuala maʻalahi" i hoʻokahi gen no ka virulence ʻuala, "paʻakikī" - ma ka liʻiliʻi ʻelua. ʻO Black et al. (1953), e hōʻuluʻulu ana i nā ʻike āpau i loaʻa iā lākou, ua ʻike he hiki i ka lāhui phytophthora ke hoʻopili i nā mea kanu me nā gen / resistes gen e pili ana i ka P. infestans virulence gen / genes, a loaʻa nā lāhui 1, 2, 3, a me 4 e hoʻopili i nā mea kanu. me nā genes R1, R2, R3 a me R4, ʻo ia hoʻi, ʻe. ka launa pū ma waena o ka parasite a me ka mea hoʻokipa ma muli o ka ʻāpana no ke ʻano kumu. Eia hou, ʻeleʻele, me ke komo pū ʻana o Gallegly a me Malcolmson, ʻike ʻia nā genes kūʻē R5, R6, R7, R8, R9, R10 a me R11, a me nā lāhui like (ʻEleʻele, 1954; ʻEleʻele & Gallegly, 1957; Malcolmson & Black, 1966; Malcolmson, 1970).
Aia kahi kino nui o ka ʻikepili ma ka ʻano lāhui o ka pathogen mai nā wahi like ʻole. Me ka nānā ʻole ʻana i kēia ʻikepili i ke kikoʻī, e hōʻike wale mākou i kahi maʻamau maʻamau: kahi i hoʻohana ʻia ai nā ʻano me nā genes kūʻē hou a i ʻole ko lākou hui ʻana, i ka manawa mua ua nāwaliwali ka hopena o ke ahi, akā ua ʻike ʻia nā lāhui me nā genulence virulence e pili ana a ua koho ʻia a hoʻomaka hou nā ʻōpala o ke ahi. ʻAʻole ʻike iki ʻia ka virulence kūʻē e kūʻē i nā ʻehā kūʻē ʻehā (R4-R1) i nā hōʻiliʻili i hōʻiliʻili ʻia ma mua o ka hoʻokomo ʻia ʻana i ka mahi ʻana o nā ʻano me kēia mau genes, akā ua piʻi nui ka helu o nā ʻano virulent i ka manawa i hoʻopili ʻia ai ka pathogen i nā ʻano e hāpai ana i kēia mau ʻāpana. ʻO Genes 4-5, ma ka ʻaoʻao ʻē aʻe, maʻa mau i nā hōʻiliʻili (Shaw, 11).
ʻO ke aʻo ʻana i ka lakio o nā lāhui like ʻole i ka wā ulu, i lawe ʻia i ka hopena o 1980, ua hōʻike ʻia i ka hoʻomaka ʻana o ka ulu ʻana o ka maʻi, nā clones me ka haʻahaʻa agriveness a me 1-2 genulence genes i lilo i ka lehulehu.
Eia hou, me ka hoʻomohala ʻana i ka hopena o ke ahi, ua emi ka nui o nā klone kumu a piʻi ka helu o nā heihei "paʻakikī" me ka huhū nui. ʻO ka hanana o ka hope i ka hopena o ke kau i 100%. Ke mālama ʻana i nā paipu, aia ka emi o ka hakakā a me ka nalo ʻana o nā genes virulence pākahi. Hiki i ka dynamics o ka pani clone ke hana i nā ʻano ʻokoʻa i nā ʻano like ʻole (Rybakova & Dyakov, 1990). Eia nō naʻe, ua hōʻike ʻia kā mākou noiʻi ʻana ma 2000-2010 i ka loaʻa ʻana o nā heihei paʻakikī mai ka hoʻomaka mua o nā epiphytotics i waena o nā kaila i hoʻokaʻawale ʻia mai nā ʻuala a me nā kōmato. Hoʻololi paha kēia i ka lehulehu o P. Infestans ma Lūkia.
Ma o 1988-1995, ke alapine o ka hanana ʻana o nā "superraces" me nā genel virulence āpau a ʻaneʻane paha i 70-100%. Ua ʻike ʻia kēlā ʻano kūlana, ma kahi laʻana, ma Belarus, ma nā moku ʻo Leningrad, Moscow, ma North Ossetia a me Kelemania (Ivanyuk et al., 2002a, 2002b; Politiko, 1994; Schober-Butin et al., 1995).
Heihei "Tomato"
I nā ʻano kōmato, he 2 wale nō mau ʻaoʻao o ke kūʻē ʻana i ka hopena o ke ahi i loaʻa - Ph1 (Gallegly & Marvell, 1955) a me Ph2 (Al-Kherb, 1988). E like me ke ʻano o nā heihei ʻuala, ka launa pū ma waena o nā ʻōmato a me P. infestans ma ke kumu kūʻauhau. Hoʻopili ka lāhui T0 i nā ʻanoʻano ʻaʻohe kūʻē kūʻē (ka hapa nui o nā ʻano ʻenehana i hoʻohana ʻia), hoʻopili ka lāhui T1 i nā ʻano me ka gen Ph1 (Ottawa), a ʻo ka lāhui T2 e hoʻopili i nā ʻano me ka ʻāpana Ph2.
Ma Rūsia, ʻaneʻane wale nō e loaʻa ʻo T0 i nā uala; Ua lanakila ʻo T0 ma nā ʻōmato i ka hoʻomaka o ke kau, akā ma hope ua hoʻololi ʻia e ka heihei T1 (Dyakov et al., 1975, 1994). Ma hope o 2000, ua hoʻomaka ʻo T1 ma kaʻuala ma nā lehulehu he nui i ka hoʻomaka mua o ka wā epiphytotic. Ma ʻAmelika Hui Pū ʻIa, ʻaʻole pathogenic ke ʻano o ka ʻuala i ke kōmato, a me nā heihei T0, T1, a me T2, ʻoiai ʻo T1 a me T2 i lanakila ma luna o nā ʻōmato (Vartanian & Endo, 1985; Goodwin et al., 1995).
ʻAno hoʻoipoipo
E alakaʻi i ke aʻo ʻana, nā mea hoʻāʻo (kūmole) me nā ʻano moena ʻike ʻia - koi ʻia ʻo A1 a me A2. Hoʻokomo ʻia ka hoʻowalewale hoʻāʻo me lākou i mau pālua i nā ipu Petri me ka meat agar. Ma hope o ka hoʻouluulu ʻana no 10 mau lā, nānā ʻia nā pā no ka loaʻa a me ka ʻole o nā oospores i ka meahana i ka wā hoʻopili o nā ʻano. Aia nā koho 4: no ka A1 ʻano moena ke ʻano o ke kanana, inā he oospores me ka mea hoʻāʻo A2, iā A2, inā he oospores me ka mea hoʻāʻo A1, iā A1A2, inā he oospores me nā mea hōʻike ʻelua, a i ʻole sterile (00), inā ʻaʻole ia he oospores. me ka mea hoʻāʻo ʻole (kākaʻikahi nā hui hope ʻelua).
I mea e hoʻoholo koke ai i nā ʻano moʻa, ua hoʻāʻo e ʻike i nā wahi o ka genome e pili ana me ke ʻano o ka moʻa ʻana, me ka pahuhopu o kā lākou hoʻohana hou ʻana e hoʻoholo i ke ʻano o ka male ʻana e PCR. ʻO kekahi o nā hoʻokolohua kūleʻa mua e ʻike ai i kēlā pūnaewele i alakaʻi ʻia e nā mea noiʻi ʻAmelika (Judelson et al., 1995). I ka hoʻohana ʻana i ka hana RAPD, ua hiki iā lākou ke hoʻomaopopo i ka moku ʻo W16 e pili ana me ka moena o nā keiki o nā kaʻawale ʻelua, a e hoʻolālā i kahi pālua o 24-bp primers e hoʻonui iā ia (W16-1 (5'-AACACGCACAAGGCATATAAATGTA-3 ') a me W16-2 (5' -GCGTAATGTAGCGTAACAGCTCTC-3 ') Ma hope o ka palena o ka huahana PCR me ka enzyme kaohi HaeIII, ua hiki ke hoʻokaʻawale i nā kaʻawale me nā ʻano pālua A1 a me A2.
ʻO kahi hoʻāʻo ʻē aʻe e kiʻi i nā māka PCR e hoʻoholo i nā ʻano o ka moʻa ʻana e nā mea noiʻi Kōlea (Kim, Lee, 2002). Ua ʻike lākou i nā huahana kikoʻī e hoʻohana ana i ka hana AFLP. A ʻo kahi hopena, ua hoʻomohala ʻia kekahi mau primers PHYB-1 (mua) (5'-GATCGGATTAGTCAGACGAG-3 ') a me PHYB-2 (5'-GCGTCTGCAAGGCGCATTTT-3'), e ʻae ana i ka hoʻonui ʻana i ka ʻāpana genome e pili ana me ka A2 moʻa. Ma hope iho, ua hoʻomau lākou i kēia hana a hoʻolālā i nā primers 5 'AAGCTATACTGGGACAGGGT-3' (INF-1, i mua) a me 5'-GCGTTCTTTCGTATTACCAC-3 '(INF-2), e ʻae ana i ka hoʻonui ʻana i ka ʻāpana Mat-A1 ʻano o nā ʻano me nā ʻano moena A1. ʻO ka hoʻohana ʻana o ka diagnostics PCR o nā ʻano moena i hōʻike i nā hopena maikaʻi i ke aʻo ʻana o P. infestans heluna kanaka ma Czech Republic (Mazakova et al., 2006), Tunisia (Jmour, Hamada, 2006), a me nā wahi ʻē aʻe. I loko o kā mākou keʻena hoʻokolohua (Mytsa, Elansky, ʻaʻole i paʻi ʻia), 34 P. infestans i hoʻokaʻawale ʻia mai nā ʻuala maʻi a me nā ʻōmato ma nā wahi like ʻole o Lūkia (Kostroma, Ryazan, Astrakhan, Moscow oblasts). ʻO nā hopena o ka hoʻopili ʻana o PCR me ka hoʻohana ʻana i nā primers kikoʻī ma mua o 90% i kūlike me nā hopena o ke kālailai ʻana i ka ʻano moena e ka hana kuʻuna ma kahi mea momona.
Papa 1. Ka loli o ke kūʻē ʻana ma waena o ka clone Sib 1 (Elansky et al., 2001)
Kahi wahi hōʻiliʻili laʻana | Helu o nā kaʻawale i kālailai ʻia | Ka helu o nā ikepili koʻikoʻi (S), nāwaliwali kūpale (SR) a me nā kūʻē kū (R), pcs (%) | ||
S | SR | R | ||
G. Vladivostok | 10 | 1 (10) | 4 (40) | 5 (50) |
G. Chita | 5 | 0 | 0 | 5 (100) |
ʻO Irkutsk | 9 | 9 (100) | 0 | 0 |
G. Krasnoyarsk | 13 | 12 (92) | 1 (8) | 0 |
Ke kūlanakauhale ʻo Yekaterinburg | 15 | 8 (53) | 1 (7) | 6 (40) |
O. Sakhalin | 66 | 0 | 0 | 66 (100) |
ʻĀina ʻo Omsk | 18 | 0 | 0 | 18 (100) |
ʻO ke kūpaʻa Metalaxyl ma ke ʻano he lehulehu
I nā makahiki 1980 i hala mua, ua ʻike ʻia nā maʻi ikaika o ka hopena o ka hopena i hopena ʻia e nā metalaxyl-resist P. infestans i nā wahi like ʻole. ʻO nā māla potato ma nā ʻāina he nui i loaʻa i nā pohō nui (Dowley & O'Sullivan, 1981; Davidse et al., 1983; Derevyagina, 1991). Mai ia manawa, i nā ʻāina he nui o ka honua, mālama ʻia ka nānā mau ʻana i ka hanana o nā ʻano kūʻē phenylamide i P. infestans heluna kanaka. Ma waho aʻe o kahi loiloi kūpono o nā prospect no ka hoʻohana ʻana i nā lāʻau i loko o ka phenylamide, ke kūkulu ʻana i kahi ʻōnaehana pale a me ka wānana ʻana i nā epiphytoties, ua lilo ke kūʻē ʻana i kēia mau lāʻau i hoʻokahi o nā māka marker i hoʻohana ākea ʻia no ka loiloi hoʻohālikelike o nā heluna o kēia pathogen. Eia nō naʻe, pono e hoʻokō ʻia ke kūpaʻa ʻana i ka metalaxyl i ka hoʻohālikelike ʻana i ka nui o ka heluna kanaka me ka noʻonoʻo ʻana i ka ʻoiaʻiʻo: 1 - ʻaʻole i hoʻoholo pono ʻia ke kumu o ke kūʻē. 2 - ke kūʻē ʻana i ka metalaxyl kahi ʻano hilinaʻi i wae ʻia e hiki ke loli i ka hoʻohana ʻana o nā phenylamides, 3 - ʻokoʻa ke kēkelē o ka noʻonoʻo i nā ʻano metalaxyl ma loko o hoʻokahi laina clonal (papa. 1).
ʻO Spectra o isozymes
Kū kaʻawale nā māka Isozyme i nā kūlana kūwaho, hōʻike i ka hoʻoilina Mendelian a he codominant, e ʻae ana e hoʻokaʻawale ma waena o homo- a me heterozygotes. ʻO ka hoʻohana ʻana o nā protein e like me nā māka gen hiki ke hoʻomaopopo i nā hoʻoponopono hou ʻelua o nā mea kūpuna, e like me nā chromosomal a me nā genomic mutations, a me nā mea hoʻokahi amino acid substitutions.
Ua hōʻike ʻia nā noiʻi electrophoretic o nā protein i ka nui o nā enzyme i loko o nā meaola i ke ʻano o nā ʻāpana i ʻokoʻa i ka neʻe electrophoretic. ʻO kēia mau hakina ka hopena o ka hoʻopāʻālua ʻana i nā ʻano hana o nā enzyme e nā loci ʻokoʻa (isozymes a i ʻole isozymes) a i ʻole nā ʻaleme like ʻole o ka locus like (allozymes a alloenzymes). ʻO ia, isozymes nā ʻano like ʻole o hoʻokahi enzyme. Loaʻa i nā ʻano ʻokoʻa ka hana catalytic like, akā ʻokoʻa iki i nā substitutions amino acid hoʻokahi i ka peptide a me ka mālama. Hōʻike ʻia ia ʻano ʻokoʻa i ka wā o ka electrophoresis.
Ke aʻo nei i nā ʻano P. infestans, ke kiʻiona o isoenzymes o ʻelua protein, peptidase a me glucose-6-phosphate isomerase, i hoʻohana ʻia (ʻo kēia enzyme he monomorphic i nā heluna Lūkini; no laila, ʻaʻole hōʻike ʻia nā ʻano hana o kāna noi i kēia hana). E hoʻokaʻawale iā lākou i nā isozymes i loko o ka māla uila, hoʻomākaukau ka hoʻomākaukau protein i hoʻokaʻawale ʻia mai nā meaola i hoʻopaʻa ʻia i kahi papa gel i hoʻonoho ʻia i kahi māla uila. ʻO ka helu o ka diffusion o kēlā me kēia protein i ka gel e pili ana i ka uku a me ke kaumaha o ka mole. No laila, i kahi māla uila, hoʻokaʻawale ʻia ka hui ʻana o nā protein i loko o nā hakina, a hiki ke ʻike ʻia me nā kala ʻokoʻa.
Lawe ʻia ke aʻo ʻana i nā isoenzymes peptidase ma ka cellulose acetate, starch a i ʻole nā polyacrylamide gels. ʻO ke kūpono loa ke ʻano e pili ana i ka hoʻohana ʻana o nā gul acetate cellulose i hana ʻia e Helena Laboratories Inc. ʻAʻole ia e koi i nā mea nui o nā mea hoʻāʻo, ʻae ia i kekahi e loaʻa i nā band contrasting ma ka gel ma hope o electrophoresis no nā loci enzyme ʻelua, ʻaʻole koi ʻia ka hoʻokō ʻana i ka manawa nui a me nā kumukūʻai waiwai (Fig 2).
Hoʻololi ʻia kahi ʻāpana mycelium i loko o kahi microtube 1,5 ml, hoʻohui ʻia he 1-2 mau kulu wai hoʻomaʻemaʻe iā ia. Ma hope o kēlā, homogenized ka hāpana (no ka laʻana, me kahi drill uila me kahi mea hoʻopili uila i kūpono no ka microtube) a hoʻoliʻiliʻi no 25 kekona ma kahi centrifuge ma 13000 rpm. 8 μl mai kēlā me kēia microtube. hoʻoili ʻia ka supernatant i ka pā applicator.
Lawe ʻia ka gel cellulose acetate mai ka pahu buffer, holoi ʻia ma waena o ʻelua pepa o ka pepa kānana a kau ʻia me ka papa hana ma luna o ke kumu ea o ka mea noi. Hoʻololi ʻia ka hopena mai ka pā e ka mea noi i ka gel 2-4 mau manawa. Lawe ʻia ka gel i kahi lumi electrophoresis,
Pākaukau 2. ʻO ke ʻano o ka hopena i hoʻohana ʻia no ka pale ʻana i ka gel cellulose acetate i ke kālailai ʻana i nā isoenzymes peptidase, kau ʻia kahi kulu o ka pena (bromophenol blue) ma ka lihi o ka gel.
TRIS HCl, 0,05M, Ph 8,0 2 ml
Peroxidase, 1000 U / ml 5 mau kulu
o-dianisidine, 4 mg / ml 8 kulu
MgCl2, 20 mg / ml 2 kulu
ʻO Gly-Leu, 15 mg / ml 10 mau kulu
L-amino-acid oxidase, 20 u / ml 2 mau kulu
Lawe ʻia ʻo Electrophoresis no 20 mau minuke. ma 200 V. Ma hope o electrophoresis, lawe ʻia ka gel i kahi papa pena kiʻi a paʻi ʻia me kahi hopena pena kūikawā (Papa 2). 10 ml o 1,6% DIFCO agar i hoʻoheheʻe ʻia i loko o ka umu umu, anuanu i 60 ° C, a ma hope iho ua hui ʻia ʻo 2 ml o ka agar me kahi kāwili pena a ninini ʻia i ka gel. Hōʻike nā kaha ma waena o 15-20 mau minuke. Hoʻohui ʻia ka L-amino-acid oxidase reagent ma mua pono o ka hoʻohuihui ʻana i ka hopena me ka agar i hoʻoheheʻe ʻia.
I nā heluna Lūkini, hōʻike ʻia ka Pep 1 locus e nā genotypes 100/100 a me 92/100. Kākaʻikahi loa ʻo Homozygote 92/92 (ma kahi o 0,1%). Ua hōʻike ʻia ʻo Locus Rehr 2 e ʻekolu mau genotypes 100/100, 100/112, a me 112/112, a ʻo nā ʻano ʻokoʻa āpau he 3 (Elanky a me Smirnov, 2003, Fig 2).
ʻImi genome
ʻO ka palena palena polymorphism lōʻihi me ka hybridization e hiki mai ana (RFLP-RG 57)
Mālama ʻia ka DNA āpau me ka Eco R1 restriction enzyme, hoʻokaʻawale ʻia nā ʻāpana DNA e electrophoresis i ka agarose gel. Nui loa ka DNA Nuclear a he nui nā kaʻina repetitive, kahi mea paʻakikī e kālailai pololei i nā ʻāpana he nui i loaʻa i ka hana o nā enzyme kaohi. No laila, hoʻoneʻe ʻia nā ʻāpana DNA i hoʻokaʻawale ʻia i ka gel i kahi membrane kūikawā a hoʻohana ʻia no ka hybridization me ka RG 57 probe, i hoʻopili ʻia me nā nucleotide i lepili ʻia me nā lekiō radioactive a i ʻole fluorescent. Hoʻopili kēia probe me nā kaʻina genomic repetitive (Goodwin et al., 1992, Forbes et al., 1998). Ma hope o ka ʻike ʻana i nā hopena o ka hybridization ma kahi kukui- a i ʻole kekahi mea radioactive, loaʻa kahi pilikino hybridization multi-locus (fingerprinting), i hōʻike ʻia e nā ʻāpana 25-29 (Forbes et al., 1998). E loaʻa i nā keiki ʻAsekual (clonal) nā like like. Ma ka hoʻonohonoho ʻana o nā kaula ma ka electrophoretogram, hoʻokolokolo ʻia nā mea like a me nā ʻokoʻa o nā meaola i hoʻohālikelike ʻia.
Nā haplotypes DNA Mitochondrial
I ka hapanui o nā hunaola eukaryotic, hōʻike ʻia ʻo mtDNA i ke ʻano o ka mole DNA ākea pālua, ka mea, ʻokoʻa ka chromosome nukelea o nā hunaola eukaryotic, replicates semi-conservative a pili ʻole i nā mole protein.
Ua hoʻokaʻawale ʻia ka genome mitochondrial o P. infestans, a ua hoʻolaʻa ʻia kekahi mau hana i ke kālailai ʻana i nā lōʻihi ʻāpana kapu (Carter et al, 1990, Goodwin, 1991, Gavino, Fry, 2002). Ma hope o Griffith a me Shaw (1998) hoʻomohala i kahi ala maʻalahi a wikiwiki hoʻi no ka hoʻoholo ʻana i nā haplotypes mtDNA, ua lilo kēia māka i kekahi o nā mea i makemake nui ʻia ma nā noi ʻo P. Infestans. ʻO ke kumumanaʻo o ka hana i loko o ka hoʻonui ʻia ʻana o nā ʻāpana DNA mitochondrial (mai ka genome maʻamau) me nā primers F2-R2 a me F4-R4 (Papa 3) a me kā lākou kapu ma hope me nā palena enzymes MspI (1st fragment) a me EcoR1 (2nd fragment). ʻAe ke ala iā ʻoe e ʻike i nā haplotypes 4: Ia, IIa, Ib, IIb. ʻOkoʻa ʻokoʻa II mai ka ʻano I e ke kū ʻana o kahi hoʻokomo o 1881 bp a me kahi ʻokoʻa o nā wahi kapu ma nā ʻāpana P2 a me P4 (Fig 3).
Mai ka 1996, i waena o nā kaha i hōʻiliʻili ʻia ma ka panalāʻau o Rūsia, ua ʻike wale ʻia nā haplotypes Ia a me IIa (Elansky et al., 2001, 2015). Hiki iā lākou ke ʻike ma hope o ka hoʻokaʻawale ʻana i nā huahana kapu me ka primer F2-R2 i kahi kahua uila (Fig 4, 5). Hoʻohana ʻia nā ʻano mtDNA i ka hoʻopili hoʻohālikelike o nā ʻano a me nā heluna. I kekahi mau ʻano noiʻi, ua hoʻohana ʻia nā ʻano mitochondrial DNA e hoʻokaʻawale i nā laina clonal a passportize iā P. infestans isolates (Botez et al., 2007; Shein et al., 2009). Ke hoʻohana nei i ka hana PCR-RFLP, ua hoʻoholo ʻia he heterogeneous ka mtDNA i ka like P. infestans ʻino (Elansky and Milyutina, 2007). Nā kūlana hoʻonui: 1x (500 kek. 94 ° C), 40x (30 kek. 90 ° C, 30 kek. 52 ° C, 90 kek. 72 ° C); 1x (5 min. 72 ° C). Hoʻohui hoʻohui: (20 μl): 0,2 U Taq DNA polymerase, 1x 2,5 mM MgCl2-Taq buffer, 0,2 mM kēlā me kēia dNTP, 30 pM primer a me 5 ng o ke kālailai ʻia DNA, wai deionized - a 20 μl.
Lawe ʻia ka palena o ka huahana PCR no 4-6 mau hola ma kahi mahana o 37 ° C. Pākuʻi palena (20 μl): 10x MspI (2 μl), 10x pā palena palena (2 μl), wai deionized (6 μl), huahana PCR (10 μl).
Pakuhi 3. Hoʻohana ʻia nā Primers no ka hoʻonui ʻana i nā wahi polymorphic mtDNA
Locus | Kinohi | ʻO ka lōʻihi Primer a me ke kau ʻana | Ka lōʻihi o ka huahana PCR | Kaohi |
---|---|---|---|---|
P2 | F2: 5'- TTCCCTTTGTCCTCTCCGAT | 21; 13619-13639 | 1070 | MspI |
R2: 5'- TTACGGCGGTTTAGCACATACA | 22; 14688-14667 | |||
P4 | F4: 5'- TGGTCATCCAGAGGTTTGGTT | 22; 9329-9350 | 964 | EcoRI |
R4:5 - CCGATACCGATACCAGCACCAA | 22; 10292-10271 |
Hoʻonui ka hoʻonui mua ʻana (RAPD)
Ke lawe nei i ka RAPD, hoʻohana ʻia kahi primer (i kekahi manawa kekahi mau primers i ka manawa hoʻokahi) me ke kaʻina nucleotide kūwaho, ʻo ka maʻamau he 10 nucleotides i ka lōʻihi, me kahi kiʻekiʻe (mai 50%) o GC nucleotides a me kahi mahana annealing haʻahaʻa (ma kahi o 35 ° C). ʻO kēlā mau primers "pae ʻāina" ma nā pūnaewele kōkua lehulehu i loko o ka genome. Ma hope o ka hoʻonui ʻana, loaʻa he nui o nā amplicons. Pili kā lākou helu i nā primer (s) i hoʻohana ʻia a me nā ʻano hopena (MgCl2 konsentration and annealing temperature).
Lawe ʻia ka ʻike ʻana i nā amplicons e ka distillation ma polyacrylamide a i ʻole agarose gel. Ke mālama nei i ka hoʻopili RAPD, pono e nānā pono i ka maʻemaʻe o nā mea i kālailai ʻia, no ka mea hiki i ka haumia me nā mea ola ʻē aʻe ke hoʻonui i ka nui o nā mea i hana ʻia, i lehulehu loa i ka nānā ʻana i nā mea maʻemaʻe (Perez et al, 1998). ʻO ka hoʻohana ʻana o kēia hana i ke aʻo ʻana i ka genome P. infestans i hōʻike ʻia i nā hana he nui (Judelson, Roberts, 1999, Ghimire et al., 2002, Carlisle et al., 2001). ʻO ke koho ʻana i nā kūlana a me nā primers (51 10-nucleotide primers i aʻo ʻia) hāʻawi ʻia i loko o ka ʻatikala e Abu-El Samen et al., (2003).
Ka Ikepili Repeat Repeat (MOSR)
Hoʻomaopopo hou ʻia ka microsatelit (nā kaʻina o nā kaʻina maʻalahi, SSR) i nā kaʻina pōkole o 1-3 (i kekahi manawa a i 6) nā nucleotide i loko o nā nukelea genome o nā eukaryotes āpau. Hiki ke loli ka helu o nā hana hou i kū mai 10 a 100. Hiki i nā loci microsatelit me kahi alapine kiʻekiʻe loa a ʻoi aku a liʻiliʻi paha i kaʻana like ʻia ma loko o ka genome (Lagercrantz et al., 1993). Pili nā Polymorphism o nā kaʻina microsatelit me nā ʻokoʻa i ka helu o nā hana hou o ke kumu kumu. He codominant nā māka microselit, a hiki iā lākou ke hoʻohana iā lākou no ke kālailai ʻana i ke ʻano o ka heluna kanaka, ka hoʻoholo ʻana i ka hanauna, nā huakaʻi neʻe genotype, a pēlā aku. Ma waena o nā mea maikaʻi ʻē aʻe o kēia mau māka, pono i kekahi e hoʻomaopopo i ko lākou polymorphism kiʻekiʻe, reproducibility maikaʻi, neutrality, a me ka hiki ke hana aunoa a me ka loiloi. Hana ʻia ka analysis o polymorphism o nā microsatelit e ka hoʻonui ʻana PCR me ka hoʻohana ʻana i nā primers i kākoʻo i nā kaʻina kūikawā e pili ana i nā microsatelit loci. I ka mua, ua hana ʻia ke kālailai me ka hoʻokaʻawale ʻana i nā huahana hopena ma kahi gel polyacrylamide. Ma hope mai, ua noi nā limahana o nā Biosystems Applied e hoʻohana i nā primers fluorescently me ka ʻike ʻana i nā huahana hopena e hoʻohana ana i kahi mea ʻike laser maʻamau (Diehl et al., 1990), a laila nā mea hoʻonoho DNA maʻamau maʻamau (Ziegle et al., 1992). ʻO ka lepili ʻana i nā primers me nā pena fluorescent ʻē aʻe e hiki ai ke kālailai i kekahi mau māka ma ka manawa hoʻokahi ma hoʻokahi ala a, no laila, hoʻonui nui i ka hana o ke ʻano a hoʻonui i ka pololei o ke kālailai.
ʻO nā puke pai mua i hoʻolaʻa ʻia i ka hoʻohana ʻana o ka anamanaʻo SSR no ke aʻo ʻana o P. infestans i kū mai i nā makahiki 2000 mua. (Knapova, Gisi, 2002). ʻAʻole hōʻike nā māka āpau e nā mea kākau i kahi kekele o ka polymorphism lawa, eia nō naʻe, ʻelua o lākou (4B a me G11) i hoʻokomo ʻia i loko o ka set of 12 SSR markers i hoʻolālā ʻia e Lees et al. (2006) a laila lawe ʻia i loko o ka pūnaewele noiʻi Eucablight (www.eucablight .org) ma ke ʻano he kūlana no P. infestans. I kekahi mau makahiki ma hope mai, ua paʻi ʻia kahi noiʻi e pili ana i ka hoʻokumu ʻana i kahi ʻōnaehana no ka hoʻonui multiplex o P. infestans DNA e pili ana i nā hōʻailona SSR ʻewalu (Li et al., 2010). ʻO ka mea hope loa, ma hope o ka loiloi ʻana i nā māka i hāpai mua ʻia a koho ʻia i ka mea ʻike nui loa o lākou, a me ka optimizing primers, nā lepili fluorescent, a me nā kūlana amplification, ua hōʻike like ka hui like o nā mea kākau i kahi ʻōnaehana o ka ʻanalike multiplex-analitiki, me nā māka 12 (Papa 4; Li et al. , 2013a). Koho ʻia nā primers i hoʻohana ʻia i kēia ʻōnaehana me ka lepili me kekahi o nā māka fluorescent ʻehā (FAM, VIC, NED, PET) no laila ʻaʻole i hoʻopili ʻia nā laulā o nā nui allele o nā primers me nā lepili like.
Ua hana nā mea kākau i ke kālailai ma ka PTC200 amplifier (MJ Research, USA) me ka hoʻohana ʻana i QIAGEN multiplex PCR kits a i ʻole QIAGEN Typeit Microsatelit PCR kits. ʻO 12.5 μL ka nui o ka hoʻohuihui. Penei nā ʻano hoʻonui: no ka QIAGEN multiplex PCR: 95 ° C (15 min), 30x (95 ° C (20 s), 58 ° C (90 s), 72 ° C (60 s), 72 ° C (20 min); no ka QIAGEN Type-it Microsatelit PCR): 95 ° C (5 min), 28x (95 ° C (30 sec), 58 ° C (90 sec), 72 ° C (20 sec), 60 ° C (30 min).
ʻO ka hoʻokaʻawale a me ka ʻike ʻana i nā huahana PCR i lawe ʻia me ka hoʻohana ʻana i kahi ABI3730 aunoa capillary DNA analyzer (Applied Biosystems).
Papa 4. Nā hiʻohiʻona o 12 mau hōʻailona SSR maʻamau i hoʻohana ʻia no ka genotyping o P. Infestans (Li et al., 2013a)
Inoa | Ka helu o nā ʻākela | Laulā nui alleles (bp) | Nā Pila |
PiG11 | 13 | 130-180 | F: NED-TGCTATTTATCAAGCGTGGG R: GTTTTCAATCTGCAGCCGTAAGA |
Pi02 | 4 | 255-275 | F: NED-ACTTGCAGAACTACCGCCC R: GTTTGACCACTTTCCTCGGTTC |
PinfSSR11 | 4 | 325-360 | F: NED-TTAAGCCACGACATGAGCTG R: GTTTAGACAATTGTTTGGGGTCGC |
D13 | 16 | 100-185 | FAM-TGCCCCCTGCTCACTC R: GCTCGAATTCATTTTCAGACTTG |
PinfSSR8 | 4 | 250-275 | F: FAM-AATCTGATCGCAACTGAGGG R: GTTTACAAGATACACACGTCCC |
PinfSSR4 | 7 | 280-305 | F: FAM-PointsTGTTCGAGTATGCGACG R: GTTTCACTTCGGGAGAAAGGCTTC |
Pi04 | 4 | 160-175 | F: VIC-AGCGGCTTTACCGATGG R: GTTTCAGCGGCTGTTTCGAC |
Pi70 | 3 | 185-205 | F: VIC-ATGAAAATACGTCAATGCTCG R: CGTTGGATATTTCTATTTCTTCG |
PinfSSR6 | 3 | 230-250 | F: GTTTTGGTGGGGCTGAAGTTTT R: VIC-TCGCCACAAGATTTATTCCG |
Pi63 | 3 | 265-280 | F: VIC-ATGACGAAGATGAAAGTGAGG R: CGTATTTTCCTGTTATCTAACACC |
PinfSSR2 | 3 | 165-180 | F: PET-CGACTTCTACATCAACCGGC R: GTTGCTTGGACTGCGTCTTTAGC |
Pi4B | 5 | 200-295 | F: PET-AAAATAAAGCCTTTGGTTCA R: GCAAGCGAGGTTTGTAGATT |
Hōʻike ʻia kahi laʻana o ka nānā ʻana i nā hopena i hōʻike ʻia ma Fig. 6. Ua kālailai ʻia nā hopena me ka hoʻohana ʻana i ka polokalamu GeneMapper 3.7 ma ka hoʻohālikelike ʻana i ka ʻike i loaʻa me ka ʻikepili o nā kaʻawale i ʻike ʻia. I mea e hoʻomaʻalahi ai i ka wehewehe ʻana i nā hopena kālailai, pono ia e hoʻokomo i nā kaʻawale kūʻē 1-2 me kahi genotype i ʻike ʻia i kēlā me kēia haʻawina.
Ua hoʻāʻo ʻia ka hana noiʻi noi ʻia ma ka helu nui o nā laʻana kahua, a ma hope iho ua hoʻokō nā mea kākau i nā kūlana ma waena o nā keʻena hoʻokolohua o nā hui ʻelua, The James Hutton Institute (UK) a me Wageningen University & Research (Netherlands), a, me ka hiki ke hoʻohana i nā kāleka FTA maʻamau no ka maʻalahi. hōʻiliʻili a me ka hoʻouna ʻana o P. infestans DNA mau laʻana, i hiki ke kamaʻilio e pili ana i ka hiki i ka hoʻohana kalepa ʻana o kēia ulu. Hoʻohui, kahi kaʻina wikiwiki a pololei hoʻi o genotyping P. infestans hoʻokaʻawale e hoʻohana ana i ka multiplex SSR analysis i hiki ai ke alakaʻi i nā loiloi maʻamau o nā lehulehu o kēia pathogen ma ka pae honua, a me ka hoʻokumu ʻana i kahi waihona honua ma ka hopena o ka blight i loko o ka papahana o ka papahana Eucablight (www.eucablight.org), me , me nā hopena o ke kālailai microsatelit, i hiki ai ke nānā i ka puka a hoʻolaha ʻana o nā genotypes hou a puni ka honua.
Hoʻonui i ka palena palena lōʻihi polymorphism (AFLP). ʻO AFLP (hoʻonui i ka lōʻihi o ka polymorphism lōʻihi) kahi ʻenehana no ka hana ʻana i nā māka mole mole me ka hoʻohana ʻana i nā primers kikoʻī. I ka AFLP, mālama ʻia ʻo DNA me ka hui pū ʻana o ʻelua mau enzymes kapu. Hoʻopili ʻia nā mea hoʻopili kikoʻī i nā wēlau paʻa o nā ʻāpana kapu.
Hoʻonui ʻia kēia mau ʻāpana me ka hoʻohana ʻana i nā primers kākoʻo i ke kaʻina adapter a me ka pūnaewele kapu, a me ka hāpai pū ʻana i hoʻokahi a ʻoi paha mau kumu kaulike ma kā lākou 3 'wēlau. ʻO ka ʻāpana o nā ʻāpana i loaʻa i ka hilinaʻi ʻana i nā enzyme ka palena a me nā nukleote i wae wale ʻia ma nā ʻaoʻao 3'o nā primers (Vos et al., 1995). Hoʻohana ʻia ʻo AFLP - genotyping e hoʻopaʻa wikiwiki i ka ʻano genetika o nā meaola like ʻole.
Hāʻawi ʻia kahi wehewehe kikoʻī o ke ʻano i nā hana a Mueller, Wolfenbarger, 1999, Savelkoul et al., 1999. ʻO ka hana nui e hoʻohālikelike ana i ka hoʻonā ʻana o nā hana AFLP a me SSR i hana ʻia e nā mea noiʻi Kina. Ua aʻo ʻia nā hiʻohiʻona phenotypic a me genotypic o 48 P. infestans i hoʻokaʻawale ʻia i ʻelima mau ʻāpana o Kina ʻĀkau. Ma muli o ka hōʻikeʻike AFLP, ʻewalu mau genotypes DNA ʻokoʻa i ʻike ʻia, ʻokoʻa i nā genotypes SSR, kahi i hōʻike ʻole ʻia ai nā mea ʻokoʻa (Guo et al., 2008).
Hoʻonui me nā primers homologous i nā kaʻina o nā mea kelepona
ʻO nā māka i loaʻa mai nā kaʻina o nā retrotransposons maʻalahi loa ia no ka palapala genetical, ke aʻo ʻana i ka ʻokoʻa a me nā kaʻina hana evolution (Schulman, 2006). Inā hana ʻia nā primers e hoʻopiha i nā kaʻina paʻa o kekahi mau mea kelepona, hiki ke hoʻonui i nā wahi genome ma waena o lākou. I ke aʻo ʻana i ka agus causative of late blight, ke ʻano o ka hoʻonui ʻana i nā wahi genome me ka hoʻohana ʻana i ka primer kōkua i ke kaʻina kumu o ka SINE (Short Interspersed Nuclear Elements) retroazone (Lavrova and Elansky, 2003). Ke hoʻohana nei i kēia hana, ua hōʻike ʻia nā ʻokoʻa a hiki i nā keiki asexual o kahi kaʻawale. E pili ana i kēia, ua hoʻoholo ʻia he kikoʻī loa ke ʻano inter - SINE - PCR a kiʻekiʻe ke ana o ka neʻe ʻana o nā mea SINE ma ka genome Phytophthora.
I ka genome o P. infestans, 12 mau ʻohana o nā retrotransposons pōkole (SINE) i ʻike ʻia; ua nānā ʻia ka ʻano laha o nā retrotransposons pōkole, ua ʻike ʻia nā mea (SINE) i loaʻa i ka genome o P. infestans wale nō (Lavrova, 2004).
Nā hiʻohiʻona o ka hoʻopili ʻana i nā ʻano hana o ka hoʻohālikelike i nā ʻano i nā aʻo heluna kanaka
Ke hoʻolālā ʻana i kahi noiʻi, pono maopopo i kekahi i nā pahuhopu e alualu a hoʻohana i nā ʻano kūpono. No laila, ʻae kekahi mau hana i ka hana ʻana i kahi nui o nā hōʻailona māka kūʻokoʻa, akā i ka manawa like he reproducibility haʻahaʻa lākou a hilinaʻi nui i nā reagents i hoʻohana ʻia, nā ʻano hopena, a me ka haumia o nā mea i hoʻopaʻa ʻia. No laila, i kēlā me kēia haʻawina o kahi hui o nā kānana, pono ia e hoʻohana i kekahi mau kaʻanalike maʻamau (kuhikuhi), akā ʻoiai i kēia hihia, paʻakikī loa nā hopena o kekahi mau hoʻokolohua e hoʻohui.
Aia kēia hui o nā kiʻina hana me RAPD, AFLP, InterSSR, InterSINE PCR. Ma hope o ka hoʻonui ʻana, loaʻa i kahi nui o nā ʻāpana DNA o nā nui like ʻole. He ʻōlelo aʻoaʻo e hoʻohana i kēlā mau ʻenehana inā pono e hoʻokumu i nā ʻokoʻa ma waena o nā kānana pili loa (makua-hānau, ʻano hihiu hihiu, a pēlā aku), a i ʻole nā hihia kahi e koi ʻia ai kahi loiloi kikoʻī o kahi hāpana liʻiliʻi. No laila, hoʻohana ākea ʻia ka hana AFLP i ka palapala palapala moʻomeheu o P. infestans (van der Lee et al., 1997) a me nā noiʻi intrapopulation (Knapova, Gisi, 2002, Cooke et al, 2003, Flier et al, 2003). Kūpono ʻole ia ʻano hana ke hoʻohana ke hana ʻana i nā waihona o nā kānana, no ka mea hiki ʻole ke hoʻohui i ka helu helu o nā hopena ke mālama ʻana i nā loiloi i nā hale hana like ʻole.
ʻOiai ka maʻalahi a me ka wikiwiki o ka hoʻokō ʻana (hoʻokaʻawale ʻana ʻo DNA me ka hoʻomaʻemaʻe ʻole, hoʻonui, ʻike ʻana i nā hopena), koi ʻia kēia ʻano hana i kahi ʻano hana kūikawā no ke kākau ʻana i nā hopena: distillation i ka gel polyacrylamide me nā mea i kapa ʻia (radioactively a luminescent paha) a me ka hōʻike ma hope o ka mea uila a radioactive paha. ʻAʻole kūpono ke ʻano maʻamau o ke kiʻi ethidium bromide agarose gel no kēia mau hana no ka mea hiki i kahi nui o nā ʻāpana DNA o nā nui like ʻole ke fuse.
Nā ʻano ʻē aʻe, i ka ʻaoʻao ʻē, ʻae iā ʻoe e hana i kahi helu liʻiliʻi o nā hiʻohiʻona me ke kiʻekiʻe o ka hana hou ʻana. Hoʻopili kēia pūʻulu i ke aʻo ʻana i nā haplotypes DNA mitochondrial (ʻelua wale nō haplotypes Ia a me IIa i kaulana ʻia ma Rūsia), nā ʻano moena (mahele ʻia ka hapa nui o nā kaʻawale i ʻelua mau ʻano: A2 a me A1, ʻaʻole ʻike ʻia ka SF momona ponoʻī) a me ka peptidase isozyme spectra (ʻelua mau loci Pep2 a me Pep1. , me nā isozymes ʻelua i kēlā me kēia) a me ka glucose-2-phosphate isomerase (i Lūkia ʻaʻohe ʻano no kēia ʻano, ʻoiai ʻo polymorphism koʻikoʻi i ʻike ʻia i nā ʻāina ʻē aʻe o ka honua). He ʻōlelo aʻoaʻo e hoʻohana i kēia mau hiʻohiʻona ke kālailai ʻana i nā hōʻiliʻili, ka hōʻuluʻulu ʻana i nā ʻikepili ʻāina a puni hoʻi. I ke ʻano o ke kālailai ʻana i nā isozymes a me nā haplotypes o ka mitochondrial DNA, hiki ke hana ʻia me ka ʻole o nā ʻano maʻamau, ʻoiai i ke kālailai ʻana i nā ʻano moena, ʻelua mau kaʻawale hoʻokolohua me nā ʻano moena ʻike ʻia.
Hiki i nā ʻano hana a me nā mea hoʻopili ke hoʻopili wale i ka hoʻohālikelike o ka huahana ma ka electrophoretogram; ʻaʻole hiki ke ʻike ʻia ka artifact i kēia ʻano o nā noiʻi.
I kēia manawa, ʻo ka hapa nui o nā heluna ma ka ʻāpana ʻEulopa o Rūsia e hōʻike ʻia e nā ʻano ʻano like ʻole o ka male ʻana (Papa 6), ma waena o lākou he mau kaʻawale me nā ʻano Ia a me IIa o ka mitochondrial DNA (ʻo nā ʻano mtDNA i loaʻa i ka honua i loaʻa ʻole ma Lūkia ma hope o 1993). ʻO nā hiʻohiʻona o pepzidase isozymes e hōʻike ʻia e nā genotypes ʻelua ma ka Pep1 locus (100/100, 92/92 a me ka heterozygote 92/100, a he kakaikahi loa ka genotype 92/92 (<0,3%)) a ʻelua mau genotypes ma ka Pep 2 locus (100/100 , 112/112 a me heterozygote 100/112, me ka genotype 112/112 e kū pinepine nei ma lalo o 100/100, akā pinepine pinepine nō hoʻi).
ʻAʻohe loli i ka laulā o isozymes o glucose-6-phosphate isomerase ma hope o 1993 (ka nalo ʻana o ka laina clonal US-1); ua loaʻa nā 100/100 genotype (Elansky a me Smirnov, 2002) i nā kaʻawale.
ʻO ke kolu o ka hui o nā hana e ʻae i ka loaʻa ʻana o kahi hui o nā hiʻohiʻona māka kūʻokoʻa me ka reproducibility kiʻekiʻe. I kēia lā, hoʻokomo pū kēia pūʻulu i ka probe RFLP-RG57, kahi e hoʻopuka ai i nā ʻāpana 25-29 DNA o nā nui like ʻole. Hiki ke hoʻohana ʻia ʻo RFLP-RG57 i ke kālailai ʻana i nā laʻana a me ka hōʻuluʻulu ʻana i nā ʻikepili. Eia naʻe, ʻoi aku ka pipiʻi o kēia hana ma mua o nā mea ma mua, he manawa nui ia, a koi i kahi nui lawa o DNA i hoʻomaʻemaʻe nui ʻia. No laila, koi ʻia ka mea noiʻi e kaupalena i ka nui o nā mea i hoʻāʻo ʻia.
ʻO ka hoʻomohala ʻana o RFLP-RG57 i nā 90 mau mua o ke kenekulia i hala i hoʻonui i ka noiʻi heluna kanaka o ka causative agent o ka hopena o ka maʻi. Ua lilo ia i kumu o ke kiʻina hana e pili ana i ke koho ʻana a me ka hōʻiliʻili ʻana o nā "laina Clonal" (e nānā ma lalo). Me RFLP-RG57, ʻano moena, DNA fingerprinting (ʻano RFLP-RG57), hōʻikeʻike o peptidase a me glucose-6-phosphate isomerase isoenzymes, a me ka ʻano mitochondrial DNA e hoʻohana ai e ʻike i nā laina clonal. Mahalo iā ia, ua hōʻike ʻia ia al., 1994), ke pani o nā heluna kahiko me nā mea hou (Drenth et al, 1993, Sujkowski et al, 1994, Goodwin et al, 1995a), ua hōʻike ʻia nā laina clonal e kū nei ma nā ʻāina he nui o ka honua. Ua hōʻike ʻia nā noiʻi o nā luhi Lūkini e hoʻohana ana i kēia hana i kahi polymorphism genotypic kiʻekiʻe o nā ʻano o ka ʻāpana ʻEulopa a me ka monomorphism o nā lehulehu o nā ʻāpana ʻAsia a me ka Hikina Hikina o Rūsia (Elansky et al, 2001). A i kēia manawa ke ʻano nui nei kēia hana i ka noiʻi heluna kanaka o P. infestans. Eia nō naʻe, keakea ʻia kāna hoʻolaha ākea e kāna kumukūʻai kiʻekiʻe a me ka ikaika o ka hana i ka hoʻokō ʻana.
ʻO kekahi ʻenehana hoʻohiki ʻē aʻe e hoʻohana pinepine ʻole ʻia i ka P. infestans haʻawina he microsatelitet repe (SSR) analysis. I kēia manawa, hoʻohana ākea ʻia kēia hana e hoʻokaʻawale i nā laina clonal. No ke kālailai ʻana i nā kānana, nā ʻano marker phenotypic e like me ke alo o nā genul virulence i nā ʻano potato (Avdey, 1995, Ivanyuk et al., 2002, Ulanova et al., 2003) a me ka ʻōmato i hoʻohana nui ʻia (a hoʻomau e hoʻohana ʻia). I kēia manawa, ua nalowale ka waiwai o nā genes no ka virulence i nā ʻano potato ma ke ʻano he māka marker no ka noiʻi heluna kanaka ma muli o ke ʻano o ka nui (a i ʻole kokoke iā ia) helu o nā genes virulence i ka hapa nui o nā kaʻawale. I ka manawa like, hoʻohana ʻiʻo ʻia ka gen virulence T1 no ka hoʻomaʻamaʻa ʻana i ke kōmike Ph1 e like me ke ʻano o ka marker (Lavrova et al., 2003; Ulanova et al., 2003).
I nā noiʻi he nui, hoʻohana ʻia ke kūpaʻa fungicide ma ke ʻano he māka. ʻAʻole makemake ʻia kēia ʻano e hoʻohana i nā noiʻi heluna kanaka ma muli o ka maʻalahi o ka hoʻololi ʻana o ke kūpaʻa ʻana i nā laina clonal ma hope o ka noi ʻana o metalaxyl- (a mefenoxam-) i loaʻa nā fungicides ma ka pā. ʻO kahi laʻana, ua hōʻike ʻia nā ʻokoʻa nui i ke kiʻekiʻe o ke kūpaʻa ma loko o ka laina clonal Sib1 (Elansky et al., 2001).
No laila, ke ʻano moena, ka peptidase isoenzyme spectrum, mitochondrial DNA type, RFLP-RG57, SSR nā hiʻohiʻona marker makemake ʻia no ka hoʻokumu ʻana i nā panakō ʻikepili a me nā lepili lepili i nā hōʻiliʻili. E hoʻohālikelike i nā laʻana palena palena, inā pono e hoʻopili i ka helu nui o nā hiʻohiʻona māka, hiki iā ʻoe ke hoʻohana iā AFLP, RAPD, InterSSR, PCR Inter-SINE (Papa 5). Eia nō naʻe, pono e hoʻomanaʻo ʻia he hana maikaʻi ʻole kēia mau ʻano hana, a i kēlā me kēia hoʻokolohua (amplification electrophoresis cycle) pono ia e hoʻohana i kekahi mau kaʻawale kaʻawale.
Papa 5. Hoʻohālikelike o nā ʻano ʻokoʻa o ka noiʻi ʻana i nā kānana P. infestans
Criterion | TS | Kāpena Isofer | MtDNA | RFLP-RG57 | RAPD | ISSR | SSR | AFLP | Hōʻike |
---|---|---|---|---|---|---|---|---|---|
Ka nui o ka ʻike | Н | Н | Н | С | В | В | С | В | В |
Reproducibility | В | В | В | В | Н | Н | С | С | С |
Hiki o nā mea hana | Н | Н | Н | Н | В | С | Н | С | В |
lilo o | Н | С | Н | В | Н | Н | Н | С | Н |
Ikaika hana | Н | Н | Н | В | NS * | NS * | Н | С | NS * |
Ka wikiwiki o ka ʻānekal ** | В | Н | Н | С | Н | Н | Н | Н | Н |
Kaha: H - haʻahaʻa, C - waena, B - kiʻekiʻe; НС * - haʻahaʻa ka ikaika o ka hana i ka hoʻohana ʻana i ka agarose gel a i ʻole akomi
genotyper, medium - e ka distillation i ka gel polyacrylamide me nā primers i lepili ʻia,
** - me ka helu ʻole ʻana i ka manawa e lilo ai i ka ulu ʻana o ka mycelium no ka hoʻokaʻawale ʻana iā DNA.
Kūlana lehulehu
Nā laina Clonal
I ka loaʻa ʻole o ka hoʻohui hou ʻana a i kāna kōkua nui ʻole paha i ka hanana heluna kanaka, ʻo ka heluna kanaka kekahi helu o nā klone, nā kūkaʻi kūʻē ma waena o nā mea maʻamau loa.
I loko o ia mau heluna kanaka, ʻoi aku ka ʻike o ka hoʻopaʻa ʻana ʻaʻole i nā alapine o nā genes pākahi, akā nā alapine o nā genotypes i loaʻa kahi kumu maʻamau (nā laina clonal a i ʻole nā laina clonal) a ʻokoʻa wale nō e nā hoʻololi mutation. ʻO nā loiloi heluna kanaka o ka hopena o ka maʻi pathogen a me ke kālailai ʻana i nā laina clonal i hōʻeleu nui ʻia mai ka hiki ʻana mai o ka hana RFLP-RG57 i nā makahiki 90 mua. Me RFLP-RG57, ke ʻano o ka moena ʻana, ka nānā ʻana o peptidase a me ka glucose-6-phosphate isomerase isoenzymes, a me ka ʻano mitochondrial DNA e hoʻohana ai e ʻike i nā laina clonal. Hōʻike ʻia nā ʻano o nā laina clonal maʻamau i ka Papa 6.
Ua hoʻomalu ʻo Clone US-1 i nā lehulehu ma nā wahi āpau a hiki i ka hopena o nā 80, a ma hope ua hoʻomaka ʻia e hoʻololi ʻia e nā klone ʻē aʻe a nalowale mai ʻEulopa a me ʻAmelika Hema. ʻIke ʻia ia i kēia manawa ma ka Hikina Hikina (Philippines, Taiwan, Kina, Iapana, Kōlea, Koh et al., 1994, Mosa et al, 1993), ma ʻApelika (Uganda, Kenya, Rwanda, Goodwin et al, 1994, Vega-Sanchez et al., 2000; Ochwo et al., 2002) a ma ʻAmelika Hema (Ecuador, Brazil, Peru, Forbes et al., 1997, Goodwin et al., 1994). ʻAʻole i ʻike ʻia kekahi mau ʻano o ka laina US-1 ma Australia wale nō. ʻOiai, ua hele mai kahi kaʻawale o P. infestans i Australia me kahi nalu o ka neʻe ʻana (Goodwin, 1997).
Ua neʻe ʻo Clone US-6 mai ka ʻākau o Mexico a Kaleponi i ka hopena o nā 70s a ua hoʻokumu ʻia kahi maʻi ahulau ma laila i ka ʻuala a me nā tōmato ma hope o 32 mau makahiki me ka maʻi ʻole. Ma muli o kona kūpikipiki kiʻekiʻe, ua kāhea ʻo ia i ka clone US-1 a ua hoʻomaka ʻo ia e hoʻomalu i ke kapakai komohana o ʻAmelika Hui Pū ʻIa (Goodwin et al., 1995a).
Ua ʻike ʻia nā genotypes US-7 a me US-8 i ʻAmelika Hui Pū ʻIa i ka makahiki 1992, a i kēia manawa ua hāʻawi ʻia ma 1994 i ka United States a me Kanada. I loko o hoʻokahi wā kau, hiki i ka clone US-8 ke aneane hoʻopau loa i ka clone US-1 i loko o nā pālala potato i pā mua ʻia me nā klone ʻelua i kahi kūlike like (Miller a me Johnson, 2000).
Ua ʻike ʻia nā klone BC-1 a hiki i BC-4 ma British Columbia ma kahi helu liʻiliʻi o nā kaʻawale mai Goodwin et al., 1995b). Ua hoʻolaha ākea ʻia ʻo Clone US-11 i United States a ua kāpae aku ʻo US-1 ma Taiwan. ʻO nā clones JP-1 a me EC-1, me ka clone US-1, he mea maʻamau ia i Iapana a me ʻEkuakua, (Koh et al., 1994; Forbes et al., 1997).
ʻO SIB-1 kahi klone i lanakila ma Rūsia ma luna o kahi panalāʻau nui mai ka moku ʻo Moscow a Sakhalin. Ma ka moku ʻo Moscow, ua ʻike ʻia ia ma ka makahiki 1993, a ʻo kekahi mau heluna nui o ke kula he ʻano nui o nā laina o kēia laina clonal, kūpaʻa loa i ka metalaxyl. Ma hope o 1993, ua emi ka nui o kēia klone. Ma waho o nā Ural i ka makahiki 1997-1998, ua loaʻa ʻo SIB-1 i nā wahi āpau, koe wale nō ka Teritori Khabarovsk (hoʻolaha ʻia ka clone SIB-2 ma laila). ʻO ka hoʻokaʻawale ākea o nā pōhaku me nā ʻano moena like ʻole e kāpae i ke kaʻina moekolohe ma Siberia a me ka Hikina hikina. Ma ka moku ʻo Moscow, ʻokoʻa i Siberia, hōʻike ʻia ka heluna kanaka e nā klone he nui; ʻaneʻane loaʻa i kēlā me kēia kaʻawale kahi genotype multilocus kūikawā (Elansky et al., 2001, 2015). ʻAʻole hiki ke wehewehe wale ʻia kēia ʻano ʻokoʻa e ka lawe wale ʻana mai i nā kānana o ka fungus mai nā wahi like ʻole o ka honua me nā mea kanu i lawe ʻia mai. Ma muli o ke ʻano o ka moʻa ʻana ʻelua i ka heluna kanaka, hiki nō paha i kona ʻano like ʻole ma muli o ka recombination. No laila, ma British Columbia, ke kuhi ʻia ʻana o nā genotypes BC-2, BC-3, a me BC-4 i manaʻo ʻia ma muli o ka hybridization o nā clone BC-1 a me US-6 (Goodwin et al., 1995b). Hiki paha ke loaʻa nā ʻano hybrid i nā heluna o Moscow. ʻO kahi laʻana, hiki i nā kaha MO-4, MO-8 a me MO-11 heterozygous no ka wahi PEP ke lilo i mau hybrids ma waena o nā kaha MO-12, MO-21, MO-22, me ka loaʻa ʻana o kahi ʻano male A2 a me ka homozygous no hoʻokahi alele o ka PEP locus a me ke kanana. ʻO MO-8, ke ʻano o ka moʻo ʻana o ka A1 a me ka homozygous no nā alele ʻē aʻe o ka locus. A inā pēlā, a i nā lehulehu o P. infestans i kēia manawa aia ka hoʻonui ʻana i ke kuleana o ka hana moekolohe, a laila e hoʻemi ʻia ka waiwai o ka ʻike o ke aniani o nā clone multilocus (Elansky et al., 2001, 2015).
Hoʻololi i nā laina clonal
A hiki i nā 90 o ke kenekulia 20, ua laha ka laina clonal US-1 i ka honua. ʻO ka hapa nui o nā māla a me nā heluna āpana kahi o nā kaupaona me ka genotype US-1. Eia nō naʻe, ua nānā ʻia nā ʻokoʻa ma waena o nā kaʻawale, a ʻo ia paha ka hopena o ke kaʻina hoʻololi. Ua loaʻa nā hoʻololi i nā DNA nukelea a me mitochondrial a hoʻopili ʻia, i waena o nā mea ʻē aʻe, ke kiʻekiʻe o ke kūpaʻa ʻana i nā lāʻau phenylamide a me ka helu o nā genulence virulence. Hōʻike ʻia nā laina ʻokoʻa mai nā genotypes kumu e nā hoʻololi ʻana e nā helu hou aʻe ma hope o ke kiko e pili ana i ka inoa o ka genotype kumu (e laʻa me ka laina mutant US-1.1 o ka laina clonal US-1). Loaʻa i nā laina DNA manamana lima US-1.5 a me US-1.6 i nā laina kōkua o nā nui like ʻole (Goodwin et al., 1995a, 1995b); ʻokoʻa ka laina clonal US-6.3 mai US-6 e hoʻokahi laina kōkua (Goodwin, 1997, Papa 7).
I ke aʻo ʻana o DNA mitochondrial, ua ʻike ʻia ma ka laina clonal US-1 wale nō ka type 1b o DNA mitochondrial i loaʻa (Carter et al., 1990). Eia nō naʻe, i ke aʻo ʻana i nā ʻōiwi o kēia moʻokūʻauhau clonal mai Peru a me Pilipine, ua ʻike ʻia nā mea kaʻawale nona nā ʻano mitochondrial DNA ʻokoʻa i ka 1b me ka loaʻa ʻana o nā hoʻokomo a me nā kāpae ʻana (Goodwin, 1991, Koh et al., 1994).
Pakuhi 6. Multilocus genotypes o kekahi P. infestans clonal laina
Inoa | ʻAno hoʻoipoipo | ʻO Isozymes | Nā manamana lima DNA | ʻAno ʻo MtDNA | |
GPI | PEP | ||||
US-1 | A1 | 86/100 | 92/100 | 1.0111010110011E + 24 | Ib |
US-2 | A1 | 86/100 | 92/100 | 1.0111010010011E + 24 | - |
US-3 | A1 | 86/100 | 92/100 | 1.0111000000011E + 24 | - |
US-4 | A1 | 100/100 | 92/92 | 1.0111010010011E + 24 | - |
US-5 | A1 | 100/100 | 92/100 | 1.0111010010011E + 24 | - |
US-6 | A1 | 100/100 | 92/100 | 1.0111110010011E + 24 | IIb |
US-7 | A2 | 100/111 | 100/100 | 1.0011000010011E + 24 | Ia |
US-8 | A2 | 100/111/122 | 100/100 | 1.0011000010011E + 24 | Ia |
US-9 | A1 | 100/100 | 83/100 | * | - |
US-10 | A2 | 111/122 | 100/100 | - | - |
US-11 | A1 | 100/111 | 92/100 | 1.0101110010011E + 24 | IIb |
US-12 | A1 | 100/111 | 92/100 | 1.0001000010011E + 24 | - |
US-14 | A2 | 100/122 | 100/100 | 1.0000000000011E + 24 | - |
US-15 | A2 | 100/100 | 92/100 | 1.0001000010011E + 24 | Ia |
US-16 | A1 | 100/111 | 100/100 | 1.0001100010011E + 24 | - |
US-17 | A1 | 100/122 | 100/100 | 1.0100010000011E + 24 | - |
US-18 | A2 | 100/100 | 92/100 | 1.0001000010011E + 24 | Ia |
US-19 | A2 | 100/100 | 92/100 | 1.0101010000011E + 24 | Ia |
EC-1 | A1 | 90/100 | 96/100 | 1.1111010010011E + 24 | IIa |
SIB-1 | A1 | 100/100 | 100/100 | 1.0001000110011E + 24 | IIa |
SIB-2 | A2 | 100/100 | 100/100 | 1.0001000010011E + 24 | IIa |
SIB-3 | A1 | 100/100 | 100/100 | 1.1001010100011E + 24 | IIa |
MO-1 | A2 | 100/100 | 100/100 | 1.0001000110011E + 24 | IIa |
MO-2 | A2 | 100/100 | 100/100 | 1.0001000010011E + 24 | Ia |
MO-3 | A1 | 100/100 | 100/100 | 1.0101000010011E + 24 | IIa |
MO-4 | A1 | 100/100 | 92/100 | 1.0101110110011E + 24 | IIa |
MO-5 | A1 | 100/100 | 100/100 | 1.0001010010011E + 24 | IIa |
MO-6 | A1 | 100/100 | 100/100 | 1.0101010010011E + 24 | Ia |
MO-7 | A1 | 100/100 | 92/100 | 1.0001000110011E + 24 | IIa |
MO-8 | A1 | 100/100 | 92/92 | 1.0101100010011E + 24 | IIa |
MO-9 | A1 | 100/100 | 92/100 | 1.0001000010011E + 24 | IIa |
MO-10 | A1 | 100/100 | 100/100 | 1.0101100000011E + 24 | Ia |
MO-11 | A1 | 100/100 | 92/100 | 1.0101010010011E + 24 | Ia |
MO-12 | A2 | 100/100 | 100/100 | 1.0101010010011E + 24 | Ia |
MO-13 | A1 | 100/100 | 100/100 | 1.0101010000011E + 24 | Ia |
MO-14 | A1 | 100/100 | 100/100 | 1.01010010011E + 22 | Ia |
MO-15 | A1 | 100/100 | 100/100 | 1.101110010011E + 23 | Ia |
MO-16 | A1 | 100/100 | 100/100 | 1.0001000000011E + 24 | IIa |
MO-17 | A1 | 86/100 | 100/100 | 1.0101010110011E + 24 | Ib |
MO-18 | A1 | 100/100 | 100/100 | 1.0101110010011E + 24 | IIa |
MO-19 | A1 | 100/100 | 100/100 | 1.0101010000011E + 24 | IIa |
MO-20 | A2 | 100/100 | 100/100 | 1.0101010000011E + 24 | IIa |
MO-21 | A2 | 100/100 | 100/100 | 1.0101010000011E + 24 | IIa |
Kaha: * - ʻaʻohe ʻikepili.
Pakuhi 7. Nā genotypes Multilocus a me kā lākou mau laina mutant
Inoa | ʻAno hoʻoipoipo | | Nā Mīkini DNA (RG57) | Nā memo | |
GPI | PEP-1 | ||||
US-1 | A1 | 86/100 | 92/100 | 1011101011001101000110011 | ʻO genotype kumu 1 |
US-1.1 | A1 | 86/100 | 100/100 | 1011101011001101000110011 | Hoʻololi i PEP |
US-1.2 | A1 | 86/100 | 92/100 | 1011101010001101000110011 | Hoʻololi i RG57 |
US-1.3 | A1 | 86/100 | 92/100 | 1011101001001101000110011 | Hoʻololi i RG57 |
US-1.4 | A1 | 86/100 | 100/100 | 1011101010001101000110011 | Hoʻololi i RG57 a me PEP |
US-1.5 | A1 | 86/100 | 92/100 | 1011101011001101010110011 | Hoʻololi i RG57 |
US-6 | A1 | 100/100 | 92/100 | 1011111001001100010110011 | ʻO genotype kumu 2 |
US-6.1 | A1 | 100/100 | 92 /92 | 1011111001001100010110011 | Hoʻololi i PEP |
US-6.2 | A1 | 100/100 | 92/100 | 1011101001001100010110011 | Hoʻololi i RG57 |
US-6.3 | A1 | 100/100 | 92/100 | 1011111001011100010110011 | Hoʻololi i RG57 |
US-6.4 | A1 | 100/100 | 100/100 | 1011011001001100010110011 | Hoʻololi i RG57 a me PEP |
US-6.5 | A1 | 100/100 | 92/100 | 1011111001001100010010011 | Hoʻololi i RG57 |
BR-1-KA | A2 | 100/100 | 100/100 | 1011101000001100001111011 | ʻO genotype kumu 3 |
BR-1.1-KA | A2 | 100/100 | 100/100 | 1010101000001100001110011 | Hoʻololi i RG57 |
Aia kekahi mau hoʻololi i ka hōʻike o isozymes. Ma ke ʻano he kumu, kumu ʻia lākou e ka haki ʻana o kahi meaola heterozygous i kinohi no kēia enzyme i nā mea homozygous. I ka makahiki 1993, ma nā hua kōmato, ʻike mākou i kahi kānana me nā ʻano o ka US-1: RG57 fingerprinting, mitochondrial DNA type, a me 86/100 genotype no glucose-6-phosphatizomerase, akā he homozygous (100/100) no ka wahi peptidase mua ma kahi o he 92/100 heterozygote ʻano o kēia laina clonal. Ua kapa mākou i ka genotype o kēia kānana MO-17 (Papa 6). ʻOkoʻa nā laina mutant US-1.1 a me US-1.4 mai US-1 e nā hoʻololi ʻana ma ka locus peptidase mua (Papa 7).
ʻO nā hoʻololi e alakaʻi i nā loli i ka helu o nā genul virulence no nā ʻano ʻuala a me nā ʻano kōmato he mea maʻamau. Ua kaulana lākou i waena o nā kaʻawale o ka laina clonal US-1 i nā lehulehu mai Netherlands (Drenth et al., 1994), Peru (Goodwin et al., 1995a), Polani (Sujkowski et al., 1991), ʻAkau ʻAkau o ʻAmelika (Goodwin et al., ., 1995b). ʻO nā ʻokoʻa i ka helu o nā genes virulence ʻuala i ʻike ʻia i waena o nā kaʻawale o nā laina clonal US-7 a me US-8 ma Kanada a me ʻAmelika Hui Pū ʻIa (Goodwin et al., 1995a), i waena o nā kaʻawale o ka laina SIB-1 ma ka ʻĀkia ʻāpana o Rūsia (Elansky et al, 2001 ).
Hoʻokaʻawale ʻia me nā ʻokoʻa ikaika i nā pae o ke kūʻē ʻana i nā lāʻau lapaʻau phenylamide i ʻike ʻia i loko o ka lehulehu monoclonal, a ʻo nā mea āpau e pili ana i ka laina clonal Sib-1 (Elansky et al, 2001, Papa 1). ʻAneʻane kokoke i nā luhi a pau o ka laina clonal US-1 e makaʻala loa i ka metalaxyl; eia nō naʻe, ua hoʻokaʻawale ʻia nā kaʻawale kūʻē kūʻē ʻole o kēia laina i ka ʻĀina Pilipino (Koh et al., 1994) a ma ʻIlani (Goodwin et al., 1996).
Nā lehulehu o P. infestans
ʻAmelika Hui Pū ʻIa (Mexico)
ʻOkoʻa ka P. infestans heluna kanaka ma Mekiko i nā heluna honua ʻē aʻe, ʻo ia ke kumu o kāna kūlana mōʻaukala. ʻO nā noiʻi he nui o kēia heluna kanaka a pili pū kekahi P. infestans species of the clade Phytophthora, a me nā kūloko o ka genus Solanum, i alakaʻi ʻia ai i ka hopena o ka ulu ʻana o ka pathogen ma ka waena konu o Mexico i hui pū ʻia me ka ulu ʻana o nā mea kanu hoʻokipa a pili pū me ka hoʻohui wahine hou (Grünwald, Flier , 2005). Hōʻike ʻia nā ʻano moena ʻelua i ka heluna kanaka, a i nā kaulike like, a me ke kū ʻana o ka oospores i ka lepo, ma nā mea kanu a me nā paipala o kaʻuala a me nā mea ulu e ulu ana i nā holoholona hihiu e hōʻoia ana i ka loaʻa ʻana o kahi hana moekolohe i ka lehulehu (Fernández-Pavía et al., 2002). ʻO nā loiloi hou o ke awāwa ʻo Toluca a me nā wahi e hoʻopuni ana (ke kikowaena manaʻo o ke kumu o ka pathogen) i hōʻoia i ka nui o nā ʻano kūpuna o ka poʻe kūloko o P. infestans (134 multilocus genotypes i kahi laʻana o 176 mau laʻana) a me ke kū ʻana o kekahi mau subpopulation ʻokoʻa i ka ʻāina (Wang et al., 2017). ʻO nā kumu e hāʻawi nei i kēia ʻokoʻa ka mahele mokuna o nā subpopulation ma nā mauna kiʻekiʻe o ke kikowaena o Mekiko, nā ʻokoʻa o nā ʻano mahi a me nā ʻano potato i hoʻohana ʻia i nā awāwa a me nā kuahiwi, a me ke kū ʻana o nā lāhui Solanum tuberous hihiu i hiki ke hana ma ke ʻano he poʻe ʻē aʻe (Fry et al ., 2009).
Eia nō naʻe, pono e hoʻomaopopo ʻia he lehulehu wale nā heluna o P. infestans ma ka ʻākau o Mekiko a ʻano like me ka heluna o ʻAmelika Hui Pū ʻIa, e hōʻike ana paha kēia nā genotypes hou (Fry et al., 2009).
ʻAmelika ʻAmelika
ʻO ka poʻe o ʻAmelika Hui Pū ʻIa o P. infestans i kahi manawa maʻalahi maʻalahi a ua hoʻokumu ʻia ko lākou ʻano clonal ma mua o ka hoʻohana ʻana o ka microsatelit analysis. A hiki i ka 1987, ua lanakila ka laina clonal US-1 i United States a me Kanada (Goodwin et al., 1995). I ka waenakonu o nā makahiki 70, i ka wā i puka mai ai nā fungicides i hoʻokumu ʻia me ka metalaxyl, ua hoʻomaka ʻia kēia clone e nā mea ʻē aʻe, nā genotypes kūʻē aʻe i neʻe mai Mexico mai (Goodwin et al., 1998). Ma ka hopena o nā 90s. ua hoʻololi loa ka genotype US-8 i ka genotype US-1 i United States a lilo i laina clonal kiʻekiʻe ma kaʻuala (Fry et al., 2009; Fry et al., 2015). ʻOkoʻa ke kūlana me nā ʻōmato, ka mea i paʻa mau i kekahi mau laina clonal, a ua loli ko lākou ʻano mai kēlā makahiki a kēia makahiki (Fry et al., 2009).
I ka makahiki 2009, ua laha mai kahi maʻi ahulau nui o ka hopena hope loa i nā tumato maʻAmelika Hui Pū ʻIa. ʻO kahi hiʻohiʻona o kēia maʻi ahulau ka ʻaneʻane like like ʻana o ka hoʻomaka ʻana ma nā wahi he nui ma ka ʻākau hikina o ʻAmelika Hui Pū ʻIa, a ua lilo ia i mea pili i ke kūʻai nui ʻia ʻana o nā hua kōmato maʻi i nā kikowaena māla nui (Fry et al., 2013). Nui loa nā pohō o nā mea kanu. Ua hōʻike ʻia ka anamanaʻo microsatellite o nā laʻana i hoʻopilikia ʻia e pili ana ka maʻi pandemi i ka laina clonal US-22 A2-type mating. I ka makahiki 2009, ʻo ka mahele o kēia genotype i ka heluna kanaka ʻAmelika o P. infestans ua hōʻea i 80% (Fry et al., 2013). I nā makahiki aʻe, ua piʻi mau ka hapa o nā genotypes huhū US-23 (ka hapanui ma nā ʻōmato) a me US-24 (ma kaʻuala) i ka heluna kanaka, akā naʻe, ma hope o 2011, ua emi ka nui o ka loaʻa o US-24, a i kēia lā, ma kahi o 90% o ka heluna pathogen i Ua hōʻike ʻia ʻo ʻAmelika Hui Pū ʻIa e ka genotype US-23 (Fry et al., 2015).
Ma Kanada, e like me ʻAmelika Hui Pū ʻIa, i ka pau ʻana o nā makahiki 90. ʻo ka genotype nui loa US-1 i hakahaka ʻia e US-8, ke kūlana koʻikoʻi i waiho mau a hiki i ka 2008. Ma Kanada, aia nā maʻi wīwī weliweli e pili pū ana me ke kūʻai aku ʻana i nā lūlū kōmato i kūpale ʻia, akā na nā genotypes US-2009 a me US-2010 (Kalischuk et al., 23) i hoʻokumu. Kupaianaha ka hoʻokaʻawale kūwaho o kēia mau genotypes: ua lanakila ʻo US-8 i nā panalāʻau komohana o Kanada (2012%), ʻoiai ʻo US-23 i hoʻomalu i nā panalāʻau hikina (68%). I nā makahiki aʻe, ua pāhola ʻo US-8 i nā ʻāina hikina; eia nō naʻe, i ka nui, ua hoʻemi iki ʻia kāna mahele o ka heluna kanaka e pili ana i ke ʻano o nā genotypes US-83 a me US-23 ma ka ʻāina (Peters et al., 22). I kēia lā, mālama ʻo US-24 i kahi kūlana koʻikoʻi ma waena o Kanada; Aia ʻo US-2014 ma British Columbia, ʻoiai ʻo US-23 a me US-8 aia ma Ontario (Peters, 23).
No laila, ʻo nā heluna o ʻAmelika ʻĀkau o P. infestans ka nui o nā laina clonal. I loko o nā makahiki he 40 i hala aku nei, ua hōʻea ka helu o nā genotypes clonal i ʻike ʻia i 24. ʻOiai ke ʻano o nā ʻano o nā moena ʻelua i ka heluna kanaka, ʻo ka likelike o ke ʻano o nā genotypes hou ma muli o ka moekolohe e hoʻomau i ka haʻahaʻa. Eia nō naʻe, i nā makahiki he 20 i hala aku nei, ua hoʻopaʻa ʻia nā hihia he nui o ka heluna o nā ephemeral recombinant (Gavino et al., 2000; Danies et al., 2014; Peters et al., 2014), a ma kekahi hihia, ʻo ka hopena o ke keʻa ʻana o ka genotype US-11 , i hoʻopaʻa ʻia ma ʻAkau ʻAmelika no nā makahiki he nui (Gavino et al., 2000). A hiki i ka 2009, ua pili nā hoʻololi ʻana i ke ʻano o nā heluna kanaka me ka puka ʻana o nā genotypes hou hou aʻe me kā lākou neʻe ʻana a me ka neʻe ʻana o nā mea i noho mua ma mua. He aha ka hana i 2009-2010 Ma USA a me Kanada, ua hōʻike nā epiphytotics no ka manawa mua i ke au o ka globalisation, hiki ke hoʻopili ʻia nā maʻi o ka maʻi me ka hoʻolaha ākea o nā genotypes hou ke kūʻai aku nei i nā mea kanu maʻi.
Amerika Hema
A hiki i kēia manawa, ʻo nā noiʻi o nā heluna o ʻAmelika Hema o P. infestans ʻaʻole i maʻamau a nui paha. ʻIke ʻia ka maʻalahi o ke ʻano o kēia mau heluna a komo pū me nā lālani clonal 1-5 no kēlā me kēia ʻāina (Forbes et al., 1998). No laila, ma ka 1998, nā genotypes US-1 (Brazil, Chile) BR-1 (Brazil, Bolivia, Uruguay, Paraguay), EC-1 (Ecuador, Colombia, Peru a me Venezuela), AR-1, AR i loaʻa ma kaʻuala -2, AR-3, AR-4 a me AR-5 (Argentina), PE-3 a me PE-7 (hema o Peru). Aia ka maʻa male A2 ma Brazil, Bolivia a me ʻAtenekia a ʻaʻole i loaʻa ma ʻō aku o ka palena ʻo Bolivian-Peruvian ma ka wahi o Lake Titicaca, ma hope o ka EC-1 A1 genotype i noho aliʻi ma Andes. Ma nā ʻōmato, ua hoʻomau ʻo US-1 i ka genotype koʻikoʻi ma ʻAmelika Hema.
Ua hoʻomau ʻia ke kūlana a i ʻole emi iki paha i nā makahiki 2000. ʻO kahi kiko nui ka ʻike ʻia ma ka ʻAkau ʻAkau ma nā ʻohana ʻuala ulu ulu (S. brevifolium a me S. tetrapetalum) o kahi laina clonal EC-2 hou o ka ʻano A2 (Oliva et al., 2010). Ua hōʻike ʻia nā noiʻi Phylogenetic ʻaʻole kūlike loa kēia laina i ka P. infestans, ʻoiai e pili pono iā ia, e pili ana i ka mea i noi ʻia e noʻonoʻo iā ia, a me kahi laina ʻē aʻe, EC-3, hoʻokaʻawale ʻia mai ka lāʻau kōmato S. betaceum e ulu nei i ka Andes, kahi ʻano hou i kapa ʻia ʻo P. andina; Eia nō naʻe, ʻaʻole maopopo ke kūlana o kēia ʻano (kahi ʻano kūʻokoʻa a i ʻole kahi hybrid o P. infestans me kekahi laina ʻike ʻole ʻia) (Delgado et al., 2013).
I kēia manawa, ʻo nā poʻe a pau o ʻAmelika Hema o P. infestans he clonal. I loko nō o ke ʻano o nā moena ʻelua, ʻaʻole i ʻike ʻia nā heluna recombinant. Ma nā ʻōmato, aia ka genotype US-1 i ʻōbiquitous, kahi i haʻalele ʻia mai nā ʻuala e nā ʻano kūloko, ʻaʻole ʻike ʻia ke kumu kikoʻī. I Brazil, Bolivia a me Uruguay aia ka genotype BR-1; ma Peru, me US-1 a me EC-1, aia kekahi mau genotypes kūloko ʻē aʻe. I ka Andes, ua hoʻomau ʻia ke kūlana poʻokela e ka laina clonal EC-1, ka pilina me ia me P. andina i ʻike hou ʻia i ʻike ʻole ʻia. ʻO kahi wale nō "paʻa ʻole" kahi no ka manawa 2003-2013. aia nā hoʻololi nui i ka heluna kanaka, lilo iā Chile (Acuña et al., 2012), kahi ma 2004-2005. ua lilo ka heluna pathogen i ke kūʻē ʻana i ka metalaxyl a me kahi haplotype DNA mitochondrial hou (Ia ma kahi o ka Ib i mua. 2006 a 2011 i ka heluna kanaka, ua loaʻa ka genotype 21 (e like me SSR), a ʻo ka mahele i hōʻea i 90%, a ma hope iho ua hala ka palama i ka genotype 20, ʻo ke alapine o ka hanana i mālama ʻia i loko o nā makahiki ʻelua e hiki mai ana ma kahi o 67% (Acuña, 2015).
ʻO Europe
I ka moʻolelo o ʻEulopa, aia ma ka liʻiliʻi he ʻelua mau nalu o ka neʻe ʻana o P. infestans mai ʻAmelika Hema: i ka kenekulia 1. (HERB-1) a me ke kenekulia 70 (US-1). ʻO ka hāʻawi ākea o nā metalaxyl-loaʻa fungicides i nā makahiki XNUMX. alakaʻi i ka neʻe ʻana o ka genotype nui US-XNUMX a me kona pani me nā genotypes hou. A ʻo kahi hopena, ma ka hapa nui o nā ʻāina o ʻEulopa Komohana, ua hōʻike nui ʻia nā lehulehu o nā pathogen e kekahi mau laina clonal.
ʻO ka hoʻohana ʻana o ka microsatelit analysis no ke kālailai ʻana i nā heluna pathogen i hiki ke ʻike i nā loli koʻikoʻi i hana ʻia ma Western Europe i ka makahiki 2005-2008. I ka 2005, ua ʻike ʻia kahi laina clonal hou ma UK, i kapa ʻia ʻo 13_A2 (a i ʻole "Blue 13") a hoʻohālikelike ʻia e ka A2 mating type , ka ikaika nui a me ke kūpaʻa ʻana i nā phenylamides (Shaw et al., 2007). Ua loaʻa ka genotype like i loko o nā laʻana i hōʻiliʻili ʻia ma 2004 ma Netherlands a me ka ʻākau o Palani, e ʻōlelo ana ua neʻe i UK mai ka ʻĀinapuniʻole ʻEulopa, malia paha me nā hua uala (Cooke et al., 2007). ʻO ke aʻo ʻana i ka genome o nā lunamakaʻāinana o kēia laina clonal i hōʻike i kahi kiʻekiʻe o polymorphism o kāna kaʻina (e 2016, ka helu o kāna mau ʻano subclonal i hōʻea i 340) a me kahi ʻano nui o ka loli i ka pae o ka hōʻike o ka ʻaoʻao, incl. nā gen effector i ka wā o ka maʻi maʻi (Cooke et al., 2012; Cooke, 2017). ʻO kēia mau hiʻohiʻona, a me ka hoʻonui ʻia o ka lōʻihi o ka pae biotrophic, hiki ke hoʻonui i ka hakakā o 13_A2 a me kona hiki ke hoʻopilikia i nā ʻano potato kūʻē i ka hopena o ke ahi.
I nā makahiki e hiki mai ana, hoʻolaha koke ka genotype ma nā ʻāina o Northwestern Europe (Great Britain, ʻIlani, Palani, Pelekiuma, Netherlands, Kelemānia) me ka neʻe like ʻana o nā genotypes mana nui ma mua 1_A1, 2_A1, 8_A1 (Montarry et al., 2010; Gisi et al. , 2011; Van den Bosch et al., 2011; Cooke, 2015; Cooke, 2017). Wahi a ka pūnaewele www.euroblight.net, ka mahele o 13_A2 i nā heluna o kēia mau ʻāina i hōʻea i 60-80% a ʻoi aku; ua hoʻopaʻa ʻia ke alo o kēia genotype i kekahi mau ʻāina o ʻEulopa Hikina a me ʻEulopa Hema. Eia nō naʻe, i ka makahiki 2009-2012. Ua lilo nā kūlana koʻikoʻi o 13_A2 ma Great Britain a me Palani, e hāʻawi ana i ka laina 6_A1 (i Ireland - 8_A1), a ma Netherlands a me Belgian ua hoʻololi ʻia ia e nā genotypes 1_A1, 6_A1, a me 33_A2 (Cooke et al., 2012; Cooke, 2017; Stellingwerf, 2017).
I kēia lā, ma kahi o 70% o ka heluna ʻEulopa o P. infestans he monoclonal. Wahi a ka pūnaewele www.euroblight.net, nā genotypes nui i nā ʻāina o North-Western Europe (UK, Palani,
Netherlands, Pelekiuma) koe, ma kahi o nā kaulike kaulike, 13_A2 a me 6_A1, a ʻo ka hope ʻaʻole kokoke e kū ma waho o ka ʻāpana i kuhikuhi ʻia (koe wale nō ʻo ʻIlani), akā aia ma kahi o 58 mau subclones (Cooke, 2017). Aia nā ʻano 13_A2 i nā helu i ʻike nui ʻia ma Kelemania, a ʻike pinepine ʻia hoʻi ma nā ʻāina o Central a me ʻEulopa ʻEulopa. ʻO Genotype 1_A1 he hapa nui o nā heluna lehulehu o Pelekiuma a me kahi hapa ʻo Netherlands a me Palani. Ua hoʻokūpaʻa ʻo Genotype 8_A1 i ka heluna o ʻEulopa ma ka pae o 3-6%, koe wale nō ʻo ʻIlani, kahi e paʻa ai kona kūlana alakaʻi a mahele ʻia i ʻelua subclones (Stellingwerf, 2017). ʻO ka mea hope loa, i ka 2016, ua hoʻonui ʻia ka hoʻonui pinepine ʻana o nā genotypes hou 36_A2 a me 37_A2, i hoʻopaʻa mua ʻia ma 2013-2014; i kēia lā, loaʻa kēia mau genotypes ma Netherlands a me Belgian a ma kekahi hapa ma Palani a me Kelemania, a ma ka ʻaoʻao hema o Great Britain (Cooke, 2017). Ma kahi o 20-30% o ka heluna nui o ʻEulopa e hōʻike ʻia e nā genotypes kūikawā i kēlā me kēia makahiki.
ʻAʻole like me ʻEulopa Komohana, i ka manawa i puka mai ai ka genotype 13_A2, ua hōʻike ʻia nā lehulehu o ka ʻĀkau ʻEulopa (Kuekene, Nolewai, Denmark, Pinilana) ʻaʻole nā laina clonal, akā na ka nui o nā genotypes kūikawā (Brurberg et al.,
2011). I loko o ka manawa o ka hoʻolaha ʻana o 13_A2 ma Western Europe, ʻaʻole i ʻike ʻia ke kū ʻana o kēia genotype ma Scandinavia a hiki i ka makahiki 2011, i ka manawa i ʻike mua ʻia ai ia ma North Jutland (Denmark), kahi e ulu nui ai nā ʻano ʻuala ʻoihana me ka hoʻohana nui o ka metalaxyl-i loko. fungicides (Nielsen et al., 2014). Wahi a www.euroblight.net, ua ʻike ʻia kekahi genotype 13_A2 i kekahi mau laʻana mai Norewai a me Denmark i ka 2014 a i kekahi mau mea Norewai i 2016; i ka hoʻohui, i ka 2013 ma Pinilana ke ʻano o ka genotype 6_A1 i kahi liʻiliʻi. ʻO ke kumu nui o ka holo ʻole o 13_A2 a me nā laina clonal ʻē aʻe i ka naʻi ʻana o Scandinavia i manaʻo ʻia ʻo ia ka ʻokoʻa kūwaho o kēia wahi mai nā ʻāina o ʻEulopa Komohana.
Hoʻohui i ka ʻoiaʻiʻo o nā kauwela anuanu a me nā hoʻoilo anuanu e paipai i ke ola ʻana o ka mycelium vegetative nui loa e like me nā oospores (Sjöholm et al., 2013), ka hau ʻana i ka lepo i ka hoʻoilo (ka mea maʻamau ʻole i nā ʻāina mehana o ʻEulopa Komohana) kōkua i ka hoʻopili like ʻana o ka ulu ʻana o ka oospores a me ke kanu ʻana. kaʻuala, kahi e hoʻonui ai i kā lākou kuleana ma ke ʻano he kumu o ka maʻi maʻi mua (Brurberg et al., 2011). Hoʻomaopopo ʻia ia, i ke kūkulu ʻākau, ka ulu ʻana o ka maʻi mai nā oospores ma mua o ka hoʻomohala ʻia ʻana o ka maʻi tuberous, ka mea e pale ai i ka noho aliʻi ʻoi loa o ka hakakā, akā ua hoʻomohala ʻia nā laina clonal (Yuen, 2012). ʻO ke ʻano o nā heluna kanaka i aʻo nui ʻia o P. infestans ma ka Hikina ʻEulopa (Polani, nā mokuʻāina ʻo Baltic) he like loa me kēlā ma Scandinavia.
Aia kekahi o nā ʻano male ʻelua ma aneʻi, a ʻo ka hapa nui o nā genotypes i hoʻoholo ʻia e ka anamanaʻo SSR kū hoʻokahi (Chmielarz et al., 2014; Runno-Paurson et al., 2016). E like me ka ʻĀkau o ʻEulopa, ka hoʻokaʻawale ʻana i nā laina clonal (ke ʻano nui o ka genotype 13_A2) pili maoli ʻole i ka lehulehu kūloko o ka pathogen, ka mea i paʻa i kahi pae kiʻekiʻe o ka like ʻole me ka ʻole o nā laina mana i haʻi ʻia.
ʻIke ʻia ke alo o 13_A2 i nā māla me nā ʻano ʻuala kalepa. Ma Rūsia, ke ulu nei ke kūlana i kahi ala like. ʻImi ʻia ka microsatelit analysis o P. infestans isolates i hōʻiliʻili ʻia ma 2008-2011 i 10 mau ʻāpana ʻokoʻa o ka ʻāpana ʻEulopa o Lūkia, ua hōʻike i kahi kiʻekiʻe o ka genotypic ʻokoʻa a me ka nele piha o nā kūlike ʻole me nā laina clonal ʻEulopa (Statsyuk et al., 2014). I kekahi mau makahiki ma hope mai, ua aʻo ʻia kahi laʻana o P. infestans i hōʻiliʻili ʻia ma ka moku ʻo Leningrad ma 2013-2014 i nā ʻokoʻa nui ma waena o lākou a me nā genotypes mai kēia ʻāpana i ʻike ʻia i loko o ka noiʻi i hala. I nā noiʻi ʻelua, ʻaʻole i loaʻa nā genotypes ʻEulopa Komohana (Beketova et al., 2014; Kuznetsova et al., 2016).
ʻO ke ʻano ʻē aʻe o nā lāhui ʻEulopa Hikina o P. infestans a me ka ʻole o nā laina clonal i loko o lākou no nā kumu he nui. ʻO ka mea mua, e like me ka ʻĀkau o ʻEulopa, nā kūlana anilā o nā ʻāina i manaʻo ʻia e hāʻawi i ka hoʻokumu ʻana i nā oospores ma ke ʻano he kumu kumu o ka maʻi (Ulanova et al., 2010; Chmielarz et al., 2014). ʻO ka lua, kahi mahele nui o kaʻuala i hana ʻia i kēia mau ʻāina i ulu ma nā pā liʻiliʻi liʻiliʻi, e hoʻopuni pinepine ʻia e nā ululāʻau a i ʻole nā mea keakea i ka neʻe manuahi ʻana o nā mea infectious (Chmielarz et al., 2014). Ma ke ʻano he kānāwai, ʻaʻole mālama pono ʻia kaʻuala me ia mau ʻano me nā kemika, a ʻo ke koho ʻana o nā ʻano like ʻole e pili ana i ka hopena o ke kūpaʻa ʻana o ka blight, ʻo ia hoʻi ʻaʻohe kaomi koho no ka huhū a me ke kūpaʻa ʻana i ka metalaxyl, kahi e hoʻonele ai i nā genotypes kūpaʻa, e like me 13_A2, o nā mea maikaʻi ma mua o nā genotypes ʻē aʻe (Chmielarz et al., 2014). ʻO ka mea hope loa, ma muli o ka liʻiliʻi o nā kihi ʻāina, ʻaʻole hoʻomaʻamaʻa ka mea nona nā mea kanu i ka hoʻohuli ʻana i nā hua, e ulu nei i kaʻuala ma kahi like no nā makahiki, kahi e kōkua ai i ka hōʻiliʻili ʻana o kahi inoculum ʻano ʻokoʻa (Runno-Paurson et al., 2016; Elansky, 2015; Elansky et al. ., 2015).
ʻO Asia
A hiki i kēia manawa, ua ʻano maʻalahi ka ʻike ʻia ʻana o ka P. P. infestans. Ua ʻike ʻia ka mea e hōʻike nui ʻia e nā laina clonal, a he liʻiliʻi loa ka hopena o ka recombination moekolohe i ka puka ʻana o nā genotypes hou. No laila, no ka laʻana, ma 1997-1998. Ma ka ʻĀkia ʻāpana o Lūkia (Siberia a me ka Hikina hikina), ua hōʻike ʻia ka heluna pathogen e ʻekolu mau genotypes wale nō me ka predominance o ka genotype SIB-1 (Elansky et al., 2001). Ua hōʻike ʻia ke kū ʻana o nā laina pathogen clonal i nā ʻāina e like me Kina, Iapana, Korea, Pilipino, a me Taiwan (Koh et al., 1994; Chen et al., 2009). Ua lanakila ka laina clonal US-1 ma luna o kahi ʻāina nui o ʻAsia i ka hopena o 90 - kakahiaka 2000. ʻaneʻane i nā wahi āpau i hoʻomaka e hoʻololi ʻia e nā genotypes ʻē aʻe, a, i ka ʻaoʻao, hāʻawi i nā ala i nā mea hou. I ka hapanui o nā hihia, pili nā hoʻololi i ka hanana a me ka hoʻohui ʻana o nā lehulehu i nā ʻāina Asia me ka neʻe ʻana o nā genotypes hou mai waho. No laila, i Iapana, koe wale nō ka genotype JP-3, nā genotypes Kepanī ʻē aʻe i ʻike ʻia ma hope o US-1 (JP-1, JP-2, JP-3) ʻoi aku ka liʻiliʻi o ka hōʻoia kūwaho kūwaho hoʻi (Akino et al., 2011) ... I kēia manawa ʻekolu mau heluna nui o nā pathogen ma Kina, i loaʻa ka mahele o ka ʻāina. ʻAʻohe a i ʻole ka nāwaliwali kahena o ke kaona ma waena o kēia mau lehulehu (Guo et al., 2010; Li et al., 2013b). Ua hōʻike ʻia ʻo Genotype 13_A2 ma ka ʻāina ʻo Kina i nā panalāʻau hema (Yunnan a me Sichuan) ma 2005-2007, a ma 2012-1014. ua ʻike ʻia ma ka ʻākau hikina o ka ʻāina (Li et al., 2013b). I Inia, ua ʻikea paha ka 13_A2 i ka manawa like me Kina, ʻo ka nui paha me nā ʻuala hua maʻi (Chowdappa et al., 2015), a ma 2009-2010. ua hoʻokumu i kahi epiphytosis koʻikoʻi o ka hopena maikaʻi i ka tōmato ma ka hema o ka ʻāina, a ma hope ua pālahalaha ʻia i ka ʻuala a ma 2014 i hoʻomaka ai ka laha ʻana o ka hopena o ke ahi i West Bengal, a ua alakaʻi ʻia i ka luku a me ka pepehi kanaka o nā mahiʻai he nui (Fry, 2016).
ʻApelika
A hiki i ka 2008-2010 ʻaʻole i hoʻokō ʻia nā noiʻi ʻōnaehana o P. infestans ma nā ʻāina ʻApelika. I kēia manawa, hiki ke hoʻokaʻawale ʻia nā heluna o ʻApelika o P. infestans i ʻelua mau hui, a pili pili kēia ʻāpana me ka ʻoiaʻiʻo o ka lawe ʻia mai o kaʻuala hua mai ʻEulopa.
Ma ʻApelika ʻĀkau, ka mea e lawe ikaika mai i nā ʻuala hua mai ʻEulopa, ua hōʻike nui ʻia ka ʻano moʻa A2 ma nā ʻāpana āpau, kahi e hāʻawi ai i kahi kūpono i ke kū ʻana o nā genotypes hou ma muli o ka hoʻohui hou ʻana o ka moekolohe (Corbière et al., 2010; Rekad et al., 2017). Hoʻohui ʻia, ma Algeria, ʻike ʻia ka loaʻa ʻana o nā genotypes 13_A2, 2_A1, a me 23_A1 me ke kūlana kiʻekiʻe o nā mea mua o lākou, a me ka hoʻoliʻiliʻi hoʻi o ka mahele o nā genotypes kūikawā e hoʻopau i ka nalo ʻana (Rekad et al., 2017). I ka hoʻohālikelike ʻana i ke koena o ka ʻāina, ma Tunisia (me ka ʻokoʻa o ka ʻĀkau hikina o ka ʻāina), ʻo ka heluna pathogen e hōʻike nui ʻia e ka ʻano male male A1 (Harbaoui et al., 2014).
ʻO ka laina clonal NA-01 ka mea nui ma aneʻi. Ma ka laulā, ʻo ka nui o nā laina clonal i ka heluna kānaka he 43% wale nō. Ma ka Hikina a me ʻApelika Hema, kahi o ka nui o ka lawe ʻana mai o nā hua i ka liʻiliʻi liʻiliʻi (Fry et al., 2009), ua hōʻike ʻia ʻo P. infestans e nā laina A1-type clonal wale nō, US-1 a me KE-1, a hoʻoneʻe ka hope i ka mea ma mua i kaʻuala ( Pule et al., 2012; Njoroge et al., 2016). I kēia lā, loaʻa i kēia mau genotypes he helu i ʻike ʻia o nā ʻano subclonal.
ʻOʻAsuria
ʻO ka hōʻike mua o ka hopena o ka hopena o ka uala ma Australia i ka makahiki 1907, a ʻo ka epiphytotia mua, i manaʻo ʻia e nā ua nui i nā mahina kau wela, i ka makahiki 1909-1911. (Drenth et al., 2002). Ma ka laulā, akā, ʻaʻohe waiwai koʻikoʻi o ka hopena o ka hopena o ka ʻāina. ʻO nā laha Sporadic o ka hopena o ke ahi, i hoʻonāukiuki ʻia e nā ʻano anilā e hāʻawi ana i ka wela kiʻekiʻe, ʻaʻole pinepine pinepine ʻia ma mua o hoʻokahi manawa i kēlā me kēia 5-7 mau makahiki a noho ʻia ma ka ʻākau o Tasmania a me Victoria waena. E pili ana i nā mea i luna, nā puke i hoʻolaʻa ʻia no ke aʻo ʻana i ke ʻano o ka heluna o ko Austraigala heluna o P. infestans. ʻO ka ʻike hou loa i loaʻa mai 1998-2000. (Drenth et al., 2002). Wahi a nā mea kākau, ka lāhui Victoria he laina clonal US-1.3, ka mea i hoʻokūpaʻa ʻole i ka neʻe ʻana o kēia genotype mai United States. Ua hoʻokaʻawale ʻia nā hiʻohiʻona Tasmanian ma ke ʻano he AU-3, ʻokoʻa i nā genotypes i kū i kēlā manawa i nā wahi ʻē aʻe o ka honua.
Nā hiʻohiʻona o ka hoʻomohala ʻana i ka hopena o ka maʻi ahi i Lūkia
I ʻEulopa, ua hoʻolauna ʻia ka maʻi me nā kūpuna hua maʻi, nā oospores i hoʻopili ʻia i ka lepo, a me ka zoosporangia i lawe ʻia e ka makani mai nā mea kanu i ulu ʻia mai nā tubers i hoʻopili ʻia i nā mahina i ka makahiki i hala (nā mea kanu "manawaleʻa"), a i ʻole nā puʻu o nā culled lepe puke no ka mālama ʻana i nā tubers. ʻO kēia mau mea, ulu ʻia nā mea kanu i luna o nā ahu o nā kūpuna i hoʻolei ʻia i manaʻo ʻia ʻo ia ke kumu weliweli o ka maʻi. ma laila, he mea nui ka helu o nā tubers kupu, a hiki i ka zoosporangia ke lawe ʻia mai iā lākou ma kahi lōʻihi. ʻO ke koena o nā kumuwaiwai (oospores, nā mea kanu "manawaleʻa") ʻaʻole makaʻu loa ia, no ka mea ʻaʻole maʻamau ka ulu ʻana i nā mea kanu i nā māla like ma mua o hoʻokahi manawa i kēlā me kēia 3-4 mau makahiki. ʻO ka maʻi mai nā tubers hua maʻi he mea liʻiliʻi hoʻi ia ma muli o kahi ʻōnaehana kaohi maikaʻi o ka hua.
Ma ka laulaha, kaupalena ʻia ka nui o ka inoculum i nā heluna o ʻEulopa, a no laila ke lohi o ka piʻi ʻana o ka maʻi ahulau a hiki ke kāohi maikaʻi ʻia me ka hoʻomākaukau ʻana i nā fungicidal kemika. ʻO ka hana nui i nā kūlana ʻEulopa ka hakakā ʻana i ka maʻi i ka wā ke hoʻomaka ka hoʻolaha nui ʻana o ka zoosporangia mai nā mea kanu i hoʻopili ʻia.
I Rūsia, ʻokoʻa loa ke ʻano. ʻO ka hapa nui o kaʻuala a me nā hua kōmato e ulu i nā māla liʻiliʻi liʻiliʻi; ʻaʻole e hoʻokō ʻia nā hana pale ma luna o lākou, a i ʻole mālama ʻia nā lāʻau fungicidal i kahi helu lawa ʻole a hoʻomaka ma hope o ke ʻano o ka hopena o ke ahi i luna. ʻO ka hopena, hana nā māla mea kanu pilikino i kumu nui o ka maʻi, a mai laila i lawe ʻia ai e ka makani i nā kanu kanu pāʻoihana. Hōʻoia kēia e kā mākou nānā pono ʻana ma Moscow, Bryansk, Kostroma, Ryazan mau wahi: ʻike ʻia ka hōʻino ʻana i nā mea kanu i nā māla ponoʻī ma mua o ka hoʻomaka ʻana o ka mālama ʻana i nā fungicide o nā kanu kanu pāʻoihana. Ma hope iho, ua kāohi ʻia ka maʻi ahulau ma nā kahua nui e ka hoʻohana ʻana o nā hoʻomākaukau fungicidal, ʻoiai ma nā māla ponoʻī ke ulu wikiwiki nei o ka hopena o ke ahi.
I ka hihia kūpono ʻole a "hoʻomalu" mālama ʻana i nā mea kanu kalepa, ʻike ʻia ka foci o ka hopena o ka hopena i nā kula. ma hope ke hoʻomōhala nei lākou, e uhi ana i nā wahi nui loa (Elansky, 2015). ʻO ka maʻi i nā māla ponoʻī ka hopena nui i nā maʻi ʻino i nā māla kūʻai. Ma nā wahi ulu ʻuala a pau o Rūsia, ʻo ka wahi i noho ʻia e kaʻuala ma nā māla ponoʻī he nui aku ka manawa ma mua o ka nui o nā wahi o nā māla o nā mea hana nui. I loko o kēlā ʻano, hiki ke ʻike ʻia nā māla mea kanu pilikino ma ke ʻano he waiwai inoculum o ka honua āpau no nā māla ʻoihana. E hoʻāʻo e ʻike i kēlā mau waiwai i ʻano o nā genotypes o nā ʻano i nā māla ponoʻī.
Ke kanu nei i ka hua ʻole a me ka quarantine ka mālama ʻana o ka ʻuala, nā hua kōmato i loaʻa mai nā mea hana haole kānalua, ka mahi ʻana o ka ʻuala a me nā tōmato ma nā wahi like, nā lāʻau fungicide kūpono ʻole a i ʻole ka loaʻa ʻole ʻana e alakaʻi ai i nā epiphytoties koʻikoʻi i ka ʻāpana pilikino, a manuahi ka hopena o ia mea kea, hybridization a me ke hoʻokumu ʻana i nā oospores i nā māla ponoʻī. ʻO ka hopena, ʻike ʻia kahi ʻano genotypic kiʻekiʻe loa o ka pathogen, ke aneane like kēlā me kēia ʻano i kāna genotype (Elansky et al., 2001, 2015). Ke kanu nei i ka ʻuala hua o nā ʻano kumu ʻē aʻe ʻaʻole paha e kū mai nā laina clonal i mākaukau e hoʻouka i kekahi ʻano. Hoʻokaʻawale ʻia nā kānana i koho ʻia i kēlā hihia e ko lākou maʻalahi e pili ana i nā ʻano loli, ʻo ka hapa nui o lākou he kokoke i ka nui o nā genes virulence. He ʻokoʻa loa kēia mai ka ʻōnaehana o "nā laina clonal" maʻamau no nā māla nui o nā ʻoihana mahiʻai me kahi ʻōnaehana hoʻonohonoho pono o ka pale ʻana i ka hopena o ke ahi. "Nā laina Clonal" (ke hōʻike ʻia nā ʻano lōkō āpau o ka pathogen blight pathogen ma ke kula e hoʻokahi a ʻoi aku paha o nā genotypes) i nā ʻāina kahi e mālama ʻia ai ka ʻuala e nā mahiʻai nui wale nō: ʻo USA, Nūlani, Denmark, a pēlā aku. ke ulu nei kaʻuala, aia kekahiʻano genotypic kiʻekiʻe aʻe i nā māla ponoʻī. I ka hopena o ke kenekulia 20, ua pālahalaha nā "laina clonal" ma nā ʻĀkia a me nā ʻĀina hikina o Rūsia (Elansky et al., 2001), no ka hoʻohana ʻia o nā ʻano ʻuala like ʻole no ke kanu ʻana. I kēia mau lā, ua hoʻomaka ka loli o ke kūlana i kēia mau wahi i ka hoʻonui ʻana i ka genotypic o nā lāhui.
ʻO ka nele o nā mālama nui ʻana me nā hoʻomākaukau fungicidal i kekahi, hopena pololei - ʻaʻohe hōʻiliʻili o nā ʻano kūpale i nā māla. ʻOiaʻiʻo, hōʻike kā mākou hopena i ka loaʻa pinepine ʻole ʻana o nā kaha metalaxyl kūʻē i nā māla ponoʻī ma mua o nā kanu kanu pāʻoihana.
ʻO ka pili kokoke o kaʻuala a me nā mea kanu kōmato, maʻamau no nā māla ponoʻī, kōkua i ka neʻe ʻana o nā ʻano ma waena o kēia mau mea kanu, ma ke ʻano he hopena, i loko o nā makahiki he 1 i hala aku nei, i waena o nā kaila i hoʻokaʻawale ʻia mai nā ʻuala, ka mahele o nā kaha e hāpai ana i ka ʻāpana no ke kūʻē ʻana i nā ʻano o ke kōmato cherry (T1), ma mua wale nō ke ʻano no " ʻōkuhi "Tomato". ʻO nā kaha me ka moʻokūʻauhau TXNUMX i ka hapanui o nā hihia he koʻikoʻi loa i nā ʻuala a me nā kōmato.
I nā makahiki i hala iho nei, ua hoʻomaka ka hopena o ka mumū ma ke kōmato i nā hanana he nui ma mua o ka uala. Hiki ke hoʻopili ʻia nā ʻano kōmato me nā oospores i ka lepo, a i ʻole nā oospores i loaʻa i nā ʻano hua kōmato a i ʻole ke pili ʻana iā lākou (Rubin et al., 2001). I nā makahiki he 15 i hala aku nei, ua ʻike ʻia ka nui o nā ʻano pūʻulu inexpensive, i lawe nui ʻia, i loko o nā hale kūʻai, a ua hoʻololi ka hapa nui o nā mea hana liʻiliʻi i ka hoʻohana ʻana iā lākou. Loaʻa paha i nā ʻanoʻano nā kānana me nā genotypes maʻamau o nā wahi o kā lākou mahi ʻana. I ka wā e hiki mai ana, ua hoʻopili ʻia kēia mau genotypes i ke kaʻina moekolohe ma nā māla ponoʻī, kahi e alakaʻi ai i ka puka ʻana o nā genotypes hou loa.
No laila, hiki ke ʻōlelo ʻia ʻo nā māla ponoʻī he "ipu hoʻoheheʻe" ākea ia, ma muli o ke kūkaʻi ʻia ʻana o nā kumuwaiwai, hana ʻia nā genotypes a ʻike ʻia nā mea hou. Eia kekahi, ke ʻano o kā lākou koho ʻana i nā ʻano ʻokoʻa ʻokoʻa mai nā mea i hana ʻia no ka ʻuala i nā mahiʻai nui: ka ʻole o ka paʻi fungicidal, ka like ʻana o nā mea kanu, ka nui o nā mea kanu e loli ʻia e nā ʻano like ʻole o ka maʻi viral a me ka maʻi koʻohune, kahi kokoke i nā ʻōmato a me nā nighthades hihiu, ka hele ʻana a me ke kūkulu ʻana i ka hale no nā oospores e hana ma ke ʻano he kumu maʻi no ka makahiki e hiki mai ana.
Ke alakaʻi nei kēia āpau i kahi ʻano genotypic kiʻekiʻe loa o nā kua kua. I nā kūlana o nā epiphytotics i nā māla mea kanu, pāhola koke ka hopena a hoʻokuʻu ʻia nā nui o nā spores, e lele i nā kanu kanu ʻoihana kokoke. Eia nō naʻe, i ke komo ʻana i nā māla ʻoihana me ka ʻōnaehana pololei o ka ʻenehana mahiʻai a me ka pale kemika, ʻaʻole hiki i nā spores i hōʻea i kahi manawa kūpono e hoʻomaka ai i nā epiphytotics ma ke kula, no ka loaʻa ʻole o nā laina clonal e kūʻē i nā fungicides a loea i ka mea i mahi ʻia.
ʻO kekahi kumuwaiwai o ka inoculum mua i maʻi tubers i paʻa i nā mea kanu kalepa. Ua ulu kēia mau tubers, ma ke ʻano he rula, i nā māla me ka ʻenehana mahiʻai maikaʻi a me ka pale kemika ikaika. Hoʻohui ʻia nā genotypes o nā kaʻawale e hoʻopili i nā tubers i ka hoʻomohala ʻana i kā lākou ʻano ponoʻī. ʻOi aku ka weliweli o kēia mau ʻano no ke kanu pāʻoihana ʻana ma mua o ka inoculum i hoʻomaka ʻia mai nā māla ponoʻī. Kākoʻo nā hopena o kā mākou aʻo ʻana i kēia kuhi. Hoʻokaʻawale ʻia nā heluna kanaka mai nā kula nui me ka mālama pono ʻana i ka kemikū a me nā ʻenehana mahiʻai maikaʻi ʻaʻole ʻokoʻa i nā ʻano genotypic kiʻekiʻe. ʻO ka manawa pinepine kēia mau laina clonal i ikaika loa.
Hiki i nā kānana mai nā mea kanu pāʻoihana ke komo i nā lehulehu i nā māla mea kanu a pili i nā hana e hana nei i loko o lākou. Eia nō naʻe, i loko o kahi māla mea kanu, e emi loa ana kā lākou hoʻokūkū ma mua o kahi pāʻoihana, a pau koke lākou e noho i ke ʻano o ka laina clonal, akā hiki ke hoʻohana ʻia ko lākou mau gen i ka "māla" heluna kanaka.
ʻO ka maʻi e hoʻomohala ma nā mea kanu "manawaleʻa" a ma nā puʻu o nā kūpe culled i ka wā e ʻohi ʻana ʻaʻole pili pono ia no Rūsia, no ka mea I nā wahi ulu ʻuala nui o Rūsia, ʻike ʻia ka hau ʻana o ka hoʻoilo i ka hoʻoilo, a ʻo nā mea kanu mai nā tubers i hoʻoheheʻe i ka lepo ʻaʻole ulu iki. Eia kekahi, e like me ka hōʻike ʻana o kā mākou hoʻokolohua, ʻaʻole e ola ke konohiki hopena hopena i nā mahana maikaʻi ʻole ma nā tubers i hoʻomau i ko lākou hiki. I ka ʻāpana maloʻo, kahi e hoʻomaʻamaʻa ai i kaʻuala, hana ʻia ka lohi ma muli o ke kau maloʻo a me ka wela.
No laila, ke nānā nei mākou i kēia manawa i ka hoʻokaʻawale ʻia o nā P. infestans i nā "pā" a me nā "māla". Eia nō naʻe, i nā makahiki i hala iho nei, ua ʻike ʻia nā kaʻina hana e alakaʻi ana i ka hoʻololi a me ka interpenetration o nā genotypes mai kēia mau lehulehu.
Ma waena o lākou, hiki i kekahi ke hoʻomaopopo i ka hoʻonui maʻamau o ka heluhelu o nā mea hana liʻiliʻi, ka puka ʻana o nā pūʻolo liʻiliʻi o nā ʻuala hua, ka hoʻolaha ʻana o nā hoʻomākaukau fungicidal i nā pūʻulu liʻiliʻi, a me ka nalo o ka makaʻu i ka "chemistry" e ka lehulehu.
Kū aʻe nā kūlana i ka wā, mahalo i ka hana ikaika a hoʻokahi mea hoʻolako, kanu ʻia nā kauhale āpau me nā tubers ʻanoʻano like a hāʻawi ʻia me nā pūʻolo liʻiliʻi o nā pesticides like. Hiki ke manaʻo ʻia e ʻike ʻia kaʻuala o nā ʻano like ma nā kanu kanu ʻoihana kokoke.
Ma ka ʻaoʻao ʻē aʻe, ke hoʻolaha nei kekahi mau ʻoihana kūʻai pesticide i nā hoʻolālā mālama lāʻau "budgetary". I kēia hihia, hoʻowahāwahā ʻia ka helu o nā lāʻau lapaʻau i manaʻo ʻia a hāʻawi ʻia nā fungicides ʻoi loa, a ʻaʻole ka manaʻo koʻikoʻi i ka pale ʻana i ka ulu ʻana o ka hopena o ke ahi a hiki i ka ʻoki ʻana i nā piko, akā no ka lohi o ka epiphytoty i mea e hoʻonui ai i ka hua. Kūpono ʻia ia ʻano papa hana i ka wā e ulu ana i kaʻuala ʻūlū mai nā mea hua lulu haʻahaʻa, ke kumu ʻole ʻaʻohe nīnau no ka loaʻa ʻana o kahi hua kiʻekiʻe. Eia nō naʻe, i kēia hihia, i ka hoʻohālikelike ʻana i ka nui o nā māla, nā kāʻei kua i hoʻohaʻahaʻa ʻia o kaʻuala i ke koho ʻana o nā lāhui physiological kikoʻī, kahi weliweli loa no kēia ʻano.
Ma ka laulaha, ʻo nā ʻano i ka hui ʻana o "māla" a me nā ʻano "māla" o ka hana ʻana i kaʻuala me he mea weliweli loa ia iā mākou. I mea e pale aku ai i ko lākou hopena maikaʻi ʻole, ma ka ʻaoʻao a me nā ʻoihana kalepa, pono ia e kaohi i ka ʻāpana o nā ʻuala hua a me ka laulā o nā fungicides i hāʻawi ʻia i nā mea nona ponoʻī i ka liʻiliʻi liʻiliʻi, a me ka ʻimi ʻana i nā hoʻolālā pale ʻuala a me ka hoʻohana ʻana o ka hoʻomākaukau fungicidal i ka ʻoihana kalepa.
I nā wahi o ka ʻāpana pilikino, aia kahi ulu ikaika ʻana o ka wela wale nō, akā ʻo Alternaria kekahi. ʻO ka hapa nui o nā mea ʻona pilikino ʻaʻole e hana i nā hana kūikawā e pale aku ai iā Alternaria, e lawe nei i ka hoʻomohala ʻana o Alternaria no ka wili maoli ʻana o nā lau a i ʻole ke hoʻomohala ʻana i ka hopena o ke ahi. No laila, me ka hoʻomohala nui ʻana o Alternaria i nā ʻano susceptible, hiki i nā pā hale ke lilo i kumu no ka inoculum no nā mea kanu pāʻoihana.
Nā hana o ka loli
Kaʻina Mutation
ʻOiai ʻo ka hanana ʻana o nā mutation kahi kaʻina maʻamau e holo nei me kahi alapine haʻahaʻa, ʻo ka hanana ʻana o nā mutation ma kēlā me kēia wahi e pili ana i ke alapine o ka mutation o kēia locus a me ka nui o ka heluna kanaka. Ke aʻo ʻana i ke alapine o nā mutations o P. infestans ke kanana nei, ʻo ka helu o nā kolone e ulu ana ma ka pāpaho nutrient wae ma hope o ka mālama ʻana me nā kemika a i ʻole nā mutagens kino e hoʻoholo pinepine ʻia. E like me ka mea i ʻike ʻia mai ka ʻikepili i hōʻike ʻia ma ka Papa 8, hiki ke hoʻololi i ke alapine mutation o nā kānana like ma nā loci ʻokoʻa e kekahi mau kauoha o ka nui. ʻO ke alapine kiʻekiʻe o ka hoʻololi ʻana i ke kūpaʻa ʻana i ka metalaxyl kekahi o nā kumu no ka hōʻiliʻili ʻana o nā ʻano kūpale kūʻē iā ia i ke kūlohelohe.
ʻO ke alapine o ka hoʻololi pinepine a i hoʻokomo ʻia paha, i helu ʻia ma ke kumu o nā hoʻokolohua hoʻokolohua, ʻaʻole kūlike mau i nā kaʻina hana i kū i nā heluna maoli, no nā kumu aʻe:
1. I nā mokuna nukilika ʻaʻohe hiki, hiki ʻole ke kuhi i ke alapine o nā hoʻololi no kēlā me kēia hanauna nukelea. No laila, hāʻawi ka hapanui o nā hoʻokolohua i ka ʻike e pili ana i ke alapine o ka hoʻololi ʻana, me ka ʻole o ka hoʻokaʻawale ʻana ma waena o nā hanana mutational ʻelua a me kekahi hanana ma hope o ka mitosis.
2. Hoʻoemi ka hoʻololi ʻana o ka ʻaneki hoʻokahi i ke kaulike o ka genome, no laila, me ka loaʻa ʻana o kahi waiwai hou, hoʻemi ke ʻano holoʻokoʻa o ka meaola. ʻO ka hapa nui o nā hoʻokolohua i loaʻa i ka hoʻokolohua i hoʻemi ʻia ka hoʻokūkū ʻole a hoʻopaʻa ʻole ʻia i nā heluna maoli. No laila, ʻo ka coefficient hoʻopili ma waena o ke kēkelē o ke kūʻē ʻana o P. infestans mutants i nā phenylamide fungicides a me ka ulu ʻana o ka mea ulu ma kahi awelika (-0,62), a me ke kūʻē ʻana i nā fungicides a me ka hakakā ma nā lau ʻuala (-0,65) (Derevyagina et al. , 1993), e hōʻike ana i ke olakino haʻahaʻa o nā mutants. ʻO nā hoʻololi i ke kūpaʻa ʻana i dimethomorph pū kekahi me ka hōʻemi nui ʻana o ka hiki (Bagirova et al., 2001).
3. Hoʻomaha hou ka hapa nui o nā hoʻololi kūlohelohe a hoʻokomo ʻia a hōʻike ʻole iā lākou iho i ka phenotypically i nā hoʻokolohua, akā he waihona huna ia o ka loli i nā lāhui kūlohelohe. Hoʻokaʻawale ʻia nā ʻano Mutant i nā hoʻokolohua hale hōʻikeʻike e lawe i nā mutation mana a i ʻole semi-domant (Kulish a me Dyakov, 1979). Kohu mea lā, ua wehewehe ka nukelika diploidy i nā hoʻāʻo kūleʻa ʻole e kiʻi i nā mutants ma lalo o ka mana o UV irradiation i virulent i nā ʻano kūʻē mua (McKee, 1969). Wahi a nā helu a ka mea kākau, hiki ke hana i kēlā mau ʻano hoʻololi me ke alapine o lalo o 1: 500000. ʻO ka hoʻololi o nā hoʻololi recessive i kahi homozygous, phenotypically expression state hiki ke hana ma muli o ka moekolohe a asexual recombination (e ʻike ma lalo). Eia nō naʻe, i kēia hihia, hiki ke uhi ʻia ka hoʻololi e nā alelela mana o nā ʻano puʻupuʻu hihiu i ka mycelium cenotic (multinucleated) a paʻa phenotypically wale nō i ka hoʻokumu ʻia ʻana o nā zoospores mononuclear.
Papa 8. ʻO ka pinepine o ka P. infestans hoʻololi i nā mea ulu ulu-pāpā ma lalo o ka hana a nitrosomethylurea (Dolgova, Dyakov, 1986; Bagirova et al., 2001)
Ke Pili | Alapine haʻalulu |
Oxytetracycline | 6,9 x 10-8 |
ʻO Blasticidin S | 7,2 m 10-8 |
Streptomycin | 8,3 x10-8 |
Trichothecin | 1,8 x 10-8 |
Cycloheximide | 2,1 x 10-8 |
Daaconil | <4 x 10-8 |
ʻO Dimethomorph | 6,3 x 10-7 |
Metalaxil | 6,9 x 10-6 |
Pākani pū kekahi i nā nui o ka heluna kanaka i ka hanana o nā hoʻololi lōkahi. I nā heluna nui loa, kahi o ka helu o nā hunaola N> 1 / a, kahi o ka helu o ka hoʻololi ʻana, pau ka hoʻololi ʻana i mea ʻano maʻamau (Kvitko, 1974).
Hōʻike nā helu ʻana me ka awelika infestation o kahi māla potato (35 mau kiko ma kēlā me kēia meakanu), hana ʻia nā spore 8x1012 i kēlā me kēia lā ma hoʻokahi hectare (Dyakov a me Suprun, 1984). ʻIke ʻia, aia nā heluna like me nā mutation āpau i ʻae ʻia e ka ʻano o ka hoʻololi ma kēlā me kēia wahi. ʻOiai kahi hoʻololi laha ʻole, e hana ʻia me ke alapine o 10-9, e loaʻa ʻia e hoʻokahi tausani mau kānaka mai nā miliona e noho ana ma hoʻokahi hectare o kahi māla potato. No nā hoʻololi e kū ana me ke alapine kiʻekiʻe (e laʻa me, 10-6), i kēlā heluna kanaka, hiki i nā hoʻololi pālua like ʻole ke hana i kēlā me kēia lā (i ka manawa ʻelua ma nā loci) e hoʻololi ka hana mutational i ka hoʻohui hou ʻana.
Nā Mea Hele
No P. infestans, ʻike ʻia ʻelua ʻano nui o ka neʻe ʻana: e pani i kahi mamao (i loko o kahi māla ʻuala a i ʻole nā kaiāulu e pili kokoke ana) ma ka hoʻolaha ʻana i ka zoosporangia e nā ʻauwai ea a i ʻole ka ʻehu ua, a i nā mamao - me ke kanu ʻana i nā paipu a i lawe ʻia paha nā hua kōmato. Hāʻawi ke ala mua i ka hoʻonui ʻana o ke kiko o ka maʻi, ka lua - ka hana ʻana i foci hou i nā wahi mamao mai ke kumu.
ʻO ka pālahalaha ʻana o ka maʻi me nā kūmato a me nā huaʻai ʻaʻole kōkua wale i ka puka ʻana o ka maʻi i nā wahi hou, akā ʻo ia nō ke kumu nui o ka ʻokoʻa kūpuna i nā lehulehu. Ma ka moku ʻo Moscow, ulu ʻia kaʻuala, lawe ʻia mai nā wahi like ʻole o Lūkia a me ʻEulopa Komohana. Lawe ʻia nā hua Tomato mai nā wahi hema o Lūkia (Astrakhan ʻāpana, Krasnodar ʻāina, North Caucasus). ʻO nā hua Tomato, hiki ke lilo i kumu o ka maʻi (Rubin et al., 2001), lawe ʻia mai nā wahi hema o Lūkia, Kina, nā ʻāina ʻEulopa a me nā ʻāina ʻē aʻe.
Wahi a nā helu ʻana a E. Mayr (1974), nā loli kūpili i kahi heluna kaiāulu i hoʻokumu ʻia e nā hoʻololi i kākaʻikahi ʻia ma mua o 10-5 no kēlā me kēia wahi, ʻoiai i ka heluna lehulehu, ʻo ka hoʻololi ma muli o ke kahe o nā genes ma kahi o 10-3 - 10-4.
ʻO ka neʻe ʻana i ka tubers maʻi ka kuleana no ke komo ʻana o P. infestans i ʻEulopa, e hoʻolaha ana i nā wahi āpau o ka honua kahi e ulu ai kaʻuala; ua hoʻokumu lākou i nā loli lehulehu loa. Ua ʻike ʻia ka hopena o ka uala ma ka ʻāina o ka Lūkini Lūkini kokoke i ka manawa like me kona hiʻohiʻona i ʻEulopa Komohana.
Ma muli o ka ʻike mua ʻia o ka maʻi i ka makahiki 1846-1847 ma nā mokuʻāina ʻo Baltic a i loko wale nō o nā makahiki e hiki mai ana i pāhola ʻia i Belarus a me nā wahi komohana ʻākau o Lūkia, maopopo leʻa kona kumu ma ʻEulopa. ʻAʻole maopopo loa ke kumu mua o ka hopena maikaʻi i ka Old World. ʻO ke kuhiakau i hoʻomohala ʻia e Fry et al. (Fry et al., 1992; Fry, Goodwin, 1995, Goodwin et al., 1994) e hōʻike ana ua hele mai ka parasite mai Mekiko a i ʻAmelike ʻĀkau, kahi i laha ai ma o nā mea kanu, a laila lawe ʻia i Western Europe (fig. 7).
Ma ke ʻano he hopena hou o ka ʻauheʻe (hopena pālua o ka "bottleneck"), ua hiki i ʻEulopa nā klona hoʻokahi, a ua lilo kahi keiki i wahi maʻi ahulau ma nā panalāʻau o ka Old World kahi e ulu ai kaʻuala. Ma ke ʻano he hōʻike o kēia kuhiakau, ua kuhikuhi nā mea kākau, ʻo ka mea mua, ka mana o hoʻokahi wale nō ʻano moena (A1) a, ʻo ka lua, ʻo ka homogeneity o nā genotypes o nā ʻano aʻo i aʻo ʻia mai nā wahi like ʻole (pili lākou āpau i nā māka mole, e like me 2 isozyme loci, nā manamana lima o DNA, a me kūlike ke ʻano o DNA mitochondrial, a kūlike i ka clone US-1 i wehewehe ʻia ma USA). Eia nō naʻe, hāpai kekahi mau ʻikepili i nā kānalua e pili ana i kekahi o nā mea i hoʻolako ʻia o ke kuhiakau i ʻōlelo ʻia. Hoʻokaʻawale ʻia ʻo P. infestans mitochondrial DNA i hoʻokaʻawale ʻia mai nā hāpana potato herbarium i pākuʻi ʻia i ka wā epiphytotic mua o nā 40 e hōʻike ana he ʻokoʻa ke ʻano o ka DNA mitochondrial mai ka clone US-1, no laila, ʻo ka liʻiliʻi loa. ʻaʻole wale ke kumu o ka maʻi i ʻEulopa (Ristaino et al, 2001).
Ua kaumaha hou ke ʻano o ka hopena blight i nā makahiki 80 o ke kenekulia XX. Ua loaʻa nā hoʻololi hou:
1) Ua mahuahua ka ʻawelika awelika o ka heluna kanaka, a ʻo ia ka mea i alakaʻi aku ai, i ka laha ākea ʻana o ke ʻano pōʻino loa o ka hopena o ke ahi - pōʻino i nā petioles a me nā koʻokoʻo.
2) Aia kahi neʻe i ka manawa o ka hopena o ka mumū ma kaʻuala - mai ka hopena o Iulai a i ka mua o Iulai a hiki i ka hopena o Iune.
3) ʻO ka ʻano moʻa moʻo A2, i hele ʻole ma mua i ka Honua Kahiko, ua lilo a puni.
ʻO nā loli i mua o nā hanana ʻelua: ka hoʻohana nui ʻana o ka metalaxyl hou fungicide (Schwinn a me Staub, 1980) a me ka puka ʻana mai o Mekiko ma ke ʻano he kūkaʻi kalo o ka honua (Niederhauser, 1993). I kulike ai me kēia, ua hāʻawi ʻia i ʻelua mau kumu no ka loli o ka heluna lehulehu - ka hoʻololi ʻana i ka ʻano moena ma lalo o ka mana o metalaxyl (Ko, 1994) a me ka hoʻolauna nui ʻana o nā ʻano hou me nā tubers maʻi maʻi mai Mexico (Fry a me Goodwin, 1995). ʻOiai ʻo interconversions o nā ʻano moena ma lalo o ka mana o metalaxyl i loaʻa ʻaʻole na Ko wale nō, akā i nā hana i hana ʻia ma ka hale hōʻikeʻike o ke Kulanui o Moscow State (Savenkova, Chherepennicova-Anikina, 2002), ʻoi aku ka maikaʻi o ke kuhiakau ʻelua. Me nā hiʻohiʻona o ka lua o ka ʻano o ka moʻa ʻana, ua hoʻololi ʻia nā loli koʻikoʻi i nā genotypes o nā ʻano infestans Lūkini P., e like me nā genes kūlohelohe (isozyme a me RFLP loci), a me ke ʻano o DNA mitochondrial. ʻAʻole hiki ke wehewehe ʻia ka paʻakikī o kēia mau hoʻololi e ka hana a metalaxyl; akā, aia kahi lawe nui ʻia o nā ʻano hou mai Mekiko mai, ʻo ia ka mea i ʻoi aku ka hakakā (Kato et al., 1997), ua hoʻoneʻe i nā ʻano kahiko (US-1), a lilo i mea nui i ka lehulehu. ʻO ka hoʻololi o ka hui ʻana o nā lehulehu o ʻEulopa i kahi manawa pōkole loa - mai 1980 a 1985 (Fry et al., 1992). Ma nā panalāʻau o ka USSR ma mua, ua loaʻa nā "ʻano hou" i nā hōʻiliʻili mai Estonia ma 1985, ʻo ia hoʻi, ma mua o Polani a me Kelemania (Goodwin et al., 1994). ʻO ka manawa hope loa i hoʻokaʻawale ʻia ka "old strain US-1" ma Lūkia mai kahi kōmato i loaʻa i ka ʻāpana ʻo Moscow i ka makahiki 1993 (Dolgova et al., 1997). Ma Palani hoʻi, ua loaʻa nā ʻano "kahiko" i nā kanu kōmato a hiki i nā makahiki 90, ʻo ia hoʻi, ma hope o ko lākou nalo lōʻihi ʻana ma kaʻuala (Leberton a me Andrivon, 1998). Ua hoʻololi nā loli i ka P. infestans i nā ʻano he nui, e like me nā mea nui o ka hana, a hoʻonui i ka pōʻino o ke ākea i hala.
Hoʻohui hou ʻana i ka moekolohe
I mea e hoʻouluulu ai ka moekolohe ʻana i ka loli, pono ia, ʻo ka mua, ke kū ʻana o ʻelua ʻano o ka male ʻana i ka heluna kanaka i ka lakio kokoke i 1: 1, a ʻo ka lua, ke kū ʻana o ka loli lehulehu.
Kūlike ka lākiō o nā ʻano moena i nā heluna like ʻole a i nā makahiki like ʻole i hoʻokahi heluna kanaka (Papa 9,10, 90). ʻO nā kumu no ka hoʻololi nui ʻana i nā alapine o nā ʻano moena i ka heluna lehulehu (ʻoiai, ma Rūsia a i ʻole i Israel i nā 2002 mua o ke kenekulia i hala) ʻike ʻole ʻia, akā manaʻo ʻia no ka hoʻokumu ʻia o nā clone hoʻokūkū hou aku (Cohen, XNUMX).
Hōʻike kekahi o nā ʻikepili pili ʻole i ka papa o ka hana moekolohe i kekahi mau makahiki a me kekahi mau wahi.
1) Ua hōʻike ʻia nā noiʻi o nā heluna mai ka ʻāina ʻo Moscow ma 13 mau heluna kahi o ka mahele o ka ʻano male male A2 ma lalo o 10%, ʻo ka nui o nā ʻano like ʻole i helu ʻia no ʻekolu isozyme loci he 0,08, a ma 14 mau heluna kahi i ʻoi aku ai ka mahele o A2. 30%, ʻelua mau manawa ke kiʻekiʻe o ka moʻokūʻauhau (0,15) (Elansky et al., 1999). No laila, ʻo ke kiʻekiʻe o ka likelika o ka moekolohe, ʻo ka ʻoi aku ka nui o nā ʻano kūpuna o ka lehulehu.
2) ʻO ka pilina ma waena o ka lakio o nā ʻano moena i ka heluna lehulehu a me ka ikaika o ka hoʻokumu ʻana i ka oospore i ʻike ʻia ma Israel (Cohen et al., 1997) a me Holland.
(Flier et al., 2004). Ua hōʻike ʻia kā mākou noiʻi ʻana i nā heluna kanaka kahi i hoʻokaʻawale ʻia ai me ka ʻano moʻa A2 i helu ʻia no 62, 17, 9, a me 6%, ua loaʻa nā oospores ma 78, 50, 30, a me 15% o nā lau uala i kālailai ʻia (me 2 a ʻoi aku nā kiko).
ʻO nā laʻana me 2 a ʻoi paha nā kiko e ʻike ʻia nā oospores ma mua o nā laʻana me 1 kiko (32 a me 14% o nā laʻana,) (Apryshko et al., 2004).
ʻOi aku ka maʻamau o Oospores i nā lau o ka papa waena a me lalo o ka mea kanu kalo (Mytsa et al., 2015; Elansky et al., 2016).
3) I kekahi mau wahi, ua ʻike ʻia nā genotypes kūikawā, ʻo ka hanana i pili me ka recombination moekolohe. No laila, ma Polani i ka 1989 a me Palani i ka makahiki 1990, hoʻoikaika i ka homozygous no ka glucose-6-
phomerate isomerase (GPI 90/90). Ma mua o ka loaʻa ʻana o 10/90 heterozygotes wale nō no 100 mau makahiki, pili ka homozygosity i ka hoʻoliʻiliʻi wahine (Sujkowski et al., 1994). Ma Colombia (USA), he mea maʻamau nā mea hoʻokaʻawale e hui pū ana i ka A2 me GPI 100/110 a me A1 me GPI 100/100, akā i ka hopena o ke kau 1994 (ʻAukake 16 a me Kepakemapa 9), nā ʻōpala me nā genotypes recombinant (A1 GPI 100/110 a me A2 GPI 100/100) (Miller et al., 1997).
4) I kekahi mau heluna mai Polani (Sujkowski et al., 1994) a me ka North Caucasus (Amatkhanova et al., 2004), ʻo ka hoʻokaʻawale ʻana o DNA loci a me nā loio protein allozyme e like me ka mahele o Hardy-Weinberg, e hōʻike ana
e pili ana i ka mahele kiʻekiʻe o ka hāʻawi o ka recombination moekolohe i ka loli o ka lehulehu. I nā wahi ʻē aʻe o Rūsia, ʻaʻohe palapala i ka hoʻokaʻawale ʻia o Hardy-Weinberg i nā heluna kanaka i loaʻa, akā ua hōʻike ʻia ke kū ʻana o ka maʻi maʻi, e hōʻike ana i ka nui o ka hana hou ʻana o ka clonal (Elansky et al., 1999).
5) ʻO ka moʻomeheu genetic (GST) ma waena o nā ʻano me nā ʻano moena like ʻole (A1 a me A2) i haʻahaʻa ma mua o waena o nā lāhui like ʻole (Sujkowski et al., 1994), ka mea hōʻike ʻole i nā keʻa kolohe.
I ka manawa like, ʻaʻole hiki ke kiʻekiʻe ka hāʻawi o ka recombination moekolohe i ka lehulehu o ka lehulehu. Ua helu ʻia kēia makana no nā lehulehu o ka ʻāina ʻo Moscow (Elansky et al., 1999). Wahi a nā helu a Lewontin (1979), "recombination, hiki ke hana i nā mea hou mai nā loci ʻelua me ke alapine e ʻoi aku i ka huahana o kā lākou heterozygosities, e lilo i mana wale nō inā kiʻekiʻe nā waiwai o ka heterozygosity no nā alela ʻelua."
Me ka lakio o nā ʻano pālua, ka mea maʻamau no ka ʻāpana o Moscow, e like me 4: 1, ʻo 0,25 ka pinepine o ka recombination. ʻO ka likelika e heterozygous kea i kēlā me kēia o nā ʻekolu i hoʻopaʻa ʻia i ka isozyme loci i loko o nā heluna i aʻo ʻia he 0,01 (2 mau hanana mai ka 177). No laila, ʻo ka likelika o ka hanana heterozygotes pālua ma muli o ka hopena o ka hoʻohui hou ʻana ʻaʻole pono e ʻoi aku i kā lākou huahana i hoʻonui ʻia e ka likelika o ke kuea (0,25x0,02x0,02) = 10-4, i. ʻaʻole hāʻule ka recombinants moekolohe i ka laʻana i aʻo ʻia o nā kanana. Ua hana ʻia kēia mau helu no nā heluna kanaka mai ka moku ʻo Moscow, kahi ʻano e ka loli kiʻekiʻe loa. I nā heluna monomorphic e like me nā Siberian, ke kaʻina moekolohe, ʻoiai inā kū ʻia i nā heluna kanaka ʻokoʻa, ʻaʻole hiki ke hoʻohuli i kā lākou ʻano ʻokoʻa.
Hoʻohui ʻia, ʻike ʻia ʻo P. infestans e ka misalignment chromosome pinepine i ka meiosis, a hiki i ka aneuploidy (Carter et al., 1999). ʻO ia mau hana hewa e hōʻemi i ka momona o nā hybrids.
Hoʻohui hou ʻana o Parasexual, hoʻololi i nā gen mitotic
I nā hoʻokolohua e pili ana i ka mahele ʻana o nā P. infestans a me nā hoʻololi ʻana i ke kūpaʻa ʻana i nā mea kūpaka ulu ʻokoʻa, ua loaʻa ka puka ʻana o nā misolates kūʻē i nā mea pāpā ʻelua (Shattock and Shaw, 1975; Dyakov, Kuzovnikova, 1974; Kulish, Dyakov,
1979). Kū nā kūʻē kū i nā mea pale ulu ʻelua ma muli o ka heterokaryotization o ka mycelium, a i kēia hihia ua pili lākou i ka wā o ka hānau ʻia ʻana e nā monosponsal zoospores (Judelson, Ge Yang, 1998), a i ʻole pili ʻole i nā keiki monozoosporous, no ka mea ua loaʻa iā lākou tetraploid (ʻoiai nā isolate mua diploid) nuclei (K , 1979). Hoʻokaʻawale ʻia nā Heterozygous diploids ma kahi haʻahaʻa haʻahaʻa ma muli o ka haploidization, chromosome nondisjunction, a me ka mitotic crossing (Poedinok et al., 1982). Hiki ke hoʻonui ʻia ke alapine o kēia mau kaʻina hana me ke kōkua o kekahi mau hana ma nā diploids heterozygous (ʻo kahi laʻana, UV irradiation o germores spores).
ʻOiai ke hoʻokumu ʻia ʻana o nā hybrids vegetative me nā pale pālua ʻaʻole kū i ka vitro wale nō, akā i nā tubers potato i pā ʻia me ka huikau o nā mutants (Kulish et al., 1978), paʻakikī e nānā i ke kuleana o ka hoʻohui hou ʻana o ka parasexual i ka hanauna o nā genotypes hou i nā lehulehu. ʻO ke alapine o ka hoʻokaʻawale ʻana ma muli o ka haploidization, nondisjunction o chromosome a me ka mitotic crossing over without special effects is negligible (ma lalo o 10-3).
Hiki ke hoʻokumu ʻia nā mea hoʻokaʻawale homozygous o nā heterozygous ma ka hele ʻana o ka mitotic a me ka hoʻololi ʻana o ka gen mitotic, ma P. sojae i loaʻa me ke alapine o 3 x 10-2 a 5 x 10-5 i kēlā me kēia wahi, kaukaʻi ʻia (Chamnanpunt et al. , 2001).
ʻOiai ʻo ke alapine o ka hanana heterokaryons a me nā heterozygous diploids i lilo i kiʻekiʻe kiʻekiʻe (hiki i nā ʻumi haneli pākēneka), ke hana ʻia kēia hana ke loaʻa nā moʻomeheu mutant i loaʻa mai nā kānana like. Ke hoʻohana nei i nā ʻano ʻokoʻa i hoʻokaʻawale ʻia mai ke kūlohelohe, ʻaʻole hiki mai ka heterokaryotization (a i ʻole me ke alapine haʻahaʻa loa) ma muli o ke kūlike ʻole o nā mea kanu (Poedinok a me Dyakov, 1981; Anikina et al., 1997b; Cherepennikova-Anikina et al., 2002). No laila, hiki ke hoʻoliʻiliʻi i ke kuleana o ka recombination parasexual i ka recombination intraclonal i nā nuclei heterozygous a me ka hoʻololi ʻana o kēlā me kēia gen i kahi kūlana homozygous me ka ʻole o ke kaʻina moekolohe. Kēia kaʻina hana paha o ka epidemiological nui i loko o nā kānana me recessive a semi-domant fungicide resistation mutations. ʻO kāna hoʻololi ʻana i kahi kūlana homozygous ma muli o ke kaʻina parasexual e hoʻonui i ke kūpaʻa o ka lawe o ka hoʻololi (Dolgova, Dyakov, 1986).
Hoʻokomo o nā gen
Hiki i nā ʻano Heterothallic Phytophthora ke hoʻopili me ka hoʻokumu ʻana i nā oospores hybrid (e ʻike iā Vorobʻeva a me Gridnev, 1983; Sansome et al., 1991; Veld et al., 1998). ʻO ka hybrid kūlohelohe o nā ʻano Phytophthora ʻelua i ikaika loa i luku ʻia he tausani mau alders ma UK (Brasier et al., 1999). Hiki iā P. infestans ke hana me nā ʻano ʻē aʻe o ka genus (P. erythroseptica, P. nicotianae, P. Cactorum, a me nā mea ʻē aʻe) ma nā mea kanu hoʻokipa maʻamau a ma ka lepo, akā he mea liʻiliʻi ka ʻike ma nā puke e pili ana i nā hybrids interspecific. Ma lalo o nā keʻena hoʻokolohua, ua loaʻa nā hybrids ma waena o P. infestans a me P. Mirabilis (Goodwin a me Fry, 1994).
Papa 9. ʻO ka mahele o nā P. infestans ʻano me nā ʻano moena A2 i nā ʻāina like ʻole o ka honua i ka manawa mai 1990 a 2000 (e like me ka ʻike o nā kumuwaiwai wehe puke a me nā pūnaewele www.euroblight.net, www.eucablight.org)
aina | 1990 | 1991 | 1992 | 1993 | 1994 | 1995 | 1996 | 1997 | 1998 | 1999 | 2000 |
---|---|---|---|---|---|---|---|---|---|---|---|
Belarus | 33 (12) | 34 (29) | |||||||||
Pelekiuma | 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) | ||||||||||
'Onelani | 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) | ||||||||||
Aialani | 4 (145) | ||||||||||
Ke Akau. Ilelani | 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) | ||||||
ʻO Scotland | 25 (147) | 11 (163) | 22 (189) | 5 (22) | |||||||
Kuekene | 25 (263) | 62 (258) | 49 (163) | ||||||||
Wales | 0 (16) | 7 (97) | 0 (48) | 0 (25) | |||||||
ʻO 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) | ||||||||||
ʻO Morocco | 60 (108, 1997-2000) | 52 (25) | 42 (40) | ||||||||
Serbia | 76 (37) | ||||||||||
Mexico (Toluca) | 28 (292, 1988-89) | 50 (389, 1997-98) |
Papa 10. ʻO ka mahele o nā P. infestans kaha me ka ʻano moena A2 i nā ʻāina like ʻole o ka honua i ka manawa mai 2000 a 2011.
aina | 2001 | 2002 | 2003 | 2004 | 2005 | 2006 | 2007 | 2008 | 2009 | 2010 | 2011 |
---|---|---|---|---|---|---|---|---|---|---|---|
Austria | 65 (83) | ||||||||||
Belarus | 42 (78) | ||||||||||
Pelekiuma | 20 (102 *) | 4 (32) | 50 (14) | 25 (16) | 62 (13) | 54 (26) | 70 (54) | 30 (23) | 29 (35) | 62 (71) | 45 (49) |
Kuikilana | 89 (19) | ||||||||||
ʻO 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) | |||||
'Onelani | 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 | |||||||
Ke Akau. Ilelani | 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) | ||
ʻO Scotland | 3 (213) | 2 (474) | 24 (135) | 86 (337) | 88 (386) | 74 (172) | |||||
Kuekene | 60 (277) | 39 (87) | |||||||||
ʻO Slovakia | 0 (36) | 14 (26) | 62 (26) | 0 (26) | |||||||
Wales | 25 (12) | 68 (106) | 80 (88) | 92 (143) | 75 (45) | ||||||
ʻO Korea | 46 (26) | ||||||||||
ʻO Brazil | 0 (49) | 0 (30) | |||||||||
China | 10 (30) | 0 (6) | 0 (6) | ||||||||
ʻO Vietnam | 0 (294, 2003-04) | ||||||||||
ʻO Uganda | 0 (8) |
ʻO Dynamics o ka genotypic hoʻohui o nā lehulehu
Hiki ke hoʻololi i nā genotypic o P. infestans i lalo o ka mana o ka neʻe ʻana o nā klone hou mai nā wahi ʻē aʻe, nā hana mahiʻai (hoʻololi i nā ʻano, noi o nā fungicides), a me nā kūlana anilā. Hoʻololi nā hopena kūwaho i nā klone ʻokoʻa i nā ʻanuʻu like ʻole o ke ola ola;
Ka hopena o ka ʻano
ʻO nā mea kanu hou me nā gen genes no ke kūʻē kūʻē (R-genes) kahi mea koho koho mana e koho i nā clones me nā genul virulence komplementary i ka P. infestans populations. I ka loaʻa ʻole o ke kūpaʻa nonspecific i loko o ka ʻano ʻuala e kāohi i ka ulu ʻana o ka heluna pathogen, hikiwawe loa ke kaʻina hana o ke kūapo ʻana i nā clones koʻikoʻi i ka heluna kanaka. No laila, ma hope o ka pālahalaha ʻana ma ka moku ʻo Moscow o ka Domodedovsky ʻano, i loaʻa ka ʻāpana kūʻē R3, ua hoʻonui ʻia ke alapine o nā clone virulent no kēia ʻano mai 0,2 a 0,82 i hoʻokahi makahiki (Dyakov, Derevjagina, 2000).
Eia nō naʻe, ʻo ka loli i nā alapine o nā genul virulence (pathotypes) i nā lehulehu ʻaʻole wale ma lalo o ka mana o nā ʻano ʻuala i mahi ʻia. ʻO kahi laʻana, ma Belarus a hiki i ka 1977, noho aliʻi nā clone me nā genul virulence 1 a me 4, i kumu ʻia e ka mahi ʻana o nā ʻano ʻuala me nā genes kūʻē R1 a me R4 (Dorozhkin, Belskaya, 1979). Eia nō naʻe, i ka hopena o nā 70 o ke kenekulia 2002, ʻike ʻia nā klone me nā genul virulence ʻokoʻa a me kā lākou hui pū ʻana, a ʻaʻole i hoʻohana ʻia nā genes resistence kompliment i ka hoʻoulu ʻana i kaʻuala (genes virulence keu) (Ivanyuk et al., XNUMX). ʻO ke kumu o ka helehelena o kēlā mau clone, no ka neʻe ʻana i ʻEulopa o nā mea maʻi mai Mekiko mai me nā kūmaka potato. Ma ka home, ua hoʻomohala ʻia kēia mau clone ma luna o kaʻuala i mahi ʻia wale nō, akā, ma nā ʻano hihiu hoʻi e halihali ana i nā ʻano kūʻē kūʻē; no laila, pono ka hui ʻana o nā genes virulence he nui i loko o ka genome e ola ai i kēlā mau ʻano.
No nā ʻano me ke kūʻē ʻole nonspecific, ʻo lākou, ma ka hoʻēmi ʻana i ka helu o ka hānau hou ʻana o ka pathogen, hoʻolohi i ka ulu ʻana o kāna mau lehulehu, a, e like me ka mea i hōʻike ʻia aʻe nei, kahi hana o ka helu. ʻOiai polygenic ka mea hoʻokūkū, nā clones i loaʻa ka nui o nā genes no ka "agresive" e hōʻiliʻili i ka wikiwiki o ka nui o ka nui o ka lehulehu. No laila, ʻaʻole nā lāhui huhū nui kahi huahana o ka hoʻopili ʻana i nā ʻano mahiʻai me ke kūʻē ʻole nonspecific, akā, i ka ʻaoʻao ʻē aʻe, e ʻike ʻia paha i nā kanu ʻana o nā ʻano maʻalahi hiki ke ʻākoakoa o nā spore parasite.
No laila, ma Lūkia, ua loaʻa nā lehulehu ʻoi loa o P. Infestans i nā ʻāpana o nā epiphytoties makahiki (nā lehulehu mai Sakhalin, Leningrad, a me nā wahi ʻo Bryansk). ʻO ke kūpikipiki o kēia mau lehulehu i ʻoi aku ke kiʻekiʻe ma mua o ka Mekiko (Filippov et al., 2004).
Eia kekahi, ʻoi aku ka liʻiliʻi o nā oospores i hoʻokumu ʻia i nā lau o nā ʻano kūpale ma mua o nā mea maʻalahi (Hanson a me Shattock, 1998), ʻo ia hoʻi, ʻo ke kūʻē ʻole nonspecific o ka ʻano e hoʻoliʻiliʻi hoʻi i nā hiki ke hoʻohui hou i nā parasite a me ka hiki ke loaʻa nā ʻano hoʻoilo hoʻoilo.
Ka hopena o nā fungicides
ʻAʻole hōʻemi wale nā fungicides i ka helu o nā fungi phytopathogenic, ʻo ia hoʻi pili i ka nui o ko lākou heluna kanaka, akā hiki iā lākou ke hoʻololi i nā alapine o nā genotypes pākahi, ʻo ia hoʻi hoʻohuli i ka hopena qualitative o nā lehulehu. Ma waena o nā hōʻailona koʻikoʻi o ka lehulehu e loli ana ma lalo o ka hopena o fungicides penei: nā hoʻololi i ke kūpaʻa ʻana i nā fungicides, nā loli i ka hakakā a me nā virulence, a me nā hoʻololi o nā ʻōnaehana hānau.
Ka hopena o nā fungicides i ke kūpaʻa a me ka hakakā o nā lehulehu
Hoʻoholo ʻia ke kēkelē o kēia mana, ka mea mua, e ke ʻano o ka fungicide i hoʻohana ʻia, a hiki ke hoʻokaʻawale ʻia i ka polysite, oligosite a me monosite.
Hoʻopili ka mea mua i nā fungicides pili. ʻO ke kūʻē ʻana iā lākou (inā he hiki i nā mea āpau) kaohi ʻia e kahi nui o nā ʻāpana nāwaliwali loa e hōʻike. Hoʻoholo kēia mau waiwai i ka loaʻa ʻole o nā loli i ʻike ʻia i ke kūpaʻa o ka heluna kanaka ma hope o ka mālama ʻana me nā fungicides (ʻoiai i loko o kekahi mau hoʻokolohua ua loaʻa kekahi piʻi ʻana o ke kūʻē ʻana). Mālama ʻia ka heluna heluna ma hope o ka pīpī ʻana me nā fungicides e hoʻopili ʻia i ʻelua mau pūʻulu o nā ʻano:
1) Mālama ʻia nā kaula i nā wahi o nā mea kanu ʻaʻole mālama ʻia me ka lāʻau. ʻOiai ʻaʻohe pilina me ka fungicide, ʻaʻole loli ka hakakā a me ke kūpaʻa ʻana o kēia mau ʻano.
2) Pākuʻi ʻia ka hoʻopili ʻana me ka fungicide, ʻo ke kukuna ma nā kiko o ka hoʻopili ma lalo o ka make. E like me ka mea i ʻōlelo ʻia ma luna, ʻaʻole i loli ke kūʻē ʻana o kēia ʻāpana o ka lehulehu, akā naʻe, ma muli o ka hopena ʻino o ka fungicide a hiki i ka sublethal concentration e pili ana i ka metabolism o ka fungal cell, ka pono laulā a me kāna mea parasite, ka huhū, hoʻemi (Derevyagina a me Dyakov, 1990).
No laila, ʻo kahi ʻāpana o ka heluna kanaka i make ʻole, hōʻike ʻia i ka hoʻopili ʻana me ka fungicide, he palupalu ka huhū a ʻaʻole hiki ke lilo i kumu no nā epiphytotics. No laila, ʻo ka mālama pono ʻana e hōʻemi ana i ke alapine o ka heluna o ka heluna kanaka i pili ʻole me ka fungicide kahi kūlana no ka kūleʻa o nā hana pale. ʻO ke kūpaʻa ʻana i nā fungicides oligosite e hoʻomalu ʻia e nā gen additive.
ʻO ka hoʻololi ʻana o kēlā me kēia ʻāpana e alakaʻi ai i kahi hoʻonui o ke kūʻē ʻana, a ʻo ka pae holoʻokoʻa o ke kūpaʻa ʻana ma muli o ka hoʻohui o ia mau hoʻololi. No laila, piʻi ka piʻi ʻana o ka kūpaʻa. ʻO kahi laʻana o ka piʻi iki o ka kūʻē ʻana, ʻo ia ka hoʻololi ʻana i ke kūpaʻa ʻana i ka dimethomorph fungicide, kahi e hoʻohana ākea ʻia e pale ai i kaʻuala mai ka hopena hope loa. ʻO ke kūpaʻa Dimethomorph polygenic a hoʻohui. Hoʻonui iki kahi hoʻololi ʻanuʻu i ke kūpaʻa.
ʻO kēlā me kēia hoʻololi hou e hōʻemi i ka nui o ka pahuhopu a, no laila, ke alapine o nā hoʻololi hou (Bagirova et al., 2001). ʻO ka piʻi ʻana o ka pale awelika o ka heluna kānaka ma hope o ka hana pinepine ʻana me ka oligosite fungicide e hele a me ka mālie. Hoʻoholo ʻia ka wikiwiki o kēia kaʻina e ʻekolu mau kumu: ke alapine o ka hoʻololi ʻana o nā ʻenemi kūʻē, ka coefficient o ke kūpale ʻana (ka lākiō o ka hopena make o kahi kānana kūpaʻa e pili ana i ka mea maʻalahi) a me ka hopena o nā hoʻololi ʻana i nā ʻeneki kūʻē i ka hoʻoikaika kino.
ʻO ke alapine o ka hanana o kēlā me kēia hoʻololi hou ma lalo o ka mea ma mua o ka mea ma mua; no laila, he ʻano hoʻoliʻiliʻi ke kaʻina (Bagirova et al., 2001). Eia nō naʻe, inā hana ʻia nā hana hoʻohui hou (moe kolohe a i ʻole parasexual) i kahi heluna kanaka, a laila hiki ke hoʻohui i nā hoʻololi like ʻole o nā mākua i kahi kānana hybrid a hoʻonui i ke kaʻina hana. No laila, loaʻa i nā lehulehu panmix ke kūʻē ʻoi aku ka wikiwiki ma mua o nā lāhui agamika, a i nā mea hope loa, nā lehulehu i loaʻa ʻole nā mea pale incompatibility vegetative i ʻoi aku ka wikiwiki ma mua o nā lehulehu i hoʻokaʻawale ʻia e kēlā mau pale. E pili ana i kēia, ke kū nei nā ʻano i nā lehulehu i ʻokoʻa i nā ʻano o ka moʻa e hoʻonui i ke kaʻina o ka loaʻa ʻana o ke kūʻē ʻana i nā fungicides oligosite.
ʻAʻole hāʻawi ka lua a me ke kolu i nā mea i ka hōʻiliʻili wikiwiki ʻana o nā ʻano dimethomorph-resistence i nā lehulehu. Kēlā me kēia hoʻololi aʻe ma kahi o ka palua o ke kūʻē, he mea nui ʻole ia, a ma ka manawa like e hoʻēmi ai i ka nui o ka ulu ʻana i kahi ʻano hana a me ka hakakā (Bagirova et al., 2001; Stem, Kirk, 2004). ʻO ia nō paha ke kumu i loaʻa ʻole ai nā ʻino kūʻē i waena o nā kūlohelohe P. infestans kūlohelohe, ʻo nā mea hoʻi i hōʻiliʻili ʻia mai nā mea kanu uala i mālama ʻia me ka dimethomorph.
ʻO ka heluna kanaka i mālama ʻia me ka fungis oligosite e komo pū i ʻelua mau ʻano o ke kanana: ʻo nā mea i launa ʻole me ka fungicide, a no laila ʻaʻole i hoʻololi i nā ʻano mua (inā loaʻa nā ʻano kūpale i waena o kēia hui, ʻaʻole lākou e hōʻiliʻili ma muli o ke kiʻekiʻe kiʻekiʻe a me ka hoʻokūkū o nā ʻano koʻikoʻi). a me nā kānana i ka pili ʻana me nā concentrations sublethal o ka fungicide. Aia ia i waena o ka hope i hiki ke hōʻiliʻili ʻia nā ʻano kūʻē kūpale, no ka mea, eia nā mea maikaʻi aʻe ma mua o nā mea maʻalahi.
No laila, ke hoʻohana nei i nā fungicides oligosite, ʻaʻole ia he lapaʻau maikaʻi loa e like me ka nui o ka lāʻau lapaʻau, ʻoi aku ka nui o nā manawa ma mua o ka lāʻau make, no ka mea me ka mutagenesis stepwise, haʻahaʻa ke kūʻē mua o nā ʻano i hoʻololi ʻia.
ʻO ka mea hope loa, hōʻike nui nā hoʻololi ʻana i ke kūpaʻa ʻana i nā fungicides monosite, ʻo ia hoʻi, hiki i kahi hoʻololi ke hōʻike i kahi kiʻekiʻe o ke kūpaʻa ʻana, a hiki i ka hoʻopau ʻana i ka nalowale o ka ʻike. No laila, wikiwiki ka hoʻonui ʻana i ke kūpaʻa o nā lehulehu.
ʻO kahi laʻana o ia mau fungicides he phenylamides, me ka metalaxyl fungisida maʻamau. Hana ʻia nā hoʻololi ʻana o ke kūʻē me ke alapine kiʻekiʻe, a kiʻekiʻe loa ke kiʻekiʻe o ke kūpaʻa ʻana i nā mutants - ua ʻoi aku ia i nā kānana maʻalahi e kahi helu o hoʻokahi kaukani aʻoi aku paha (Derevyagina et al., 1993). ʻOiai ke emi nei ka helu ulu a me ka hakakā o nā mutants kūʻē e kūʻē i ke kumu o ka make ʻana o nā ʻano susceptible mai kahi fungicide ʻōnaehana, ke ulu wikiwiki nei ka helu o ka lehulehu kūpaka a ke piʻi nei kona huhū i ka like. No laila, ma hope o kekahi mau makahiki o ka hoʻohana ʻana i ka fungicide, ʻaʻole hiki i ka hoʻokūkū paʻakikī ke kūlike i ka huhū o nā mea maʻalahi, akā ʻoi aku hoʻi ma mua (Derevyagina, Dyakov, 1992).
ʻO ka hopena ma ka recombination moekolohe
Ma muli o ka hanana pinepine o ka ʻano moʻa A2 i ka P. infestans heluna kanaka i kūlike me ka hoʻohana nui ʻana o ka metalaxyl e kūʻē i ka hopena o ke ahi, ua ʻōlelo ʻia e hoʻohuli ka metalaxyl i ka hoʻohuli ʻano male ʻana. I P. parasitica, ua hoʻāʻo hoʻokolohua ʻia kēlā ʻano hoʻololi ma lalo o ka hana a chloroneb a me metalaxyl (Ko, 1994). ʻO kahi alahele ma kahi mehana me ka hoʻohaʻahaʻa haʻahaʻa o metalaxyl i alakaʻi ʻia i ka puka ʻana o ka homothallic isolates mai kahi ʻano o P. infestans sensitive i ka metalaxyl me ka moʻo ʻano A1 (Savenkova a me Cherepnikova-Anikina, 2002). I loko o nā paukū e hiki mai ana ma luna o ka pāpaho me kahi kiʻekiʻe o ka metalaxyl, ʻaʻole i ʻike ʻia kahi kaʻawale o ke ʻano pālua A2, eia nō naʻe, ʻo ka hapa nui o nā kaʻawale, ke hele ʻia me nā kaʻawale A2, ma kahi o nā oospores, hana i nā hōʻiliʻili mycelium ʻino a he manuahi. Nā ʻāpana o kahi kānana kūpaʻa e loaʻa ana i ka ʻano moʻa A2 ma ka pāpaho me kahi kiʻekiʻe o ka metalaxyl i ʻae iā mākou e ʻike i ʻekolu mau ʻano o nā loli ʻano moena: 1) hoʻopau piha i ka wā kea me nā kaʻawale A1 a me A2; 2) homotallism (ke hoʻokumu ʻana i nā oospores i kahi monoculture); 3) ka hoʻohuli ʻana o ka ʻano pilina A2 i A1. No laila, hiki i ka metalaxyl ke hana i nā loli i ke ʻano o ka moʻa ʻana i ka P. infestans heluna kanaka, a no laila, ke kū ʻana o ka moekolohe i loko o lākou.
Ka hopena i ka recombination vegetative
Ua hoʻonui kekahi mau genes resistence antibiotic i ke alapine o ka hehalokaryotization hyphal a me ka diploidization nukelea (Poedinok a me Dyakov, 1981). E like me ka mea i hōʻike ʻia ma mua, heterokaryotization o hyphae i ka hoʻohui ʻana o nā ʻano like ʻole o P. infestans he mea liʻiliʻi loa ia ma muli o ke ʻano o ka incompatibility vegetative i kēia fungus. Eia nō naʻe, hiki i nā genes no ke kūʻē ʻana i kekahi mau antibiotic ke loaʻa nā hopena ʻaoʻao, i hōʻike ʻia i ka lanakila ʻana i ka incompatibility vegetative. Ua loaʻa kēia waiwai e ka ʻenemi kūʻē streptomycin 1S-1 mutant. Hiki ke hoʻonui i ke kahe o nā genes ma waena o nā kānana a me ka wikiwiki i ka hoʻololi ʻana o ka lehulehu holoʻokoʻa i nā ʻano hou a i ʻole fungicides.
Hiki i kekahi mau fungicides a me nā antibiotic ke hoʻohuli i ke alapine o ka mitotic recombination, kahi hiki ke hoʻololi i nā alapine genotype i nā lehulehu. Hoʻopaʻa ka benomyl fungicide i hoʻohana ʻia i ka beta-tubulin, kahi protein kahi e kūkulu ʻia ai nā microtubules o ka cytoskeleton, a ma laila e hoʻopilikia ai i nā kaʻina hana o ka hoʻokaʻawale chromosome i ka anaphase o mitosis, e hoʻonui ana i ke alapine o ka mitotic recombination (Hastie, 1970).
ʻO ka fungicide para-fluorophenylalanine, hoʻohana ʻia e mālama i ka maʻi Dutch i nā elms, he waiwai like kāna. Ua hoʻonui ka Para-fluorophenylalanine i ke alapine o ka recombination ma heterozygous diploids P. infestans (Poedinok et al., 1982).
Hoʻololi nā Cyclic i ka genotypic hoʻohui o nā heluna i ke ola o P. infestans
ʻO ke kaʻina hoʻomohala moʻokalaleo o P. infestans i ke kaona ʻoiaʻiʻo he 4 mau hanana.
1) pae o ka ulu exponential o ka heluna kanaka (pae polycyclic) me nā hanauna pōkole. Hoʻomaka pinepine kēia pae i Iulai a mau ʻo 1,5-2 mau mahina.
2) Ke kaʻina o ka hoʻokū ʻana i ka ulu o ka heluna kānaka ma muli o ka hōʻemi nui ʻana o ka mahele o nā mea i hoʻopili ʻole ʻia a i ʻole ka hoʻomaka ʻana o nā kūlana maikaʻi ʻole o ke aniau. ʻO kēia pae i nā mahiʻai e lawe i ka wehe mua ʻana o ka lau i ka lau i waho o ka pōʻai makahiki.
3) Ke kaʻina o ka hoʻoilo i loko o nā tubers, me ka hoʻoliʻiliʻi nui o ka nui o ka heluna kanaka ma muli o ka maʻi pōpilikia o nā tubers, ka ulu lohi o ka maʻi i loko o lākou, ka loaʻa ʻole o ka maʻi hou o nā tubers, ka pala a me ka culling o nā tubers i pili ma lalo o nā kūlana mālama maʻamau.
4) Ke kaʻina o ka lohi o ka hoʻomohala ʻana i ka lepo a me nā mea kanu (pae monocyclic), kahi e hiki ai i ka lōʻihi o ka hanauna i hoʻokahi mahina a ʻoi paha (hopena o Mei - Iulai mua). ʻO ka maʻamau i kēia manawa, paʻakikī ka ʻike ʻana i nā lau maʻi, ʻoiai me nā nānā kūikawā.
ʻO ka pae o ka ulu lehulehu exponential (pae polycyclic)
Nā hōʻike he nui (Pshedetskaya, Kozubova, 1969; Borisenok, 1969; Osh, 1969; Dyakov, Suprun, 1984; Rybakova, Dyakov, 1990) i ka hoʻomaka ʻana o ka epiphytoty, nā pali hoʻohaʻahaʻa haʻahaʻa a me nā mea hōʻeuʻeu iki, a laila pani ʻia e nā mea ʻino a me nā mea hakakā. ʻo ka helu o ka ulu ʻana o ka hakakā o ka lehulehu ka mea kiʻekiʻe, ʻoi aku ka liʻiliʻi o ke kūpaʻa o nā ʻano mea kanu o ka mea hoʻokipa.
I ka ulu ʻana o ka heluna kanaka, piʻi ka nui o nā genes selectively important i loko o nā ʻano pāʻoihana (R1-R4) a me ke kūlike ʻole (R5-R11). No laila, i nā lehulehu kokoke i Moscow i ka makahiki 1993, ua hoʻonui ʻia ka virulence awelika mai ka hopena o Iulai a i waena o ʻAukake mai 8,2 a i 9,4, a ua ʻike ʻia ka hoʻonui nui loa no ka gen Rul virulence i wae ʻia (mai 5 a 31% o nā clone virulent) (Smirnov, 86 ).
Hele pū ka hōʻemi ʻana o ka helu ulu o kahi heluna kanaka me ka hoʻoliʻiliʻi o ka hana parasite a ka heluna kanaka. No laila, i nā makahiki depressive, ʻo ka helu nui o nā lāhui a me ka mahele o nā lāhui virulent haʻahaʻa ma lalo o nā epiphytotic (Borisenok, 1969). Inā ma ke kiʻekiʻe o ke ʻano o ka manawa epiphytotic e loli i ka maikaʻi ʻole no ka lohi o ka hōʻino ʻana o ka uala a me ka hoʻohaʻahaʻa ʻana o ka uala, e hoʻoliʻiliʻi hoʻi ka nui o nā clone virulent a me ka hakakā (Rybakova et al., 1987).
ʻO ka piʻi ʻana o nā alapine o nā gen e pili ana i ka virulence a me ka hakakā o ka heluna kanaka ma muli o ke koho ʻana o nā clone virulent a me ka hakakā i ka heluna lehulehu. E hōʻike i ke koho ʻana, ua hoʻomohala ʻia kahi hana no ke kālailai ʻana i nā hoʻololi kūlike ʻole, i hoʻohana maikaʻi ʻia i nā heluna cheastat o ka hū (Adams et al., 1985) a me Fusarium graminearum (Wiebe et al., 1995).
ʻO ke alapine o nā mutants kūpaʻa i ka blasticidin S i ka heluna nui o P. infestans i hoʻoliʻiliʻi like me ka ulu ʻana o ka agresive o ka heluna kanaka, kahi e hōʻike ai i ka loli o nā clone mana i ka wā o ka ulu ʻana o ka heluna kanaka (Rybakova et al., 1987).
ʻO ka hoʻoilo i ka tubers
I ka hoʻoilo i loko o nā kūmaka potato, hoʻemi ka virulence a me ka hakakā o P. infestans, a ʻoi aku ka lohi o ka hōʻino ma mua o ka hoʻokūkū (Rybakova a me Dyakov, 1990). Kohu mea lā, ma lalo o nā kūlana kūpono i ka ulu wikiwiki ʻana o ka heluna kanaka (r-select), pono ke kōkua ʻia nā genes virulence "extra", no laila ke hele pū ʻia nei ka epiphytotics me ke koho ʻia ʻana o nā clone virulent a me ka hakakā. I ke kūlana saturation o ke kaiapuni, inā ʻaʻole ka helu o ka hānau hou ʻana, akā ʻo ka hoʻomau ʻana o ka noho ʻana i nā kūlana maikaʻi ʻole (K-koho) he kuleana nui, "nā keu" genes o ka virulence a me ka hakakā e hōʻemi i ke olakino, a ʻo nā klona me kēia mau ʻāpana ka mea mua e make, i mea e make ai ka awelika awelika a ke hāʻule nei ka virulence o ka lehulehu.
ʻO ka lau mea kanu i ka lepo
ʻO kēia pae ka mea pohihihi loa i ke ola ola (Andrivon, 1995). Hoʻolaha wale ʻia kona ola ʻana - ma muli o ka nele o ka ʻike e pili ana i ka hopena o ka maʻi pathogen ma kahi lōʻihi (i kekahi manawa ʻoi aku ma mua o hoʻokahi mahina) - mai ka puka ʻana o nā lau ʻuala i ke ʻano o nā wahi mua o ka maʻi ma luna o lākou. Ma ke kumu o nā nānā a me nā hoʻokolohua, ua kūkulu hou ʻia ke ʻano o ka fungus i kēia wā o ke ola (Hirst and Stedman, 1960; Boguslavskaya, Filippov, 1976).
Hiki ke hana i ka sporulation o ka fungus i nā tubers maʻi i loko o ka lepo. Kūkulu nā spores hopena me ka hyphae, hiki ke hoʻoulu no ka manawa lōʻihi i ka lepo. ʻO ke kula kiʻekiʻe (hoʻokumu ʻia ma nā tubers) a me ke kula kiʻekiʻe (ma ka mycelium i loko o ka lepo) piʻi nā ʻūlū i ka papa lepo e nā kahe capillary, akā loaʻa ka hiki ke hoʻopili i nā ʻuala ma hope wale o ka iho ʻana o kāna lau lalo a pili i ka papa o ka lepo. ʻO ia mau lau (ʻo ia hoʻi, nā kiko mua o ka maʻi i loaʻa ma luna o lākou) ʻaʻole kū koke, akā ma hope o ka ulu lōʻihi ʻana a me ka hoʻomohala ʻana o nā piko potato.
No laila, i ka pōʻaiapuni o P. infestans, hiki ke noho i ka pae kanu saprotrophic. Inā i ka ʻāpana parasitika o ka pōʻaiapuni ke ola nui ka mea nui o ke olakino, a laila ma ka ʻāpana saprotrophic e koho ʻia e hoʻoliʻiliʻi i nā waiwai parasite, e like me ka hoʻokolohua i hōʻike ʻia no kekahi fungi fitopathogenic (e ʻike iā Carson, 1993). No laila, i kēia kaʻina o ke pōʻaiapuni, pono e nalo ikaika loa ʻia nā waiwai hakakā. Akā i kēia manawa ʻaʻole i lawe ʻia nā hoʻokolohua pololei e hōʻoia i nā manaʻo i luna.
Hoʻololi nā hoʻololi o ka wā i nā waiwai pathogenic wale nō o P. infestans, akā i ke kūʻē ʻana i nā fungicides, e ulu nei i ka polycyclic phase (i ka wā epiphytoties), a e emi i ka wā hoʻoilo (Derevyagina et al., 1991; Kadish and Cohen, 1992). Ua ʻike ʻia kahi hāʻule ikaika i ke kūʻē ʻana i ka metalaxyl ma waena o ke kanu ʻana i nā tubers i hoʻopili ʻia a me ke ʻano o nā wahi mua o ka maʻi i ke kula.
Hana kūikawā Intraspecific a me kona ulu ʻana
Ke hoʻokumu nei ʻo P. infestans i nā maʻi ʻino i nā mea kanu waiwai ʻelua, ʻo kaʻuala a me kōmato. Hoʻomaka koke nā Epiphytoties ma kaʻuala ma hope koke iho o ka komo ʻana o ka fungus i nā wahi hou. Ua ʻike ʻia ka lilo ʻana o ke kōmato ma hope koke iho o ka ʻike ʻia ʻana o ka maʻi ma kaʻuala, akā ua ʻike ʻia nā epiphytoties ma ke kōmato hoʻokahi haneli mau makahiki wale nō i hala - i ka waena o ke kenekulia XNUMX. Eia ka mea a Hallegli lāua ʻo Niederhauser i kākau ai e pili ana i ka pio ʻana o nā tōmato ma USA
(1962): "Ma kahi o 100 mau makahiki ma hope o ka epiphytoty koʻikoʻi o 1845, kakaikahi a ʻaneʻane paha ʻaʻole i hoʻāʻo ʻia e kiʻi i nā ʻano kūpale kūpato. ʻOiai ua kākau mua ʻia ka hopena maikaʻi loa i nā ʻōmato ma mua o 1848, ʻaʻole ia i lilo i mea e nānā nui ai nā mea hānai ma kēia mea kanu a hiki i ka wā o ka maʻi nui i ka makahiki 1946. Ma nā panalāʻau o Rūsia i hoʻopaʻa ʻia i ka 60 kenekulia. "No kekahi manawa lōʻihi, ʻaʻole nānā ka poʻe noiʻi i kēia maʻi, ʻoiai ʻaʻole ia he kumu no ka hōʻino waiwai nui. Akā i nā makahiki 70 a me 1979. ʻIke ʻia hoʻi nā epiphytoties XX kenekulia o ka hopena o ka maʻi i loko o ka Soviet Union, ma ka hapa nui o ka Lower Volga, ma Ukraine, North Caucasus, ma Moldova ... "(Balashova, XNUMX).
Mai ia manawa, ua lilo ka maʻi kōmato e ka hopena maikaʻi loa i makahiki, ua pālahalaha ʻia ma nā ʻāina āpau o ka mahi ʻana i nā ʻoihana a me nā home a kumu i ka hōʻino waiwai nui i kēia hua. He aha i hana ai? No ke aha i ʻike ʻia ai ke ʻano mua o ka parasite ma kaʻuala a me ka liona epiphytotic o kēia hua i like i ka manawa like, a no ke aha i hala kahi kenekulia no ka epiphytotic e ʻike ʻia ma ka tumato? Kākoʻo kēia mau ʻokoʻa i kahi Mekiko ma mua o kahi kumu maʻi maʻi ʻAmelika Hema. Inā ua hoʻokumu ʻia nā ʻano Phytophthora infestans ma ke ʻano he parasite o nā lāhui tuber Mekiko o ka genus Solanum, a laila he mea maʻalahi ke kumu o ka mahi ʻana i kaʻuala e pili ana i ka ʻāpana like o ka genus e like me ka mea i laha nui ʻia nā ʻano Mekiko, akā ma muli o ka loaʻa ʻole o ka co-evolution me ka parasite, ka mea i hoʻomohala ʻole i nā ʻano kūʻē a kūʻē ʻole nonspecific.
ʻO Tomato kahi ʻāpana ʻokoʻa o ka genus, ke ʻano o kāna kūkaʻi he ʻokoʻa nui kona mai nā ʻano tuberous, no laila, ʻoiai ʻo ka ʻoiaʻiʻo ʻaʻole ʻo ka kōmato ma waho o ka meaʻai i hoʻomākaukau ʻia o P. infestans, ʻaʻole lawa ka ikaika o kona pohō no nā pohō waiwai nui.
ʻO ka puka ʻana o epiphytoties ma ka ʻōmato ma muli o nā loli koʻikoʻi maoli i ka parasite, kahi i hoʻonui ai i ka hiki ke hoʻololi ʻia (pathogenicity) ke paras parasised. Manaʻo mākou ʻo ka palapala hou i hana kūikawā ʻia no ka parasitizing i ka ʻōmato ka lāhui T1 i wehewehe ʻia e M. Gallegly, e loli ana i nā ʻanoʻano o ke kōmato cherry (Red Cherry, Ottawa), kūpaʻa i ka heihei T0 i hoʻolaha nui ʻia ma kaʻuala (Gallegly, 1952). Kohu mea lā, kahi hoʻololi (a i ʻole kahi moʻo o nā hoʻololi) i hoʻohuli i ka heihei T0 i ka heihei T1 a alakaʻi i ke ʻano o nā clones i hoʻomaʻamaʻa nui ʻia i ka eo ʻana o ka ʻōmato. E like me ka pinepine, ʻo ka hoʻonui ʻana o ka pathogenicity i kekahi mea hoʻokipa i ukali ʻia me ka hoʻoliʻiliʻi ʻana i kahi ʻē aʻe, ʻo ia hoʻi, kahi hoʻomaka mua, ʻaʻole naʻe i kū i ka manawa kūikawā intraspecific - i nā uala (heihei T0) a i ke kōmato (heihei T1).
He aha nā hōʻike no kēia kuhi?
- Ke hanana ma kaʻuala a me nā kōmato. Ma nā lau kōmato, lanakila ka heihei T1, ʻoiai ma nā lau ʻuala he laha ʻole ia. Wahi a S.F.Birirova a me T.A. ʻO Oreshonkova (ʻaʻole i paʻi ʻia) ma ka moku ʻo Moscow ma 1991-1992, ʻo ka hanana o ka heihei T1 ma nā mea kanu uala ʻo 0%, a i nā kanu kōmato - 100%; i ka 1993-1995 - 33% a me 90%, i kēlā me kēia; i 2001 - 0% a me 67%. Loaʻa nā ʻikepili like i loko o Israel (Cohen, 2002). ʻO nā hoʻokolohua me ka maʻi o nā kūmaka ʻuala me nā kaʻawale o ka heihei T1 a me ka huikau o nā kaʻawale T0 a me T1 i hōʻike ʻia e mālama maikaʻi ʻole ʻia nā kaʻawale o nā lāhui T1 i nā tubers a ua hoʻololi ʻia e nā kaʻawale o ka lāhui T0 (Dyakov et al., 1975; Rybakova, 1988).
2) Nā Dynamics o ka heihei T1 i nā mea kanu kōmato. Lawe ʻia ka maʻi nui o nā lau kōmato e nā kaʻawale o ka lāhui T0, ka mea e hoʻomalu i ka ʻimi ʻana o ka maʻi i nā kiko mua i hana ʻia i nā lau. Hōʻoia kēia i ka papahana i ʻae ʻia o ka neʻe parasite: ʻo ka nui nui o ka maʻi mai ka ʻuala i hana ʻia e ka lāhui T0, akā naʻe, he hapa liʻiliʻi o nā clone T1 i mālama ʻia i loko o kaʻuala, hoʻokahi manawa ma ke kōmato, hoʻoneʻe i ka heihei T0 a hōʻiliʻili i ka hopena o ka wā epiphytotic. Hiki nō paha ke loaʻa kahi kumu ʻē aʻe o ka maʻi ʻomaʻoma lau me ka lāhui T1, ʻaʻole ikaika e like me ka ʻuala a me nā lau, akā mau. No laila, he hopena nāwaliwali kēia kumu i ka hanana genetika o ka heluna kanaka e hoʻopili ana i ka ʻōmato, akā hoʻoholo hope i ka hōʻiliʻili o ka lāhui T1 (Rybakova, 1988; Dyakov et al., 1994).
3) Hoʻohui i ka uala a me nā kōmato. ʻO ka maʻi artipisyal o ka lau ʻōmato a me kaʻuala me nā kaʻawale o nā lāhui ʻo T0 a me T1 i hōʻike ʻoi aku ka hakakā o ka mea mua no kaʻuala ma mua o ka ʻōmato, a ʻoi aku ka ikaika o ka hope no ka ʻōmato ma mua o kaʻuala. Hōʻike ʻia kēia mau ʻokoʻa i ka neʻe ʻana o nā kaʻawale o kahi lāhui ʻona ponoʻī ʻole mai kahi lāhui huikau i ka manawa o nā ala lau i loko o kahi hale kōmaʻomaʻo (D'yakov et al., 1975) a me nā pā kula (Leberton et al., 1999); nā ʻokoʻa i ka ukali infectious liʻiliʻi, ka manawa latency, ka nui o nā wahi infectious a me ka hana 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).
ʻO ka huhū o nā kaʻawale o ka heihei T1 i nā ʻano kōmato i nele i nā genes kūʻē ke kiʻekiʻe loa a hoʻoneʻe ʻia kēia mau mea hoʻokaʻawale i nā lau e like me ka mea momona me ka ʻole o ka necrotizing o ka maʻi i loaʻa i ka maʻi (Dyakov et al., 1975; Vega-Sanchez et al., 2000).
4) Virulence no kaʻuala a me nā kōmato. Hoʻopili ka lāhui T1 i nā ʻano kōmato cherry me ka ʻaoʻao kūʻē Ph1, ʻoiai ʻaʻole hiki i ka lāhui T0 ke hoʻopili i kēia mau ʻano, ʻē. he palupalu ka virulence. E pili ana i nā mea ʻokoʻa
Pili ʻokoʻa nā lāina R o nā uala, ʻo ia hoʻi ʻoi aku ka maikaʻi ʻole o nā kaha i hoʻokaʻawale ʻia mai nā lau kōmato ma mua o nā kānana "ʻuala" (Papa 11).
5) Nā hōʻailona kūleʻa. ʻO ke kālailai ʻana o nā māka kūpaʻa i nā heluna kanaka o P. infestans parasitizing ma kaʻuala a me nā tōmato hoʻi e hōʻike nei i ke koho intraspecific multidirectional. I ka heluna lehulehu o P. infestans o Brazil, no ka laina clonal US-1 nā kaʻawale kaʻanalā, a ʻo nā mea mai nā lau ʻuala no ka laina BR-1 (Suassuna et al., 2004). Ma Florida (USA), mai ka 1994, ua hoʻomaka ka clone US-90 e hoʻomake ma luna o kaʻuala (me kahi hanana i ʻoi aku ma mua o 8%), a me nā clone US-11 a me US-17 ma ke kōmato, a ʻoi aku ka hakakā o nā mea kaʻawale no ka ʻōmato ma mua o kaʻuala (Weingartner , Tombolato, 2004). Ua hoʻokumu ʻia nā ʻokoʻa nui o nā alapine genotype (DNA fingerprints) ma ka ʻuala a me nā kaʻawale kōmato no 1200 P. infestans i hōʻiliʻili ʻia i United States mai 1989 a 1995 (Deahl et al., 1995).
ʻO ka hoʻohana ʻana i ka hana AFLP ua hiki ke hoʻokaʻawale iā 74 mau kaha i ʻohiʻohi ʻia mai kaʻuala a me nā lau kōmato i ka makahiki 1996-1997. ma Palani a me Kuikilana, i loko o 7 mau pūʻulu. ʻAʻole i ʻokoʻa ke ʻano o kaʻuala a me nā ʻano kōmato, akā ʻo nā ʻano "ʻuala" i ʻoi aku ka nui o ka genetically ma mua o nā "tōmato". Ua loaʻa nā mea mua i nā pūpū ʻehiku, a ʻo ka hope, i loko o nā ʻehā wale nō, kahi e hōʻike ai i kahi genome loea o ka hope (Knapova and Gisi, 2002).
6) Nā ʻano hana o ke kaʻawale. Inā hoʻololi ka lehulehu o ka parasite i nā ʻano mea kanu ʻelua i ka hoʻopili ʻana o ka hana pilikino i kā lākou "host" ponoʻī, a laila ala aʻe nā ʻano hana mua a me nā postmeiotic e pale ai i nā hoʻololi kūpuna interpopulation (Dyakov a me Lekomtseva, 1984).
Ua ʻimi nā noiʻi he nui i ka hopena o ke kumu o nā ʻano makua ma ka pono o ka hybridization. Ke hoʻokaʻawale ʻia nā kaha o kēlā me kēia ʻano o ka genus Solanum ma Ecuador (Oliva et al., 2002), ua ʻike ʻia ʻo nā kaha me ka moʻo ʻano A2 mai ka Solanaceae hihiu (laina clonal EC-2) ka mea ʻoi loa i hele aʻe me nā ʻōpū mai ke kōmato (laina EC -3), a kea ʻoi aku ka maikaʻi me ke kānana ʻuala (EC-1).
Loaʻa nā hybrids āpau i non-pathogenic. Manaʻo nā mea kākau i ka pākēneka haʻahaʻa o ka hybridization a me ka hōʻemi ʻana o ka pathogenicity i nā hybrids ma muli o nā ʻano postmeiotic o ka hoʻokaʻawale hānau ʻana o nā lehulehu.
I nā hoʻokolohua o Bagirova et al. (1998), ua hele a nui ka nui o nā ʻuala a me nā ʻano kōmato me nā waiwai o nā heihei T0 a me T1. ʻO nā momona loa nā keʻa o T1xT1 mau ʻano i hoʻokaʻawale ʻia mai ka ʻōmato (36 oospores i ka māla microscope o ka nānā, 44% o ka ulu ʻana o ka oospore), ʻo ka mea liʻiliʻi loa ke keʻa o nā lāhui T0xT1 i hoʻokaʻawale ʻia mai nā pūʻali ʻokoʻa (kahi helu haʻahaʻa o nā oospores ulu a germination, kahi mahele kiʻekiʻe o nā oospores abortive a underdeveloped) ... ʻO ka maikaʻi o nā keʻa i waena o nā kaʻawale o ka lāhui T0 i hoʻokaʻawale ʻia mai kaʻuala i waena. Ma muli o ke kino nui o nā ʻano o ka heihei T0 e pili ana i kaʻuala, aia kahi kumu hilinaʻi o ka hoʻoilo - tubers potato, ma muli o ka mea nui o nā oospores e like me nā hoʻoiho infectious wintering no nā heluna mai kaʻuala. Hiki i ke "ʻano kōmato" i hoʻopili ʻia i ka hoʻoilo ma ke kōmato ma ke ʻano o nā oospores (e ʻike ma lalo) a no laila e paʻa ana i kahi huahana kiʻekiʻe aʻe o ka hana moekolohe. Ma muli o kona momona kiʻekiʻe, loaʻa iā T1 kahi mana kūʻokoʻa no ka maʻi mua i ka ʻōmato. ʻO nā hopena i loaʻa iā Knapova et al. (Knapova et al., 2002) hiki ke unuhi ʻia i ke ala like. ʻO nā keʻa o nā ʻano kaʻawale i hoʻokaʻawale ʻia mai kaʻuala me nā kānana mai ke kōmato i hāʻawi i ka helu kiʻekiʻe o nā oospores - 13,8 no ka sq. Mm. waena (me ka pālahalaha o 5-19) a me kahi pākēneka waena o ka germination o oospores (6,3 me ka hoʻolaha ʻana o 0-24). ʻO nā keʻa o nā kaha i hoʻokaʻawale ʻia mai ka kōmato i hāʻawi i ka pākēneka haʻahaʻa o nā oospores (7,6 me ka pālahalaha o 4-12) me ka pākēneka kiʻekiʻe loa o kā lākou germination (10,8). ʻO nā keʻa i waena o nā kaha i hoʻokaʻawale ʻia mai kaʻuala i hāʻawi i kahi helu waena o nā oospores (8,6 me kahi kiʻekiʻe o ka ʻikepili - 0-30) a me ka hapa haʻahaʻa o ka germination o oospores (2,7). No laila, ʻoi aku ka momona o nā kānana mai kaʻuala ma mua o nā kōmato, akā ʻo nā keʻa interpopulation ʻaʻole i hāʻawi i nā hopena maikaʻi ʻole ma mua o nā intrapopulation. Hiki i nā ʻokoʻa me nā ʻikepili ma luna aʻe na Bagirova et al. ua wehewehe ʻia e ka ʻoiaʻiʻo o ka hana ʻana o nā mea noiʻi Lūkini me nā ʻano i hoʻokaʻawale ʻia i ka hoʻomaka ʻana o 90 o ke kenekulia iwakālua, a me nā mea noiʻi mai Switzerland - me nā maʻi i hoʻokaʻawale ʻia i ka hopena o 90.
ʻO ke kumu no ka momona haʻahaʻa ka heteroploidy o nā ʻano. Inā ma nā lehulehu Mekiko, kahi e maʻamau ai ka moekolohe a me ka maʻi mua me nā keiki oosporous, ʻo ka hapa nui o nā ʻano aʻo i aʻo ʻia o P. Infestans e diploid, a laila ma nā ʻāina o ka Old World intrapopulation polymorphism o ploidy e mālama ʻia (di-, tri- a me tetraploid mau ʻano, a me nā heterokaryotic a me nā heteroploid nuclei) , a me nā kaha i loaʻa nā ʻano moena like ʻole, ʻo ia hoʻi momona momona, ʻokoʻa i nā ploidy nukelea (Therrien et al., 1989, 1990; Whittaker et al., 1992; Ritch, Daggett, 1995). ʻO ka loli o nā mea nui i ka antheridia a me ka oogonia ke kumu o ka momona haʻahaʻa.
E pili ana i nā hoʻololi nuklea ma waena o ka hyphae i ka wā o anastomoses, pale ʻia kēia e ka incompatibility vegetative, ka mea e wāwahi i nā poʻe asexual i loko o nā clone hoʻokaʻawale genetically (Poedinok a me Dyakov, 1987; Gorbunova et al., 1989; Anikina et al., 1997b).
7) Hoʻohui o nā lehulehu. Hōʻike ka ʻikepili i luna aʻe hiki i ka hybridization ma waena o "potato" a me "Tomato" P. infestans ke hiki. Hiki nō hoʻi ke loaʻa hou i ka maʻi Reciprocal o nā pūʻali like ʻole, ʻoiai me ka hōʻemi o ka hakakā.
Ua hōʻike ʻia kahi noiʻi ʻana i nā māka lehulehu ma nā wahi kaʻawale mai nā ʻuala pili a me nā māla kōmaka i ka makahiki 1993 e pili ana i ka hapahā o nā kaʻawale i hoʻokaʻawale ʻia mai nā lau kōmato mai kahi māla ʻuala e pili kokoke ana (Dolgova et al., 1997). Ma ke ʻano kumuhana, hiki ke manaʻo ʻia e piʻi ka divergence o nā lehulehu ma luna o nā pūʻali ʻelua a alakaʻi i ka puka ʻana o nā ʻano intraspecific kūikawā (f.sp. potato and f.sp. Tomato), ʻoiai ʻoiai hiki i nā oospores ke hoʻomau i nā ʻōpala o nā mea kanu (Drenth et al., 1995 ; Bagirova, Dyakov, 1998) a me nā hua kōmato (Rubin et al., 2001). No laila, loaʻa i nā ʻōmato kahi kumu o ka hoʻoulu hou ʻana o ka puna i kū kūʻē ʻole i nā kōpoki uala.
Eia naʻe, ʻokoʻa nā mea āpau. ʻO ka overwintering me nā oospores i ʻae i ka parasite e hōʻalo i ka pae haʻahaʻa loa o kāna pōʻaiapuni - ke kahua monositolo o nā mea kanu i ka lepo, kahi e hoʻemi ai nā waiwai parasite, i hoʻihoʻi hou ʻia i ka polycyclic phase i ke kauwela.
Pākaukau 11. Nā alapine o nā genul virulence i nā ʻano ʻokoʻa potato ma nā ʻano P. infestans
aina | Makahiki | Ka helu maʻamau o nā genul virulence i nā kānana | mea kākau | |
mai kaʻuala | mai kōmato | |||
France | 1995 | 4.4 | 3.3 | Leberton et al., 1999 |
1996 | 4.8 | 3.6 | Leberton, Andrivon, 1998 | |
Palani, Kuikilana | 1996-97 | 6.8 | 2.9 | ʻO Knapova, Gisi, 2002 |
United States | 1989-94 | 5 | 4.8 | Goodwin et al., 1995 |
USA, Zap. Kālāʻo DC DC | 1996 | 4.6 | 5 | Dorrance et al., 1999 |
1997 | 6.3 | 3.5 | " | |
Ecuador | 1993-95 | 7.1 | 1.3 | Oyarzun et al., 1998 |
Israel | 1998 | 7 | 4.8 | Cohen, 2002 |
1999 | 6 | 5.7 | " | |
2000 | 6.7 | 6.1 | " | |
Lūkia, Mosk. ʻāpana | 1993 | 8.9 | 6.7 | Smirnov, 1996 |
Lūkia, nā ʻāina ʻokoʻa | 1995 | 9.4 | 8 | Kozlovskaya a me kekahi poʻe ʻē aʻe. |
1997 | 9.2 | 9.2 | " | |
2000 | 8.7 | 4.8 | " |
ʻO ka zoosporangia kumu a me nā zoospores, ka mea e ulu i nā oospores, he pae kiʻekiʻe o ka hana parasite, keu hoʻi inā i hoʻokumu ʻia nā oospores ma lalo o ka mana o nā pheromones o kahi kānana me ka ʻano ʻē aʻe o ka male ʻana. No laila, ʻo nā mea infectious ma luna o nā pua kōmato i ulu aʻe mai nā ʻanoʻano i hoʻopili ʻia me nā oospores he pathogenic nui ia no nā ʻōmato a me kaʻuala.
Ua alakaʻi kēia mau hoʻololi i kahi hoʻoponopono hou ʻana o ka heluna kanaka, i hōʻike ʻia i loko o nā loli koʻikoʻi e pili ana mai kahi manaʻo epidemiological:
- Ua lilo nā kumulāʻau kōmato maʻi i kumu waiwai nui no ka maʻi o kaʻuala (Filippov, Ivanyuk, nā leka pilikino).
- Hoʻomaka e ʻike ʻia nā Epiphytoties ma kaʻuala ma mua o Iune, ma kahi o hoʻokahi mahina ma mua o ka maʻamau.
- I nā kanu uala, ua hoʻonui ka pākēneka o ka heihei T1, i kū mua ʻia ma laila ma kahi nui kūpono ʻole (Ulanova et al., 2003).
- Hoʻokaʻawale ʻia nā ʻano kaʻawale mai nā lau kōmato mai nā ʻano ʻuala i ka virulence ma nā ʻokoʻa potato o nā genes virulence a hoʻomaka i ka ʻoi aku i nā ʻano "potato" i loko o ka hoʻokūkū ʻaʻole wale ma ke kōmato, akā ma nā uala hoʻi (Lavrova et al., 2003; Ulanova et al. , 2003).
No laila, ma kahi o ka divergence, aia kahi o ka lehulehu o ka lehulehu, ka puka ʻana o kahi heluna kanaka hoʻokahi ma luna o nā mea kanu ʻelua me ka nui o ka virulence a me ka hakakā i nā ʻano ʻelua.
hopena
No laila, ʻoiai ʻoi aku ma mua o 150 mau makahiki o ke aʻo ikaika ʻana o P. infestans, i ka biology, me ka biology heluna kanaka o kēia causative agena o nā maʻi nui loa o nā mea kanu solanaceous i mahi ʻia, nui ka ʻike ʻole ʻia. ʻAʻole maopopo i ke ʻano o ka hele ʻana o kēlā me kēia ʻāpana o ke ola pōʻaiapili e pili ana i ka hanana o nā heluna kanaka, he aha nā ʻano hanauna o ka hiki ke hoʻololi ʻia o ka hakakā a me ka virulence, he aha ka lakio o nā ʻōnaehana reproductive a me clonal i nā taupuna maoli, pehea ka hoʻoilina o ka vegetative, he aha ke kuleana o kaʻuala a me nā tōmato i ka maʻi nui o kēia mau mea kanu a i he aha ka hopena i ka hanana o nā heluna parasite. I kēia manawa, ʻo nā pilikia kūpono e like me nā ʻano hana kūpuna no ka hoʻololi ʻana i ka hakakā o ka parasite a i ʻole ke erosion o ka kūpaʻa potato nonspecific ʻaʻole i hoʻonā ʻia. Me ka hohonu a me ka hoʻonui ʻana o ka noiʻi ʻana ma ka uala i ka lohi o ka maikaʻi, hāʻawi ka parasite i nā mea hou i nā mea noiʻi. Eia nō naʻe, ka hoʻomaikaʻi ʻana o ka hiki ke hoʻokolohua, ke kū ʻana o nā ala hana hou e pili ana i ka manipula me nā gen a me nā protein e ʻae iā mākou e lana i kahi hopena kūleʻa o nā nīnau i hāpai ʻia.
Ua paʻi ʻia kēia ʻatikala ma ka puke pai "Protection Potato" (No. 3, 2017)