S.N. Elansky, L.Yu. Kokaeva, N.V. Statsyuk, Yu.T. Dyakov
Horudhac
Oomycete Phytophthora infestans (Mont.) De Bary - oo ah wakiilka sababa duqeynta goor dambe, cudurka dhaqaale ahaan ugu muhiimsan ee baradhada iyo yaanyada - ayaa soo jiitay indho dhow cilmi baarayaasha dalal kala duwan in ka badan qarni iyo badh. Si lama filaan ah ayay uga soo muuqatay Yurub badhtamihii qarnigii XNUMXaad, waxay sababtay cudur baradho ah oo ku sii jiray xusuusta jiilal badan.
Illaa iyo hadda, waxaa badanaa loogu yeeraa "boqosha gaajada Irishka". Ku dhowaad boqol sano ka dib cudurradii ugu horreeyay, noocyada baradhada duurjoogta ah ee u adkaysta dillaaca goor dambe ayaa la ogaaday, habab looga gudbo baradhada la beeray ayaa la sameeyay (Muller, 1935), iyo noocyadii ugu horreeyay ee u adkaysata u-adkaysiga waa la helay (Pushkarev, 1937). Si kastaba ha noqotee, wax yar ka dib bilowgii ganacsigoodii ganacsi, jinsiyadaha cudurkii halista ahaa ee daba-dheeraaday ee u nugul noocyada iska caabbinta ayaa ururay. iyo soo bandhigida hiddo-wadahayaal cusub oo iska caabin ah oo ka soo baxa baradhada duurjoogta ah ee Mexico noocyada kala duwan waxay bilaabeen inay si dhaqso leh u lumiyaan waxtarkoodii
Ku guuldareysiga adeegsiga monogenic (toosan) caabbinta qasab kalkaaliyeyaasha si ay u raadiyaan siyaabo aad u adag oo looga faa'iideysto caabbinta polygenic aan caadiga ahayn (jiifka). Sanadihii la soo dhaafay, jinsiyadaha dagaalka daran waxay bilaabeen inay ku soo ururaan shakhsiyaadka shakhsiyaadka dulinka ah, taasoo keeneysa nabaad guurka xitaa iska caabin aan qeexneyn. Soo bixitaanka noocyada u adkaysta fungicide-ka ayaa dhibaatooyin ku keenay isticmaalka kiimikada ka hortagga baradhada.
Farqiga weyn ee udhaxeeya oomycetes iyo fungi ee halabuurka kiimikada, Ultrastructure, iyo metabolism, sunta fangaska, gaar ahaan kuwa nidaamka loo isticmaalo si dhirta looga ilaaliyo cudurada fangaska badan, waxtar uma leh oomycetes.
Sidaa darteed, difaaca kiimikada ee ka soo baxa duqeynta goor dambe, tiro badan (illaa 12 jeer xilli ciyaareedkiiba ama ka badan) ku buufinta diyaarinta xiriirka xiriir ballaaran oo ficil ballaaran ah ayaa la isticmaalay. Tallaabo kacaan ah ayaa ahayd adeegsiga phenylamides, kuwaas oo sun ku ah oomycetes kuna faafa si nidaamsan dhirta. Si kastaba ha noqotee, isticmaalkooda baahsan ayaa si dhakhso leh u horseeday isku-soo-ururinta noocyo adkaysi u leh dadka fungal ah (Davidse et al., 1981), taas oo si weyn u murugsan ilaalinta dhirta. P. infestans ficil ahaan waa dulinka kaliya ee aagga cimilada, dhibaatada ka soo gaarto beerashada dabiiciga ah lama dhex dhexaadin karo iyadoo aan la adeegsan hab difaac kiimiko ah (Van Bruggen, 1995).
Waxyaabaha kor ku xusan waxay sharraxayaan dareenka weyn ee ay cilmi-baarayaasha ka kala yimid dalal kala duwan ay siinayaan daraasadda P. dadka ku dhex nool, firfircoonida badan ee ay leeyihiin iyo isku-dhafka hidde-wadaha, iyo sidoo kale hababka hidda-socodka ee kala-duwanaanta.
Wareegga nolosha ee R. INFESTANS
Oomycete Phytophthora infestans waxay soo saarta is-dhex-dhexaadiyaha 'mycelium' oo leh haustoria gudaha caleemaha baradhada. Ku quudinta unugyada caleenta, waxay sababtaa samaysma dhibco madow, oo isu beddela madow oo ku qudhmaa cimilada qoyan. Guuldarro xoog leh, caleen dhan ayaa dhimata. Muddo quudinta ka dib, soo bixitaanno ayaa ka soo baxa 'mycelium - sporangiophores' - oo ka baxa dibedda illaa stomata. Xilliga cimilada qoyan, waxay sameeyaan ubax cad oo ku wareegsan dhibcaha caleemaha hoostooda. Dhammaadka sporangiophores, zoosporangia oo u eg liin ayaa la sameeyay, kuwaas oo kala baxa oo ay qaadaan buufinta roobka (Sawirka 1). Ku soo dhaca dhibco biyo ah dusha sare ee caleen baradho ah, sporangia waxay ku dhalataa 6-8 zoospores, kuwaas oo, ka dib muddo dhaqdhaqaaq ah, la soo koobo, oo lagu daboolo qolof kuna soobaxay tuubada biqilka. Dhirta waxay soo dhex martaa stomata-ga illaa unugyada caleen. Xaaladaha qaarkood, sporangia waxay ku kori kartaa tuubada koritaanka si toos ah unug caleen ah. Xaaladaha wanaagsan awgood, waqtiga infakshanka illaa sameynta manida cusub waa 3-4 maalmood oo keliya.
Markaad dhulka joogtid oo aad sifeeysid ciidda, sporangia waxay awood uleeyihiin inay qaadaan cudurka digriiga. Digriiga sida aadka ah u saameeyay wuu xumaadaa inta lagu jiro keydinta; in si taban u saameysey, infekshanku wuu sii jiri karaa illaa xilliga soo socda Intaas waxaa sii dheer, wakiilka sababa duqeynta goor dambe wuxuu kusii jiri karaa xilliga jiilaalka qaab loo yaqaan 'oospores' (fallaaro galmo nasasho leh oo adag oo nasasho leh) oo ku dhex jira ciidda qashinka dhirta iyo miraha yaanyada. Oospores-ka waxaa lagu sameeyaa xubnaha nool ee dhirta marka noocyada noocyada kala duwan ee isku-dhafka la kulmaan qoyaan badan. Guga, xawada asexual-ka waxaa lagu sameeyaa digriiga la beeray iyo haraaga dhirta oo leh oospores; zoospores waxay soo galaan ciidda waxayna sababaan infekshinka caleemaha hoose ee dhirta. Xaaladaha qaarkood, mycelium ayaa ka soo bixi kara tuber-ka cudurka qaba ee ku dhaca qaybta cagaaran ee geedka waxayna badanaa ka muuqdaan qaybta sare ee asliga.
Farqi weyn oo udhaxeeya oomycetes iyo fangaska badankood wuxuu kujiraa badnaanta diplophase ee wareega noloshooda oo leh gamio meiosis iyo biqilka zygotes (oospores) iyadoon lakala goynin nukliyeerka. Tilmaamahan, oo lagu daray beddelka heterotallism ee beddelaya bisexuality, waxay umuuqataa inay suurtogal ka dhigeyso in lagu dabaqo oomycetes habab loo sameeyay barashada dadweynaha eukaryotes ka sareeya (falanqaynta panmixia iyo kala-qeybsanaanta dadweynaha, qulqulka hiddo-wadaha iyo qulqulka qulqulka, iwm.). Si kastaba ha noqotee, seddex arrimood ma oggola in gebi ahaanba lagu wareejiyo hababkan daraasadda P.
1. Ayadoo ay weheliso oospores isku jira, iskood isu dhalaalaya iyo oo loo yaqaan 'parthenogenetic oospores' ayaa laga sameeyay dadweynaha (Fife iyo Shaw, 1992; Anikina et al., 1997a; Savenkova, Cherepnikoba-Anirina, 2002; Smirnov, 2003), iyo inta jeer ee samaysankoodu ku filnaan karo inay saameyn ku yeeshaan natiijooyinka baaritaanka.
2. Nidaamka galmada ee P. infestans wuxuu waxtar aan qiimo lahayn ku yeelanayaa dhaqdhaqaaqa tirada dadka, maxaa yeelay fangaska wuxuu ku tarmaa inta badan dhirtu, isagoo sameysmaya in ka badan 90% natiijooyinka falanqaynta nooca isku-tagga iyadoo loo adeegsanayo habka dhaqameed ee nafaqada leh ... xilliga sii kordhaya ee jiilal dhowr ah oo xaamilo ah (horumarka cudurka polycyclic). Oospores waxay door muhiim ah ka ciyaaraan ilaalinta noolaha inta lagu jiro mudada aysan jirin dhir cagaaran (xilliga qaboobaha) iyo infekshanka asaasiga ah ee geedo. Kadib, inta lagu jiro xilliga kuleylaha, taranka clonal iyo kororka ama, liddi ku ah, hoos u dhac ku yimaada tirada isku-xirnaanta shakhsiyadeed ee ku soo kordhay natiijada galmada dib-isu-keenista ayaa dhacda, taas oo inta badan lagu go'aamiyo xulashada kuwa ugu habboon. Sidaa darteed, saamiga midabyada shakhsiga ee dadweynaha bilowga iyo dhammaadka epiphytotics way gabi ahaanba ka duwanaan karaan.
3. Wareegga la sharaxay wuxuu caan ku yahay dadka u dhashay P. ee ku nool dalkooda, Bartamaha Ameerika. Meelaha kale ee adduunka, habka galmada lama aqoon in ka badan 100 sano; mycelium daaqsinka ku jira digriiga baradhada qaba waa marxaladdii qaboobaha. Wareega noloshu wuxuu ahaa mid gebi ahaanba agamic ah, faafitaankuna wuxuu ahaa mid udub dhexaad u ah dabeecadda: infekshinka ka imaaday digriiga kaliya ee la beeray oo u gudbay caleemaha, sameysana ujeeddooyinka koowaad ee cudurka, kaas oo ku milmi kara horumarka ballaaran ee cudurka.
Sidaa darteed, gobollada qaarkood waxaa jiri kara isku-beddel wareegyo galmo iyo isu-tag ah, halka kuwa kalena - kaliya wareega wareega.
Asalka P. INFESTANS
P. infestans waxay ka soo muuqdeen Yurub dhammaadkii qeybtii hore ee qarnigii 1991aad. Maaddaama baradhadu ay u dhalatay waqooyiga-bari ee Koonfurta Ameerika, waxaa loo qaatay in dulinka laga keenay halkaas loo keenay Yurub inta lagu gudajiray kor u qaadista milixda cusbada ee Chile. Si kastaba ha noqotee, daraasado lagu sameeyay xarunta baradhada ee Rockefeller Center oo ku taal dooxada Toluca, Mexico ayaa ku qasabtay aragtidan in dib looga fiirsado (Niederhauser, 1993, XNUMX).
1. Dooxada Toluca, noocyada baradhada tuubbada ee deegaanka (Solanum demissum, S. bulbocastanum, iwm.) Waxay leeyihiin noocyo hiddo sidayaal ah oo iska caabin toosan ah oo ay weheliso heer sare oo iska caabin aan qeexnayn, taas oo muujineysa wada-noolaansho dheer oo dulin leh. Noocyada Koonfurta Ameerika, oo ay ku jiraan baradhada dalagga, ma laha hiddo-wadayaal iska caabin ah.
2. Dooxada Toluca, go'doominno leh noocyada isweheshiga ee A1 iyo A2 ayaa laga helaa, taas oo ay keentay in dadka ku dhex-dhexaadsan ee P. infestans ay ku baahsan yihiin; halka dhulkii hooyo ee baradhada la beeray, Koonfurta Ameerika, dulinku si wada jir ah ugu faafo.
3. Dooxada Toluca, waxaa jira cudurro faafa oo aad u daran oo sanad kasta dhaca. Sidaa darteed, ka mid ah cilmi baarayaasha Waqooyiga Ameerika (Jaamacadda Cornell), fikradda ku saabsan Mesoamerica (Bartamaha Ameerika) oo ah halka uu ku dhashay baradhada phytophthora ayaa la aasaasay (Goodwin et al., 1994).
Baarayaasha Koonfurta Ameerika ma wadaagaan fikraddan. Waxay rumeysan yihiin in baradhada la beeray iyo dulinimadiisa P. infestans ay leeyihiin dal ay wadaagaan - Koonfurta Ameerika. Waxay ku taageereen aragtidooda daraasad molikal ah oo ku saabsan falanqaynta DNA polymorphisms ee hiddo-wadaha mitochondrial (mtDNA) iyo hiddo-wadaha nukliyeerka RAS iyo β-tubulin (Gomez-Alpizar et al., 2007). Waxay muujiyeen in noocyada laga soo ururiyay meelo kala duwan oo adduunka ka mid ahi ay ka soo kala jeedaan saddex sadar oo awooweyaal kaladuwan oo saddexda ahba laga helo Koonfurta Ameerika. Noocyada kala duwan ee 'Andean haplotypes' waa farac ka kooban laba sadar: go'doominta ugu da'da weyn mtDNA waxaa laga helaa Solanaceae oo duurjoog ah oo ka soo jeeda qaybta Anarrhicomenum ee Ecuador, halka go'doominta khadka labaad ay ku badan yihiin baradhada, yaanyada iyo habeennada duurjoogta ah. Toluca, xitaa nuucyada noocyada dhifka ah waxay ka soo farcameen hal nasab oo keliya, iyadoo kala duwanaanshaha hidde-wadaha ee noocyada Toluca (hoos u dhaca hooseeya ee bogagga qaarkood ee isbeddelka leh) ay muujineyso saameyn aasaasihii aasaasiga ahaa sababo la xiriiray dhowaantan.
Intaa waxaa dheer, nooc cusub oo ah P. andina ayaa laga helay Andes, qaab ahaan iyo hidde ahaanna la mid ah P. infestans, taas oo, sida ay qorayaashu tilmaamayaan, Andes inay tahay meel kulul oo takhasus gaar ah u leh isir ahaanta Phytophthora. Ugu dambeyntiina, Yurub iyo Mareykanka, P. dadka infestans-ka ku nool waxaa ka mid ah labada abtirsiin ee Andean, halka Toluca kaliya ay tahay hal.
Daabacaaddan ayaa keentay jawaab ka timid koox cilmi-baarayaal ah oo ka kala yimid dalal kala duwan, kuwaas oo qabtay howlo badan oo tijaabo ah si ay dib ugu eegaan daraasaddii hore (Goss et al., 2014). Shaqadan, marka hore, taxane dheeri ah oo microsatelite DNA ah ayaa loo isticmaalay in lagu barto polymorphisms-ka DNA; marka labaad, falanqaynta isku-urursiga, waddooyinka socdaalka, waqtiga kala-duwanaanta dadka, iwm. moodooyin aad u horumarsan ayaa la adeegsaday (F-tirakoobka, qiyaasta Bayesian, iwm.) , laakiin sidoo kale leh noocyada dabiiciga ah ee reer Mexico P. mirabilis, P. Ipomoeae, iyo Phytophthora phaseoli - hiddo ahaan u dhow P. ciyaal isir isku mid ah (Kroon et al., 2012). Natiijooyinka ka soo baxa falanqeyntan, waxaa si aan mugdi ku jirin loo muujiyey in qaybta asalka u ah geedka phylogenetic ee dhammaan noocyada genus Phytophthora ee lagu qaatay daraasadda, marka laga reebo isugeynta P. andina, waxaa iska leh noocyada Mexico, socodka socdaalkuna wuxuu leeyahay jihada Mexico - Andes, mana ahan taa cakiskeeda, bilowgeeduna wuxuu la mid yahay Yurub gumaysiga Dunida Cusub (300-600 sano ka hor). Marka, soo bixitaanka noocyada P. infestans, oo ku takhasusay guuldarada baradhada, ayaa ka dhacay xarunta hidda-socodka labaad ee sameynta habeynnada tuubbada leh, i.e. Bartamaha Ameerika.
Genome ee P. INFESTANS
Sannadkii 2009, koox caalami ah oo saynisyahanno ah ayaa taxay dhammaystirka 'P infestans genome' (Haas et al, 2009), oo cabirkiisu ahaa 240 MB. Tani waxay dhowr jeer ka badan tahay noocyada dhow ee la xiriira P. sojae (95 Mb), oo sababa xididdada soybeanska, iyo P. Ramorum (65 Mb), oo saameynaya noocyada qiimaha leh ee geedka sida geed, beech, iyo qaar kale. Xogta la helay waxay muujisay in hiddo-wadaha uu ku jiro tiro badan oo nuqullo ah oo isdaba-joog ah - 74%. Hidda-wadaha wuxuu ka kooban yahay 17797 hiddo-wadayaal hiddo-wadayaal ah, badankooduna waa hiddo-wadayaal ku lug leh geedi socodka gacanta, oo ay ku jiraan soo-saaridda DNA, qorista iyo tarjumaadda borotiinnada.
Isbarbardhiga genomes ee genus Phytophthora wuxuu shaaca ka qaaday urur aan caadi aheyn oo hiddo-wadaha ah, oo ka kooban jajabyo taxane ah oo hiddo-wadayaal la ilaaliyay ah, kaas oo cufnaanta hiddo-wadaha ay aad u sarreyso isla markaana waxa ku jira taxanaha soo noqnoqda ay aad u hooseyso, iyo gobollada shaqsiyaadka leh taxadarka hiddo-wadaha aan la ilaalin, oo leh cufnaanta hiddo-wadaha hoose iyo waxyaabo badan oo gobollada ku celcelinaya Baloogyada loo yaqaan 'Conservative blocks' ayaa lagu xisaabtamaa 70% (12440) dhammaanba hiddo-wadayaasha 'protein-coding gen'. Qeybaha muxaafidka ah, hiddo-wadaha ayaa sida caadiga ah si aad ah ugu kala fog celcelis ahaan masaafada dhexdhexaadka ah ee 604 bp. Meelaha u dhexeeya baloogyada muxaafidka ah, masaafada u dhaxeysa ayaa ka weyn (3700 bp) sababtoo ah kororka cufnaanta walxaha soo noqnoqda. Hiddo-wadayaasha sirta dhaqsaha leh ee isbeddelaya ayaa ku yaal gobollada hidda-wadaha saboolka ah.
Falanqaynta isku xigxiga ee P. Infestans genome ayaa muujisay in ku dhowaad saddex meelood hal meel hiddo-wadaha uu ka tirsan yahay xubno la is-gudbin karo. Genome-ka P. infestans wuxuu ka kooban yahay qoysas aad u kala duwan oo transposons ah genomes-yada kale ee la yaqaan Inta badan P. infestans transposons waxaa iska leh qoyska 'Gypsy'.
Tiro badan oo ka mid ah qoysaska hiddo-wadaha qaaska ah ee ku lug leh cudur-sidaha ayaa lagu aqoonsaday P. infestans genome. Qeyb muhiim ah oo ka mid ah ayaa qiraya borotiinno waxtar leh oo beddelaya jir ahaanta dhirta martida loo yahay isla markaana gacan ka geysta infekshankiisa. Waxay ka tirsan yihiin laba qaybood oo ballaadhan: Waxyeeleyaasha apoplastic-ka, kuwaas oo ka shaqeeya meelaha isku-xidhka ah (apoplasts), iyo saamileyaasha cytoplasmic, ee gala unugyada iyada oo loo marayo haustoria. Waxyeelada Apoplastic-ka waxaa ka mid ah enzymes hydrolytic oo qarsoodi ah sida proteases, lipases iyo glycosylases oo burburiya unugyada dhirta; kahortagayaasha enzymes difaaca dhirta martida loo yahay; iyo sunta necrotizing sida borotiinada Nep1-u eg (NPLs) iyo Pcf-sida borotiinada yar-yar ee qaniga ku ah qanjirrada (SCRs).
P infestans genes genor waa kuwo badan oo badiyaa ka weyn hiddo-wadayaasha aan faafin. Kuwa ugu caansan ayaa ah saamaynta cytoplasmic RXLR iyo Crinkler (CNR). Saameynaha cytoplasmic ee caadiga ah ee oomycetes waa borotiinada RXLR. Dhammaan hiddo-wadayaasha RXLR ee la helay illaa hadda waxaa ku jira kooxda amino-terminal Arg-XLeu-Arg, halkaasoo X ay tahay amino acid. Daraasadda awgeed, waxaa la soo jeediyay inay jiraan 563 RXLR hiddo-sideyaal ah oo ku jira P. infestans genome, taasoo 60% ka badan tan P. sojae iyo P. ramorum. Qiyaastii nus ka mid ah hiddo-wadayaasha RXLR ee ku jira P. infestans genome waa nooc gaar ah. RXLR saameyn waxay leeyihiin noocyo kala duwan oo taxane ah. Kuwaas waxaa ka mid ah, mid weyn iyo 150 qoysas yaryar ah. Si ka duwan sida loo yaqaan 'proteome-ka' weyn, hiddo-wadaha RXLR-ka wax-ku-oolka ah badanaa waxay ku yaalliin hiddo-wadaha saboolka ah iyo soo noqnoqda-qaniga ah ee genome Curiyeyaasha mobilada ee go'aamiya firfircoonaanta gobolladaas waxay fududeeyaan isku-soo-noqoshada hiddo-wadahaas.
Saameynaha 'Cytoplasmic CRN' waxaa asal ahaan lagu aqoonsaday P. infestans transcripts codaynta nudaha unugyada necrosis peptides. Tan iyo markii la helay, wax yar baa laga ogaa qoyska waxtarradan. Falanqaynta hiddo-wadaha 'P. Infestans genome' ayaa shaaca ka qaaday qoys aad u tiro badan oo ka kooban 196 hiddo-wadaha CRN, oo si weyn uga ballaaran kan P. sojae (100 CRN) iyo P. ramorum (19 CRN). Sida RXLRs, CRNs waa borotiinno qaabaysan oo ka kooban N-terminal LFLAK domain aad loo ilaaliyo (50 amino acids) iyo DWL ku dhow oo leh hiddo-wadayaal kala duwan. Inta badan CRNs (60%) waxay leeyihiin astaan peptide.
Suurtagalnimada CRNs kala duwan oo lagu carqaladeeyo geedi socodka gacanta ee warshadda martida loo yahay ayaa la darsay. Falanqaynta necrosis-ka dhirta, ka saarista borotiinada CRN2 waxay suurta galisay in la aqoonsado gobolka C-terminal oo ka kooban 234 amino acids (jagooyinka 173-407, DXG domain) oo keena dhimashada unugyada. Falanqaynta P. infestans hiddo-wadaha CRN ayaa shaaca ka qaaday afar gobol oo kala duwan oo C-terminal ah, oo waliba sababa dhimashada unugyada geedka dhexdiisa. Kuwaas waxaa ka mid ah meelaha cusub ee la aqoonsaday ee DC (P. Infestans waxay leeyihiin 18 hiddo iyo 49 been abuur), iyo sidoo kale D2 (14 iyo 43) iyo DBF (2 iyo 1) xayndaabyo la mid ah borotiinka kinases. Barootiinnada CRN-da ee lagu muujiyey warshad waa la ilaaliyaa (maqnaanshaha astaamaha peptides) ee unugga dhirta waxayna kiciyaan dhimashada unugyada iyadoo la adeegsanayo farsamada gudaha. 255 isku xigxig oo kale oo ay kujiraan aaladaha CRN waxay u badan tahay inaysan u shaqeyn sida hiddo ahaan.
Kordhinta tirada iyo cabirka qoysaska hidda-wadaha RXLR iyo CRN waxay u badan tahay inay sabab u tahay dib-u-soo-kicinta isku-midka ah ee aan isku midka ahayn iyo isku-darka hidda-wadaha. In kasta oo xaqiiqda ah in hiddo-wadaha ay ku jiraan tiro badan oo ka mid ah walxaha dhaqdhaqaaqa ee firfircoon, haddana weli ma jiraan caddayn toos ah oo ku saabsan wareejinta hiddo-wadayaasha wax-ku-oolka ah.
Hababka loo adeegsaday daraasadda qaab-dhismeedka dadweynaha
Daraasada qaabdhismeedka hidaha ee dadka ayaa hada ku saleysan falanqaynta dhaqamada saafiga ah ee noocyadeeda. Falanqaynta dadweynaha iyada oo aan la kala soocin dalagyada saafiga ah ayaa sidoo kale loo sameeyaa ujeedooyin gaar ah, sida, tusaale ahaan, barashada gardarrada dadweynaha ama jiritaanka noocyo u adkaysta fungicides-ka (Filippov et al., 2004; Derevyagina et al., 1999). Cilmi baarista noocan ahi waxay ku lug leedahay adeegsiga habab gaar ah, sharraxaaddeeda oo ka baxsan baaxadda dib-u-eegidgan. Falanqaynta isbarbardhiga ee noocyada, habab dhowr ah ayaa loo adeegsadaa iyadoo lagu saleynayo falanqaynta qaabdhismeedka DNA iyo daraasadda muuqaalka muuqaalka ah. Falanqaynta isbarbardhiga ee dadweynaha waa inay la macaamilaan tiro badan oo go'doomin ah, kuwaas oo ku soo rogaya shuruudo gaar ah hababka loo adeegsaday. Fikrad ahaan, waa inay buuxiyaan shuruudaha soo socda (Cooke, Lees, 2004, Mueller, Wolfenbarger, 1999):
- noqo mid raqiis ah, fududahay in la hirgaliyo, uma baahna kharashyo waqti badan, oo ku saleysan tikniyoolajiyadda guud ahaan la heli karo (tusaale ahaan, PCR);
- waa inuu abuuraa tiro ku filan oo ah astaamo calaamadeeye calaamadeysan;
- ay leeyihiin dib-u-soo-saar sare
- Isticmaal xaddiga ugu yar ee unugyada ah ee la baarayo;
- si gaar ah ugu ekaado substrate-ka (faddarada ku jirta dhaqanka waa inaysan saameyn ku yeelan natiijooyinka);
- uma baahnid adeegsiga hababka halista ah iyo kiimikooyin aad u suntan.
Nasiib darrose, ma jiraan habab u dhigma dhammaan cabbirka kor ku xusan. Daraasad isbarbardhig ah oo ku saabsan noocyada waqtigeenna, hababka waxaa loo adeegsadaa iyadoo lagu saleynayo falanqaynta astaamaha loo yaqaan 'phenotypic traits': virulence to baradho iyo noocyo tamaandho ah (baradho iyo tartamo yaanyo ah), nooca mating, spectra of peptidase isoenzymes iyo glucose-6-phosphate isomerase, iyo falanqaynta qaab dhismeedka DNA: dhererka polymorphism Qaybta xaddidaadda (RFLP), oo badanaa lagu daro baaritaanka isku-dhafka RG 57, falanqaynta ku celcelinta microsatellite (SSR iyo InterSSR), ku-dheellitirnaanta asaasiga ah (RAPD), kordhinta jajabyada xaddidaadda (AFLP), ku-dheellitirnaanta asaasiga ah ee la mid ah qaybaha walxaha moobiilka (tusaale ahaan, Inter SINE PCR), go'aaminta mitir-mitirka 'DNA haplotypes'.
Sharaxaad kooban oo ku saabsan qaababka daraasadda isbarbardhiga ee noocyada loo isticmaalo shaqada lala yeelanayo P. Infestans
Astaamaha calaamadeynta Phenotypic
Tartanka "baradhada"
Tartamada "Baradho" waa calaamadee guud ahaan la baaray oo la adeegsaday. Jinsiyadaha "Baradho fudud" waxay leeyihiin hal hiddo-wadaha barafka, kuwa "kakan" - ugu yaraan labo. Black et al. (1953), oo soo koobaya dhammaan xogta ay heli karaan, waxay ogaadeen in tartanka phytophthora uu awood u leeyahay inuu ku faafiyo dhirta hiddo-wadaha / hiddo-wadaha iska caabinta ee u dhigma P. infestans hiddo-wadaha / hiddo-wadaha, wuxuuna helay jinsiyadaha 1, 2, 3, iyo 4 ee ku dhaca dhirta. oo leh hiddo-wadaha R1, R2, R3 iyo R4, siday u kala horreeyaan, i.e. isdhexgalka ka dhexeeya dulinka iyo martida loo yahay waxay ku dhacdaa iyadoo loo eegayo hiddo-wadaha mabda'a hiddo-wadaha. Dheeraad ah, Madow, oo ay kaqeybgaleen Gallegly iyo Malcolmson, waxay ogaadeen hiddo-wadayaasha iska caabinta R5, R6, R7, R8, R9, R10 iyo R11, iyo sidoo kale jinsiyadaha u dhigma (Madow, 1954; Madow & Gallegly, 1957; Malcolmson & Black, 1966; Malcolmson, 1970).
Waxaa jira xog ballaaran oo ku saabsan isir ahaanta jinsiga ee cudur-sidaha ka yimid gobollada kala duwan. Anaga oo aan si faahfaahsan u falanqeyneyn xogtaan, waxaan ku tuseynaa kaliya isbedel guud: halkaasoo noocyo leh hidde-sideyaal cusub oo iska caabin ah ama iskudhafkooda la adeegsaday, markii hore waxaa jiray xoogaa daciif ah oo soo daahay goor dambe, laakiin markaa tartammo ay la socdaan hiddo-wadayaal u dhigma ayaa u muuqday oo la soo xulay oo dillaaca dillaaca dambe ayaa dib u billowday. Xanuun gaar ah oo ka dhan ah 4 hidde-sideyaasha ugu horreeya ee iska caabinta (R1-R4) ayaa marar dhif ah lagu arkay ururinta la ururiyey ka hor inta aan la bilaabin abuurista noocyo hidde-sideyaashan ah, laakiin tirada noocyada murugada leh ayaa si aad ah u kordhay markii cudur-sidaha cudur-sidaha lagu daaweynayo noocyada hiddo-sideyaashaan. Hidda-wadaha 5-11, dhanka kale, wuxuu ku badnaa guud ahaan aruurinta (Shaw, 1991).
Daraasad ku saabsan saamiga jinsiyadaha kaladuwan inta lagu jiro xilliga koritaanka, oo la qabtay dhammaadkii 1980-meeyadii, ayaa muujisay in bilowga horumarinta cudurka, kloneyaal leh gardarro hooseeya iyo 1-2 hiddo-wadayaasha ku badan dadka.
Dheeraad ah, marka la soo gaadho nabarada goor dambe, isku soo ururinta midabyada asalka ah ayaa hoos u dhacaya tirada tirooyinka "kakan" ee leh rabshad badan ayaa sii kordheysa. Dhacdooyinka dambe dhamaadka xilli ciyaareedka ayaa gaaraya 100%. Markaad keydinaysid digriiga, waxaa jira hoos u dhac ku yimaada dagaalka iyo luminta hiddo-wadayaasha shakhsiga ah. Dhaqdhaqaaqa beddelka dhirta ayaa ku dhici kara noocyo kala duwan siyaabo kala duwan (Rybakova & Dyakov, 1990). Si kastaba ha noqotee, daraasaddeenii 2000-2010 waxay muujisay in jinsiyadaha isku dhafan laga helo bilowga hore ee epiphytotics ee noocyada ka go'doonsan baradhada iyo yaanyada labadaba. Tani waxay u badan tahay inay sabab u tahay isbeddelada ku yimid dadka P. Infestans ee ku nool Ruushka.
Markii la gaadho 1988-1995, inta jeer ee ay dhacaan "superraces" oo leh dhammaan ama ku dhowaad dhammaan hidda-wadayaasha firirrada ee gobollada kala duwan ayaa gaadhay 70-100%. Xaaladda noocaas ah ayaa lagu xusay, tusaale ahaan, Belarus, gobollada Leningrad, Moscow, Waqooyiga Ossetia iyo Jarmalka (Ivanyuk et al., 2002a, 2002b; Politiko, 1994; Schober-Butin et al., 1995).
Tartanka "Yaanyo"
Beerista yaanyada, kaliya 2 hiddo-wadaha oo iska caabin ah goor dambe ayaa laga helay - Ph1 (Gallegly & Marvell, 1955) iyo Ph2 (Al-Kherb, 1988). Sida kiisaska baradhada, isdhexgalka u dhexeeya yaanyada iyo P. infestans waxay ku dhacdaa hiddo-sidaha. Tartanka T0 wuxuu ku dhacaa noocyo aan lahayn hiddo-wadayaal iska caabin ah (inta badan noocyada warshadaha loo isticmaalo), T1 wuxuu ku dhacaa noocyo hiddo-wadaha Ph1 (Ottawa), iyo T2-ga wuxuu ku dhacaa noocyo hiddo-wadaha Ph2.
Ruushka dhexdiisa, ku dhowaad gaar ahaan T0 ayaa laga helay baradhada; T0 wuxuu ku adkaaday yaanyada bilowgii xilli ciyaareedka, laakiin markii dambe waxaa lagu beddelay T1 tartanka (Dyakov et al., 1975, 1994). Ka dib 2000, T1 baradho dad badan ayaa bilaabay inuu dhaco bilowga hore ee epiphytotics. Mareykanka gudahiisa, noocyada baradhada waxay ahaayeen kuwa aan cuduro ku aheyn yaanyada, sidoo kale jinsiyadaha T0, T1, iyo T2, halka T1 iyo T2 ay ku badnaayeen yaanyada (Vartanian & Endo, 1985; Goodwin et al., 1995).
Nooca rimay
Si loo sameeyo daraasadda, tijaabiyaha (tixraaca) muruqyada leh noocyada isweheshiga ee la yaqaan - A1 iyo A2 - ayaa loo baahan yahay. Go'doominta tijaabada ah waxaa lagu tallaalay iyaga oo labo-labo ah suxuunta Petri oo leh oat agar dhexdhexaad. Ka-dib markii la soo xareynayo 10 maalmood, taarikada waxaa lagu baarayaa joogitaanka iyo maqnaanshaha oospores-ka dhexdhexaadka aagga xiriirka ee noocyada. Waxaa jira 4 ikhtiyaar: culeyska waxaa iska leh nooca isku midka ah ee loo yaqaan 'A1 mating', haddii uu ka sameysmo daroogo tijaabiyaha A2, illaa A2, haddii uu ka sameysmo walxo leh tijaabiyaha A1, A1A2, haddii uu ka sameysmo dheecaan labada tijaabo, ama nadiif yahay (00), haddii uusan sameysan oospore iyada oo aan lahayn tijaabo (labada koox ee ugu dambeeya waa dhif).
Si dhakhso leh loo go'aamiyo noocyada isutagga, isku dayga ayaa la sameeyay si loo aqoonsado gobollada hiddo-wadaha la xiriira nooca isku-tagga, iyadoo looga dan leeyahay isticmaalkooda dheeraadka ah si loo go'aamiyo nooca isugeynta PCR. Mid ka mid ah tijaabooyinkii ugu horreeyay ee lagu guuleysto si loo aqoonsado goobta noocan oo kale ah waxaa sameeyay cilmibaarayaal Mareykan ah (Judelson et al., 1995). Iyagoo adeegsanaya habka RAPD, waxay awoodeen inay aqoonsadaan gobolka W16 ee laxiriira nooca isugeynta ee faraca labada gooniya oo isdhaafsan, iyo naqshadeynta labo astaamo 24-bp ah oo loogu talagalay xoojinta (W16-1 (5'-AACACGCACAAGGCATATAAATGTA-3 ') iyo W16-2 (5' -GCGTAATGTAGCGTAACAGCTCTC-3 ') Kadib xaddidaadda alaabada PCR ee la socota xayiraadda enzyme HaeIII, waxaa suurtagal noqotay in lakala saaro go'doominta noocyada lammaanaha A1 iyo A2.
Isku day kale oo lagu helayo calaamadaha PCR si loo go'aamiyo noocyada isutagga waxaa sameeyay cilmibaarayaal Kuuriyaan ah (Kim, Lee, 2002). Waxay aqoonsadeen alaabooyin gaar ah iyagoo adeegsanaya habka AFLP. Natiijo ahaan, labo nooc oo hordhac ah PHYB-1 (horay) (5'-GATCGGATTAGTCAGACGAG-3 ') iyo PHYB-2 (5'-GCGTCTGCAAGGCGCATTTT-3') ayaa la soo saaray, taasoo u oggolaaneysa kordhinta xulashada ee genome gen ee la xiriira isugeynta A2. Kadib, way sii wadeen shaqadan waxayna qorsheeyeen astaamaha 5 'AAGCTATACTGGGACAGGGT-3' (INF-1, horay) iyo 5'-GCGTTCTTTCGTATTACCAC-3 '(INF-2), taas oo u oggolaaneysa kordhinta xulashada ee gobolka Mat-A1 astaamaha noocyada noocyada nooca A1. Isticmaalka baaritaanka cudurka PCR ee noocyada isutagga waxay muujiyeen natiijooyin wanaagsan oo ku saabsan daraasadda P. dadka ku soo duulay Czech Republic (Mazakova et al., 2006), Tunisia (Jmour, Hamada, 2006), iyo gobollada kale. Shaybaarkayaga (Mytsa, Elansky, lama daabicin), 34 P. infestans noocyo laga soocay baradhada iyo xubnaha yaanyada ee gobollada kala duwan ee Ruushka (Kostroma, Ryazan, Astrakhan, Moscow oblasts) ayaa la falanqeeyay. Natiijooyinka falanqaynta PCR iyadoo la adeegsanayo astaamo gaar ah oo ka badan 90% waxay la jaanqaadeen natiijooyinka falanqaynta nooca isku-tagga ee habka dhaqanka ee nafaqada leh.
Jadwalka 1. Kala duwanaanshaha caabbinta ee ku dhexjirta aaladda 'Sib 1 clone' (Elansky et al., 2001)
Goobta tusaalaha | Tirada go'doominta la falanqeeyay | Tirada xasaasiga ah (S), daciifka u adkaysta (SR) iyo noocyada iska caabinta (R), kumbiyuutarada (%) | ||
S | SR | R | ||
G. Vladivostok | 10 | 1 (10) | 4 (40) | 5 (50) |
G. Chita | 5 | 0 | 0 | 5 (100) |
Irkutsk | 9 | 9 (100) | 0 | 0 |
G. Krasnoyarsk | 13 | 12 (92) | 1 (8) | 0 |
Magaalada Yekaterinburg | 15 | 8 (53) | 1 (7) | 6 (40) |
O. Sakhalin | 66 | 0 | 0 | 66 (100) |
Gobolka Omsk | 18 | 0 | 0 | 18 (100) |
U adkaysiga Metalaxyl calaamadeeyaha dadweynaha
Horraantii 1980-meeyadii, dillaac xoog leh oo ku yimid duqeymo goor dambe ka dhashay metalaxyl-u adkaysta noocyada infestans ayaa lagu xusay gobollo kala duwan. Beeraha baradhada ee wadamo badan waxaa ku dhacay khasaaro fara badan (Dowley & O'Sullivan, 1981; Davidse et al., 1983; Derevyagina, 1991). Tan iyo markaas, dalal badan oo adduunka ah, kormeer joogto ah oo ku saabsan dhacdooyinka noocyada u adkaysta phenylamide-ka ee dadka ku dhaca P. Marka lagu daro qiimeyn wax ku ool ah oo ku saabsan rajada isticmaalka daawooyinka ay ku jirto phenylamide, dhisida nidaam ka mid ah tallaabooyinka ka hortagga iyo saadaalinta epiphytoties, iska caabinta daawooyinkan ayaa noqotay mid ka mid ah astaamaha calaamadaha si ballaaran loogu adeegsado falanqaynta isbarbardhiga ee dadka cudurkan keena. Si kastaba ha noqotee, isticmaalka iska caabbinta metalaxyl ee daraasaadka tirada dadka waa in la fuliyaa iyadoo la tixgelinayo xaqiiqda ah: 1 - aasaaska hidda-wadaha ee caabbinta aan weli si sax ah loo go'aamin, 2 - caabbinta metalaxyl waa astaamo xulasho ahaan ku tiirsan oo ku kala duwanaan kara isticmaalka phenylamides, 3 - kala duwan heerka xasaasiga ah ee noocyada metalaxyl ee ku jira hal xariijin hal wadh (miiska 1).
Muuqaalka isozymes
Calaamadaha Isozyme badanaa way ka madax bannaan yihiin xaaladaha dibadda, waxay muujiyaan dhaxalka Mendelian waana kuwo caan ah, oo u oggolaanaya in la kala saaro homo- iyo heterozygotes. Isticmaalka borotiinada sida astaamaha hiddo-wadaha ayaa suurtogal ka dhigeysa in la aqoonsado labada habayn ee isku-dhafan ee hiddo-wadaha, oo ay ku jiraan koromosoomyada iyo is-beddelka hidde-wadaha, iyo beddelka hal amino acid.
Daraasadaha electrophoretic ee borotiinada waxay muujiyeen in enzymes badankood ay ku jiraan noolaha qaab jajabyo kaladuwan oo ku kala duwan dhaqdhaqaaqa electrophoretic. Jajabkaani waa natiijada ka dambeynta qaabeynta noocyo badan oo enzymes ah oo ay leeyihiin loci kala duwan (isozymes ama isozymes) ama luuqado kala duwan oo isku meel ah (allozymes ama alloenzymes). Taasi waa, isozymes waa noocyo kala duwan oo hal enzyme ah. Noocyada kaladuwan waxay leeyihiin waxqabad isku mid ah, laakiin waxyar ayey ku kala duwan yihiin beddellada amino acid ee peptide iyo mas'uuliyadda. Kala duwanaanshaha noocan ah ayaa la muujiyaa inta lagu jiro electrophoresis.
Markaad baraneyso noocyada kala duwan ee 'P. infestans', muuqaalka 'isoenzymes' ee laba borotiin, peptidase iyo glucose-6-phosphate isomerase, ayaa la isticmaalaa (enzyme-kan ayaa monomorphic ku ah dadka reer Ruushka; sidaa darteed, hababka daraasaddiisa laguma soo bandhigin shaqadan). Si loo kala saaro isozymes-ka aagga korantada, u diyaargarowga borotiinka ee laga soocay noolaha la barto ayaa lagu dabaqayaa saxanka jel ee aagga korantada ku jira. Qiyaasta faafitaanka borotiinnada shaqsiyeed ee jelku waxay kuxirantahay qarashka iyo culeyska molikaliska; sidaa darteed, aagga korantada, isku-darka borotiinnada waxaa loo qaybiyaa jajabyo shaqsiyeed, oo lagu arki karo iyadoo la adeegsanayo midabbo gaar ah
Daraasadda peptidase isoenzymes waxaa lagu sameeyaa cellulose acetate, istaarijka ama polyacrylamide jel. Midda ugu habboon ayaa ah habka ku saleysan adeegsiga jel cellulose acetate oo ay soo saartay shirkadda Helena Laboratories Inc. Uma baahna qaddar badan oo ah qalabka baaritaanka, waxay u oggolaaneysaa mid ka mid ah inuu helo kooxo isku dhejisan oo ku jira jelka ka dib electrophoresis labadaba labadaba enzyme loci, hirgelinteeda uma baahna waqti badan iyo kharashka sheyga (Sawirka 2).
Qayb yar oo mycelium ah ayaa loo wareejiyaa microtube 1,5 ml, 1-2 dhibcood oo biyo ah ayaa la siiyaa. Intaa ka dib, muunada waa la isku qurxiyo (tusaale ahaan, layliga korantada oo leh lifaaq balaastig ah oo ku habboon microtube) oo la mariyo 25 ilbidhiqsi centrifuge 13000 rpm. 8 μl microtube kasta. supernatant-ka waxaa loo wareejiyaa saxanka qaboojiyaha.
Jellada 'cellulose acetate gel' ayaa laga soo saaraa weelka keydka ah, oo lagu tirtiraa inta u dhexeysa labada warqadood ee warqadda shaandhada ah waxaana lagu ridayaa lakabka shaqada ilaa salka caagga ah ee qalabka wax lagu buuxiyo. Xalka saxanka waxaa ku wareejinaya codsadaha gelinta jeer 2-4 jeer. Jel waxaa loo wareejiyaa qolka electrophoresis,
Jadwalka 2. Isku-darka xalka loo adeegsaday wasakheynta jel cellulose acetate falanqaynta peptidase isoenzymes, dhibic rinji ah (bromophenol buluug) ayaa la dhigayaa geeska geeltiisa.
TRIS HCl, 0,05M, Ph 8,0 2 ml
Peroxidase, 1000 U / ml 5 dhibcood
o-dianisidine, 4 mg / ml 8 dhibcood
MgCl2, 20 mg / ml 2 dhibcood
Gly-Leu, 15 mg / ml 10 dhibcood
L-amino-acid oxidase, 20 u / ml 2 dhibcood
Electrophoresis waxaa la sameeyaa 20 daqiiqo. at 200 V. Ka dib electrophoresis, jelka waxaa loo wareejiyaa miiska rinjiyeynta oo lagu wasakheeyaa xalka rinjiyeynta gaarka ah (Shaxda 2). 10 ml oo ah 1,6% DIFCO agar ayaa si hordhac ah loogu dhalaalay foornada microwave, oo la qaboojiyey ilaa 60 ° C, ka dib 2 ml oo agar ah ayaa lagu qasan yahay rinjiga isku darka oo lagu shubay jel. Karbaashyo waxay soo muuqdaan 15-20 daqiiqo gudahood. Daawada loo yaqaan 'L-amino-acid oxidase reagent' ayaa isla markiiba lagu daraa ka hor intaan lagu darin xalka agar la shubay.
Dadka reer Ruushka, Pep 1 goobtiisa waxaa matala genotypes 100/100 iyo 92/100. Homozygote 92/92 waa dhif iyo naadir (qiyaastii 0,1%). Locus Rehr 2 waxaa matalaya seddex genotypes 100/100, 100/112, iyo 112/112, dhamaan 3 noocana waa wax iska caadi ah (Elanky iyo Smirnov, 2003, Jaantus 2).
Cilmi baarista
Xakamaynta dhererka polymorphism oo leh isku-darka soo socda (RFLP-RG 57)
Wadarta DNA-da waxaa lagu daaweeyaa enzyme xaddidaadda Eco R1, jajabka DNA-ga waxaa kala soocaya electrophoresis ku jira geel agarose ah. Nukliyeerka DNA waa mid aad u ballaaran oo leh taxaneyaal badan oo soo noqnoqda, taas oo adkeyneysa in si toos ah loo falanqeeyo jajabyada badan ee ay heleen ficil-celinta xaddidaadda enzymes. Sidaa darteed, jajabyada DNA-da ee loo kala saaray jellada waxaa loo wareejiyaa xuub qaas ah waxaana loo adeegsadaa isku-darka baaritaanka RG 57, oo ay ku jiraan nukleotides ay ku calaamadeysan yihiin calaamadaha shucaaca ama nalka. Baadhitaankani wuxuu ku dhexjiraa taxanaha genomic soo noqnoqda (Goodwin et al., 1992, Forbes et al., 1998). Ka dib markii la muujiyey natiijooyinka isku-darka aaladda iftiinka ama shucaaca, muuqaal isku-dhaf ah oo isku-dhafan (faro-qaadis) ayaa la helaa, oo ay matalaan jajabyada 25-29 (Forbes et al., 1998). Carruurta Asexual (clonal) waxay yeelan doonaan isla astaamo. Marka la isku habeeyo kooxaha ku xiran electrophoretogram, qofku wuxuu xukumi karaa isku ekaanshaha iyo kala duwanaanshaha noolaha isbarbar dhiga.
Noocyada 'Mitochondrial DNA haplotypes'
Inta badan unugyada 'eukaryotic', mtDNA waxaa lagu soo bandhigaa qaab labo-xarigle ah oo wareeg ah oo loo yaqaan 'molecule DNA', taas oo ka duwan koromosoomyada nukliyeerka ee unugyada 'eukaryotic', oo celcelis ahaan u sameeya qayb ahaan isla markaana aan la xiriirin maaddooyinka borotiinka.
Hidda-wadaha mitochondrial ee P. infestans ayaa la isku xigxigay, tiro shaqooyin ahna waxaa loo hibeeyay falanqaynta dhererka jajabka jajabka (Carter et al, 1990, Goodwin, 1991, Gavino, Fry, 2002). Ka dib markii Griffith iyo Shaw (1998) ay soo saareen hab fudud oo dhakhso leh oo lagu ogaanayo mtDNA haplotypes, astaamahani wuxuu noqday mid ka mid ah kuwa ugu caansan ee daraasadaha P. Infestans. F2-R2 (Jadwalka 4) iyo xaddiditaankooda xiga ee xaddidaadda enzymes MspI (jajabka 4aad) iyo EcoR3 (jajabka 1aad). Qaabka ayaa kuu ogolaanaya inaad aqoonsato 1 nooc oo haplotypes ah: Ia, IIa, Ib, IIb. Nooca II wuxuu kaga duwan yahay nooca I jiritaanka meesha la geliyay 2 bp cabir ahaan iyo meel kale oo ka mid ah goobaha xaddidaadda ee gobollada P4 iyo P1881 (Sawirka 2).
Laga soo bilaabo 1996, oo ka mid ah noocyada laga soo ururiyey dhulka Ruushka, kaliya waxaa la xusay haplotypes Ia iyo IIa (Elansky et al., 2001, 2015). Waa la aqoonsan karaa ka dib kala soocida alaabooyinka xaddidaadda leh aasaasiga F2-R2 ee aagga korantada (Sawir 4, 5). Noocyada mtDNA waxaa loo isticmaalaa falanqaynta isbarbardhiga noocyada iyo dadka. Daraasado dhowr ah, noocyada mitochondrial DNA ayaa loo isticmaalay in lagukala saaro khadadka cnal iyo passportize P. infestans isolates (Botez et al., 2007; Shein et al., 2009). Iyadoo la adeegsanayo habka PCR-RFLP, waxaa lagu soo gabagabeeyay in mtDNA ay ku kala duwan tahay isla iskujirka P. infestans (Elansky iyo Milyutina, 2007). Xaaladaha Kordhinta: 1x (500 sec. 94 ° C), 40x (30 sec. 90 ° C, 30 sec. 52 ° C, 90 sec. 72 ° C); 1x (5 daqiiqo. 72 ° C). Isku darka falcelinta: (20 μl): 0,2 U Taq DNA polymerase, 1x 2,5 mM MgCl2-Taq buffer, 0,2 mM kasta dNTP, 30 pM Primer iyo 5 ng ee DNA-da la falanqeeyay, biyaha la karkaray - ilaa 20 μl.
Xaddidaadda alaabada PCR waxaa lagu fuliyaa 4-6 saacadood heerkulka 37 ° C. Xaddidaadda xaddidaadda (20 μl): 10x MspI (2 μl), 10x buffer buffer (2 μl), biyo la gooyey (6 μl), wax soo saarka PCR (10 μl).
Jadwalka 3. Astaamaha loo adeegsado kordhinta gobollada polymorphic mtDNA
Meelo | Horudhac | Dhererka aasaasiga ah iyo meelaynta | Dhererka alaabada PCR | Xaddid |
---|---|---|---|---|
P2 | F2: 5'- TTCCCTTTTGGCCCCTAGCGATT | 21; 13619-13639 | 1070 | MspI |
R2: 5'- TTGGGGGTTTAGAGCACATACA | 22; 14688-14667 | |||
P4 | F4: 5'- TGGTCATCCAGAGGTTTATGTT | 22; 9329-9350 | 964 | EcoRI |
R4: 5 - CCGATACCGATACCAGCACACA | 22; 10292-10271 |
Kordhinta aasaasiga ah ee Random (RAPD)
Markaad fulineyso RAPD, hal nooc ayaa la adeegsadaa (mararka qaarkood dhowr astaamo isku mar) oo leh taxane aan kala sooc lahayn oo nucleotide ah, badanaa 10 nucleotides oo dherer ah, oo leh maadad sare (laga bilaabo 50%) GC nucleotides iyo heerkul hoose oo qaboojin ah (qiyaastii 35 ° C). Hordhacyada noocan oo kale ahi waxay ku "degaan" goobo fara badan oo is kaabaya oo ku jira hiddo-wadaha. Kordhinta ka dib, tiro badan oo kiciyeyaal ah ayaa la helaa. Tiradoodu waxay kuxirantahay aasaasiga (yada) la isticmaalay iyo xaaladaha falcelinta (diirada MgCl2 iyo heerkulka annealing).
Muuqaalka amplicon waxaa lagu sameeyaa iyadoo lagu nadiifiyo polyacrylamide ama geel agarose ah. Marka la sameynayo falanqaynta RAPD, waxaa lagama maarmaan ah in si taxaddar leh loola socdo nadiifnimada waxyaabaha la falanqeeyay, maxaa yeelay wasakheynta walxaha kale ee nool waxay sababi kartaa koror weyn oo ku yimaada tirada farshaxanimada, kuwaas oo aad ugu badan falanqaynta waxyaabaha saafiga ah (Perez et al, 1998). Isticmaalka qaabkan daraasadda ee 'infestans genome' ayaa ka muuqda shaqooyin badan (Judelson, Roberts, 1999, Ghimire et al., 2002, Carlisle et al., 2001). Xulashada xaaladaha falcelinta iyo hordhaca (51 10-nucleotide primers ayaa la darsay) waxaa lagu siiyay maqaalka Abu-El Samen et al., (2003).
Falanqaynta Kucelcelinta Microsatellite (SSR)
Microsatellite ku celcelinta (soo noqnoqoshada isku xigxiga ee fudud, SSR) ayaa si isdaba joog ah loogu celcelinayaa taxane gaagaaban oo ah 1-3 (mararka qaarkood ilaa 6) nukleotides oo ku jira genomes-ka nukliyeerka ee dhammaan eukaryotes. Tirada soo noqnoqda ee isdabajoogga ah way kala duwanaan kartaa 10 ilaa 100. Microsatellite loci waxay ku dhacdaa si aad u sarreeysa oo si aad ah ama ka yar ayaa loo qaybiyaa dhammaan genome (Lagercrantz et al., 1993). Polymorphism ee taxanaha microsatelite waxay laxiriiraa kaladuwanaanshaha tirada soo noqnoqoshada ujeedada aasaasiga ah. Calaamadeeyayaasha Microsatelite-ku waa kuwo cod-bixiyayaal ah, taas oo suurtagal ka dhigaysa in loo isticmaalo falanqaynta qaab-dhismeedka dadweynaha, go'aaminta qaraabada, waddooyinka socdaalka genotype, iwm. Faa'iidooyinka kale ee astaamahan, mid waa inuu ogaadaa polymorphism-kooda sare, dib-u-soocid wanaagsan, dhexdhexaadnimo, iyo awood u leh inuu sameeyo falanqeyn otomaatig ah iyo qiimeyn. Ugu horreyntii, falanqaynta waxaa lagu sameeyay kala-soocidda alaabada falcelinta ee ku jirta gel polyacrylamide. Markii dambe, shaqaalaha Applied Biosystems waxay soo jeediyeen inay isticmaalaan astaamo dhalaalaya oo leh calaamado muujinaya astaamaha falcelinta iyadoo la adeegsanayo qalabka baaritaanka laser-ka ee otomaatiga ah (Diehl et al., 1990), ka dibna taxanaha caadiga ah ee DNA tooska ah (Ziegle et al., 1992). Calaamadeynta astaamaha leh midabyo dhaadheer oo kala duwan ayaa suurtagal ka dhigaya in lagu falanqeeyo dhowr astaamo hal mar hal haad, isla markaana, si weyn u kordhiya wax soo saarka habka iyo kordhinta saxnaanta falanqaynta.
Daabacadihii ugu horreeyay ee loogu talagalay isticmaalka falanqaynta SSR ee daraasadda P. infestans waxay soo muuqdeen horaantii 2000s. (Knapova, Gisi, 2002). Dhammaan astaamaha ay soo jeediyeen qorayaashu muu muujin heer ku filan oo polymorphism ah, hase yeeshe, labo ka mid ah (4B iyo G11) ayaa lagu soo daray calaamadaha 12 SSR ee ay soo jeediyeen Lees et al. (2006) oo markii dambena ay qaateen shabakadda cilmi baarista ee Eucablight (www.eucablight) .org) oo halbeeg u ah P. infestans. Dhowr sano ka dib, daraasad ayaa lagu daabacay abuurista nidaam loogu talagalay falanqaynta multiplex ee P. infestans DNA oo ku saleysan sideed calaamadaha SSR (Li et al., 2010). Ugu dambeyntiina, ka dib markii la qiimeeyay dhammaan astaamihii hore ee la soo jeediyay isla markaana la xushay kuwa ugu badan ee xog-ogaal u ah iyaga, iyo sidoo kale in la wanaajiyo astaamaha, astaamaha dhaldhalaalka, iyo xaaladaha sii ballaadhinta, isla kooxda qorayaashu waxay soo bandhigeen nidaam loogu talagalay falanqaynta hal-tallaabada ah, oo ay ku jiraan calaamadaha 12 (Shaxda 4; Li et al. , 2013a). Hordhacyada loo adeegsaday nidaamkan waxaa lagu xushay oo lagu calaamadeeyay mid ka mid ah afar astaamooyin dhaadheer (FAM, VIC, NED, PET) sidaa darteed kala duwanaanta cabbirka allele ee horyaallada leh calaamado isku mid ah ma aysan is-dul-marin.
Qorayaashu waxay ku sameeyeen falanqaynta kumbuyuutar PTC200 (MJ Research, USA) iyagoo adeegsanaya xirmooyinka QIAGEN multiplex PCR ama xirmooyinka QIAGEN Typeit Microsatellite PCR. Muga isku dhafka falcelinta wuxuu ahaa 12.5 μL. Xaaladaha kordhinta waxay ahaayeen sidan soo socota: QIAGEN multiplex PCR: 95 ° C (15 min), 30x (95 ° C (20 sec), 58 ° C (90 sec), 72 ° C (60 sec), 72 ° C (20 min); for QIAGEN Type-it Microsatellite PCR: 95 ° C (5 daqiiqo), 28x (95 ° C (30 sec), 58 ° C (90 sec), 72 ° C (20 sec), 60 ° C (30 min).
Kala saarista iyo muuqaalka alaabada PCR waxaa lagu fuliyay iyadoo la adeegsanayo ABI3730 otomaatig ah falanqeeyaha DNA-ga madaxbanaan (Applied Biosystems).
Jadwalka 4. Astaamaha 12 astaamaha SSR ee caadiga ah ee loo adeegsado genotyping ee P. Infestans (Li et al., 2013a)
Title | Tirada alleles | Baaxadda cabbirka alleles (bp) | Astaamaha |
PiG11 | 13 | 130-180 | F: NED-TGTTTTGGCGGGGGGGGG R |
Pi02 | 4 | 255-275 | F: NED-ACAGCAGAACAGTACCGCCC R: GTTGACCAGTTTTGCGGGCTC |
PinfSSR11 | 4 | 325-360 | F: NED-TTGGGGGGATGGGGGGCGGGG CAGCAGGAGGAGGGGGGGGCAGG |
D13 | 16 | 100-185 | F: FAM-TGCCCCCTGCTCACTCTC R: GCTCGAATTTTGGAGAGAGTTG |
PinfSSR8 | 4 | 250-275 | F: FG-AATGATGGGGCAGCAGGGGGCAGCAGTGGGGGG CAGCAGCAGGGGGGGCAG |
PinfSSR4 | 7 | 280-305 | F: CAATTTGTGGGGAGGATGATGGGGGGGGGAGGAGGGGGGGGC CAGCAGCAGCAG |
Pi04 | 4 | 160-175 | F: VIC-GAGCTGGCGGCAGGGGGG R: GTTAGAGCGGAGCAGCAGCAGGGG |
Pi70 | 3 | 185-205 | F: VIC-ATGAAAATACGATATATCATGCTGGG RGTGGGATATTTGGGGGCGGCAGAGAG |
PinfSSR6 | 3 | 230-250 | F: GTTTGGGGGGGGGGGATGATGGGGGGGCGGGGGGGCGGGGCAGGTGTGTGGG |
Pi63 | 3 | 265-280 | F: VIC-ATGGGGGGGGAGAGAGTGAGG R |
PinfSSR2 | 3 | 165-180 | F: PET-TGGGGATGATCGGCGGCAGCAGCTGCGCGGCGGCGCAGAGCAGC: GGGGAGCAGCAGC |
Pi4B | 5 | 200-295 | F: PET-AAAATAAAGCCTTGGTGTTGAATA RGGGGGGTGGAGAGTT |
Tusaalaha indha indheynta natiijooyinka gorfaynta waxaa lagu muujiyey Jaantuska. 6. Natiijooyinka waxaa lagu lafa guray iyadoo la adeegsanayo barnaamijka GeneMapper 3.7 iyadoo la isbarbar dhigayo xogta la helay iyo xogta gooni u goynta la yaqaan. Si loo fududeeyo fasiraadda natiijooyinka falanqaynta, waxaa lagama maarmaan ah in lagu daro 1-2 tixraacyada go'doominta leh genotype la yaqaan daraasad kasta.
Habka cilmi baarista ee la soo jeediyay ayaa lagu tijaabiyay tiro muunad ah oo tijaabo ah, ka dib markii ay qoreyaashu sameeyeen jaangooyooyinka hab-maamuuska u dhexeeya labo-laboratarrada, The James Hutton Institute (UK) iyo Wageningen University & Research (Netherlands), taas oo ay weheliso suurtagalnimada isticmaalka kaararka caadiga ah ee FTA ee la fududeeyay. ururinta iyo shixnadaha P. infestans tijaabooyinka DNA, waxay suurta galisay in laga hadlo suurtagalnimada isticmaalka ganacsi ee horumarkan. Intaa waxaa sii dheer, hab dhakhso leh oo sax ah oo loo yaqaan 'genotyping P. infestans' ayaa go'doomiya iyadoo la adeegsanayo falanqaynta 'multiplex SSR' taasoo suurta galisay in la sameeyo daraasado isku dheelitiran oo ku saabsan dadka cudurkan keena oo heer caalami ah, iyo abuuritaanka xog-ururin caalami ah oo ku saabsan goor-dambe oo ku saabsan qaab dhismeedka mashruuca Eucablight (www.eucablight.org), oo ay ku jiraan , oo ay kujirto natiijooyinka falanqaynta mikrosatelite-ka, waxay suurta galisay in lala socdo soo ifbaxa iyo faafitaanka genotypes cusub aduunka oo dhan.
Polymorphism dhererka jajabka xakamaynta (AFLP). AFLP (polymorphism dhererka jajabka ballaadhan) waa tiknoolajiyad loogu talagalay soo saarista astaamo molikal ah oo aan kala sooc lahayn iyadoo la adeegsanayo astaamo gaar ah. AFLP, DNA waxaa lagu daaweeyaa iskudhaf ah laba enzymis xaddidan. Adabtarada qaaska ah waxay kuxiran yihiin dhamaadka dariiqyada jajabyada xadidan.
Jajabyadaas ayaa markaa la xoojinayaa iyadoo la adeegsanayo hormoonno isku dhafan oo isku xigxiraya adabtarada iyo goobta xaddidaadda iyo iyadoo weliba lagu sido hal ama in ka badan oo saldhigyo aan kala sooc lahayn ah '3' dhammaadkooda. Qaybaha jajabka ah ee la helay waxay kuxirantahay insaymisyada xaddidan iyo nucleotides si aan kala sooc lahayn loo xushay ee 3'-dhamaadka ee hordhaca (Vos et al., 1995). AFLP - genotyping waxaa loo isticmaalaa in si deg deg ah loogu barto kala duwanaanta hidde ee noolaha kala duwan.
Sharaxaad faahfaahsan oo ku saabsan habka ayaa lagu siiyay shaqooyinka Mueller, Wolfenbarger, 1999, Savelkoul et al., 1999. Shaqooyin badan oo isbarbardhigaya xallinta hababka AFLP iyo SSR ayaa waxaa fuliyay cilmi baarayaasha Shiinaha. Astaamaha dabeecadda iyo hidde-wadaha ee 48 P. infestans isolates laga soo ururiyay shan gobol oo ku yaal Waqooyiga Shiinaha ayaa la darsay. Iyadoo lagu saleynayo muuqaalka AFLP, sideed nooc oo hiddo-wadaha DNA-ga ah ayaa la aqoonsaday, marka loo eego hiddo-wadaha SSR, oo aan lagu soo bandhigin kala duwanaansho (Guo et al., 2008).
Kordhinta oo leh astaamo la mid ah taxanaha walxaha moobiilka
Calaamadaha laga soo qaatay taxanaha 'retrotransposons' ayaa aad ugu habboon khariidaynta hidde-wadaha, daraasadda kala duwanaanta hidde-wadaha iyo geeddi-socodka isbeddelka (Schulman, 2006). Haddii horudhac loo sameeyay in lagu kabo taxadarka xasilloon ee walxaha qaarkood ee mobilada, waxaa suurtagal ah in la ballaadhiyo gobollada hidde-wadaha ee u dhexeeya. Daraasadaha ku saabsan wakiilka sababa ee duqeynta goor dambe, habka loo kordhiyo gobollada hiddo-wadaha iyadoo la adeegsanayo hordhac hordhac u ah taxanaha aasaasiga ah ee SINE (Waxyaabaha Nuclearka Gaaban ee Dhex-dhexaad ah) retroazone ayaa si guul leh loo isticmaalay (Lavrova iyo Elansky, 2003). Iyadoo la adeegsanayo qaabkan, kala duwanaanshaha ayaa la muujiyay xitaa faracyada asalka ah ee mid gooni ah. Marka la eego arrintan, waxaa lagu soo gabagabeeyay in habka isku xidhka - SINE - PCR uu yahay mid si gaar ah loo qeexay iyo heerka dhaqdhaqaaqa ee walxaha SINE ee ku jira genome Phytophthora uu sarreeyo.
Genome-ka P. infestans, 12 qoys oo ka mid ah retrotransposons gaagaaban (SINEs) ayaa lagu aqoonsaday; qaybinta noocyada retrotransposons gaagaaban ayaa la baaray, canaasiirta (SINEs) ayaa la aqoonsaday kuwaas oo laga helo genomka kaliya P. infestans (Lavrova, 2004).
Noocyada ku saabsan habka loo adeegsado qaababka daraasadda isbarbardhiga ee noocyada daraasadaha dadweynaha
Markaad qorsheyneyso daraasad, qofku waa inuu si cad u fahmaa yoolalka ay hiigsaneyso una adeegsanayo hababka ku habboon. Sidaa awgeed, hababka qaarkood ayaa suurtogal ka dhigaya in la soo saaro tiro badan oo calaamado calaamadeeyaal madaxbannaan ah, laakiin isla markaa waxay leeyihiin dib-u-soo-celin hooseeya waxayna si xoog leh ugu tiirsan yihiin reagent-yada la isticmaalay, xaaladaha falcelinta, iyo faddarada maaddada baaritaanka. Sidaa darteed, daraasad kasta oo ka mid ah koox murugo leh, waxaa lagama maarmaan ah in la isticmaalo dhowr jaangooyo (tixraac) go'doomin, laakiin xitaa kiiskan, natiijooyinka dhowr tijaabo ayaa aad u adag in la isku daro.
Kooxdan hababka ah waxaa ka mid ah RAPD, AFLP, InterSSR, InterSINE PCR. Kordhinta ka dib, tiro aad u tiro badan oo ah jajabyada DNA ee cabbirro kala duwan ah ayaa la helaa. Waxaa lagugula talinayaa inaad isticmaasho farsamooyinka noocaas ah haddii ay lagama maarmaan tahay in la sameeyo kala duwanaansho u dhexeeya noocyada isku dhow (waalid-faracooda, nooca duurjoogta, iwm), ama kiisaska falanqaynta faahfaahsan ee sambal yar loo baahan yahay. Sidaa darteed, habka AFLP waxaa si ballaaran loogu adeegsadaa khariidaynta hidde-wadaha ee P. infestans (van der Lee et al., 1997) iyo daraasadaha isdhaafsiga (Knapova, Gisi, 2002, Cooke et al, 2003, Flier et al, 2003). Hababka noocan oo kale ah kuma haboona in la isticmaalo marka la abuurayo keydka macluumaadka ee noocyada kala duwan, maxaa yeelay ficil ahaan waa wax aan macquul aheyn in la mideeyo xisaabinta natiijooyinka marka la sameynayo falanqaynta shaybaarada kala duwan.
In kasta oo ay u muuqato fudeyd iyo xawaare dil (go'doominta DNA iyada oo aan la sifeyn wanaagsan, ballaadhinta, aragtida natiijooyinka), kooxdan hababku waxay u baahan yihiin adeegsiga hab gaar ah oo loogu talagalay diiwaangelinta natiijooyinka: kala-saaridda ku jirta polyacrylamide gel oo leh astaamo (shucaac ama iftiin) horudhac ah iyo iftiinka dambe ee iftiinka ama qalabka shucaaca. Sawirka caadiga ah ee 'ethidium bromide agarose gel imaging' guud ahaan kuma haboona hababkaan maxaa yeelay tiro aad u tiro badan oo ah jajabyada DNA-da ee cabbirkoodu kala duwan yahay ayaa fuseyn kara
Hababka kale, liddi ku ah, waxay suurtogal ka dhigayaan in la soo saaro tiro yar oo astaamo ah oo leh dib-u-soo-celin heer sare ah. Kooxdan waxaa ka mid ah daraasadda mitrooga mitirka maskaxda ee DNA (kaliya laba haplotypes Ia iyo IIa ayaa lagu xusay Ruushka), noocyada isutagga (kala-goysyada badankood waxaa loo qaybiyaa noocyada 2: A1 iyo A2, is-abuurka SF-ka ayaa dhif ah in la helo) iyo peptidase isozyme spectra (labo loci Pep1 iyo Pep2) , oo ka kooban laba isozymes midkiiba) iyo gulukoos-6-fosfate isomerase (Ruushka gudahiisa malaha kala duwanaansho sifadan, in kasta oo polymorphism muhiim ah lagu xusay wadamada kale ee adduunka). Waxaa lagugula talinayaa inaad isticmaashid astaamahan markaad falanqeyneyso ururinta, ururinta keydka macluumaadka gobolka iyo adduunka. Marka laga hadlayo falanqaynta isozymes-ka iyo haplotypes-ka mitochondrial DNA, waxaa suurtagal ah in la sameeyo iyadoo aan la helin noocyo caadi ah gabi ahaanba, halka falanqaynta noocyada isutagga, loo baahan yahay laba tijaabo oo kala sooc ah oo leh noocyo caan ah oo la yaqaan.
Xaaladaha falcelinta iyo reagentyadu waxay kaliya saameyn ku yeelan karaan kala-duwanaanta alaabta electrophoretogram; muujinta astaamaha noocyada daraasaddan ayaan u badnayn.
Waqtigan xaadirka ah, dadka ugu badan ee ku nool qaybta Yurub ee Ruushka waxaa matala noocyo ka mid ah labada nooc ee isweydaarsiga (Shaxda 6), iyaga waxaa ka mid ah go'doominno noocyada Ia iyo IIa ah ee mitochondrial DNA (noocyada kale ee mtDNA ee laga helay adduunka laguma helin Ruushka wixii ka dambeeyay 1993). Muuqaalka 'peptidase isozymes' waxaa matalaya laba nooc oo genotypes ah oo ku yaal Pep1 locus (100/100, 92/92 iyo heterozygote 92/100, iyo genotype 92/92 waa dhif iyo naadir) , 0,3/2 iyo heterozygote 100/100, oo leh genotype 112/112 oo dhaca wax ka yar 100/112, laakiin sidoo kale marar badan).
Ma jirin wax isbeddel ah oo ku yimid muuqaalka 'isozymes' ee gulukooska-6-fosfate isomerase wixii ka dambeeyay 1993 (luminta khadka dhexe ee loo yaqaan 'US-1'); dhammaan barashada go'doominta waxay leeyihiin nooca 100/100 (Elansky iyo Smirnov, 2002).
Kooxda saddexaad ee hababku waxay suurtogal ka dhigeysaa in la helo koox ku filan oo ah astaamo sumadeeyaal madax-bannaan oo leh dib-u-soo-saar sare. Maanta, kooxdan waxaa ka mid ah baaritaanka RFLP-RG57, oo soo saaraya 25-29 qaybo DNA ah oo cabbirro kala duwan ah. RFLP-RG57 waa la isticmaali karaa labadaba marka la falanqeynayo shaybaarada iyo keydinta keydka macluumaadka. Si kastaba ha noqotee, qaabkani waa mid aad uga qaalisan kuwii hore, waa waqti qaadasho, wuxuuna u baahan yahay qaddar aad u tiro badan oo DNA aad u nadiif ah. Sidaa darteed, cilmibaaraha waxaa lagu qasbay inuu xadido mugga maadada la tijaabiyay.
Horumarinta RFLP-RG57 horaantii 90-meeyadii qarnigii la soo dhaafay waxay si aad ah u xoojisay daraasadaha dadweynaha ee ku saabsan wakiilka sababa duqeynta goor dambe. Waxay noqotay aasaaska habka ku saleysan xulashada iyo falanqaynta "Khadadka Qabiilka" (eeg hoos). Iyada oo ay weheliso RFLP-RG57, nooca isugeynta, sawirka faraha DNA (habka RFLP-RG57), muuqaalka 'peptidase' iyo 'glucose-6-phosphate isomerase isoenzymes', iyo nooca mitochondrial DNA ayaa loo isticmaalaa in lagu garto khadadka cnal. Waad ku mahadsan tahay isaga, waxaa la tusay al., 1994), bedelida dadkii hore iyo kuwa cusub (Drenth et al, 1993, Sujkowski et al, 1994, Goodwin et al, 1995a), ayaa shaaca ka qaaday khadadka cnal ee ka jira dalal badan oo adduunka ah. Daraasado lagu sameeyay noocyada Ruushka ee adeegsanaya qaabkan ayaa muujiyay polymorphism genotypic sare ah oo ka mid ah muruqyada qaybta Yurub iyo monomorphism ee dadweynaha qaybaha Aasiya iyo Bariga Fog ee Ruushka (Elansky et al, 2001). Haddana qaabkani waa kan ugu weyn ee daraasadaha dadweynaha ee P. infestans. Si kastaba ha noqotee, baahintiisa ballaaran waxaa hor istaagay qiimaheeda qaaliga ah iyo xooggeeda shaqo ee fulinta.
Farsamo kale oo rajo leh oo si dhif ah loogu adeegsado daraasadaha P. infestans waa falanqaynta microsatelite-ka (SSR). Waqtigan xaadirka ah, qaabkan waxaa si ballaaran loogu adeegsadaa in lagu sooco khadadka cnal. Falanqaynta noocyada, astaamaha noocan oo kale ah astaamaha sida joogitaanka hiddo-wadayaasha noocyada baradhada (Avdey, 1995, Ivanyuk et al., 2002; Ulanova et al., 2003) iyo yaanyo aad ayaa loo isticmaali jiray (waana la sii isticmaalayaa). Waqtigaan la joogo, hiddo-wadayaasha noocyada baradhada ayaa lumiyay qiimahoodii sida astaamaha calaamadaha loogu talagalay daraasadaha dadweynaha iyadoo ay ugu wacan tahay muuqaalka ugu badan (ama ku dhow) tirada hiddo wadayaasha ee ku badan inta badan go'doominta. Isla mar ahaantaana, hiddo-wadaha 'T1' virulence hiddo-wadaha ee yaanyada ku xambaara hidda-wadaha u dhiganta 'Ph1' ayaa wali si guul leh loogu adeegsaday astaamo sumadeed (Lavrova et al., 2003; Ulanova et al., 2003).
Daraasado badan, caabbinta kaneecada waxaa loo isticmaalaa calaamadee ahaan. Sifadaani waa mid aan loo baahnayn in loo isticmaalo daraasadaha dadweynaha iyadoo ay ugu wacan tahay muuqaalka fudud ee is-beddelka caabbinta ee khadadka isku-xirnaanta ka dib marka la isticmaalo metalaxyl- (ama mefenoxam-) oo ay ku jiraan sunta lagu buufiyo berrinka. Tusaale ahaan, kala duwanaansho weyn oo ku saabsan heerka iska caabinta ayaa lagu muujiyey gudaha xariiqda 'Sib1 clonal line' (Elansky et al., 2001).
Sidaa awgeed, nooca isugeynta, peptidase isoenzyme spectrum, nooca mitochondrial DNA, RFLP-RG57, SSR ayaa ah astaamaha calaamadaha doorbida ee abuurista bangiyada xogta iyo calaamadeynta noocyada kaladuwan ee uruurinta. Si loo barbardhigo shaybaarada xaddidan, haddii ay lagama maarmaan tahay in la adeegsado tirada ugu badan ee astaamaha calaamadaha, waxaad isticmaali kartaa AFLP, RAPD, InterSSR, Inter-SINE PCR (Shaxda 5). Si kastaba ha noqotee, waa in la xusuusnaado in hababkaani ay yihiin kuwo liidata oo dib-loo-soo-saari karo, iyo tijaabo kasta oo shaqsi ah (ballaadhinta wareegga elektroforesis) waa lagama maarmaan in la isticmaalo dhowr go'doomin tixraac ah.
Jadwalka 5. Isbarbardhigga hababka kala duwan ee baaritaanka noocyada P. infestans
Shuruudaha | TS | Isofer cops | MtDNA | RFLP-RG57 | RAPD | ISSR | SSR | AFLP | Rev |
---|---|---|---|---|---|---|---|---|---|
Tirada macluumaadka | Н | Н | Н | С | В | В | С | В | В |
Taranka | В | В | В | В | Н | Н | С | С | С |
Suurtagalnimada alaabooyin | Н | Н | Н | Н | В | С | Н | С | В |
qiimaha | Н | С | Н | В | Н | Н | Н | С | Н |
Awoodda foosha | Н | Н | Н | В | NS * | NS * | Н | С | NS * |
Xawaaraha Falanqaynta ** | В | Н | Н | С | Н | Н | Н | Н | Н |
Fiiro gaar ah: H - hoose, C - dhexdhexaad, B - sare; * - Awoodda foosha ayaa hooseysa markii la isticmaalayo geel agarose ama otomaatig ah
genotyper, dhexdhexaad - by muxaadaro ku jirta polyacrylamide gel leh astaamo summad leh,
** - ma tirin waqtiga lagu qaatay sii kordheynta mycelium ee go'doominta DNA.
Qaab dhismeedka dadka
Khadadka qabiilooyinka
Maqnaanshaha dib-u-ururinta ama waxtarkeeda aan yareyn ee qaab-dhismeedka dadweynaha, dadku waxay ka kooban yihiin tiro cayiman oo isku mid ah, is-weydaarsiga hidda-socodka ee u dhexeeya oo aad u dhif ah.
Dadyowga noocan oo kale ah, waa macluumaad aad u badan oo la barayo maaha inta jeer ee hiddo-wadaha shakhsiga ah, laakiin inta jeer ee genotypes ee leh asal guud (khadadka loo yaqaan 'clonal lines or clonal lineages') oo waxay ku kala duwan yihiin oo keliya is-beddelka dhibcaha. Daraasadaha dadweynaha ee cudurka dabaysha ee dambe iyo falanqaynta khadadka isku xidhka ayaa si xawli ah u soo degay tan iyo markii uu soo ifbaxay habka RFLP-RG57 horaantii 90-meeyadii qarnigii la soo dhaafay. Iyada oo ay weheliso RFLP-RG57, nooca mating, spectra of peptidase iyo glucose-6-phosphate isomerase isoenzymes, iyo nooca mitochondrial DNA ayaa loo isticmaalaa in lagu garto khadadka cnal. Sifooyinka khadadka ugu caansan ee caadiga ah waxaa lagu muujiyey Shaxda 6.
Clone US-1 ayaa ku badnaa dadka meel kasta ilaa dhammaadkii 80-yadii, ka dib oo ay bilawday in lagu beddelo mid kale oo laga waayo Yurub iyo Waqooyiga Ameerika. Hadda waxaa laga helaa Bariga Fog (Filibiin, Taiwan, Shiinaha, Jabbaan, Kuuriya, Koh et al., 1994, Mosa et al, 1993), Afrika (Uganda, Kenya, Rwanda, Goodwin et al, 1994, Vega-Sanchez et al., 2000; Ochwo et al., 2002) iyo Koonfurta Ameerika (Ecuador, Brazil, Peru, Forbes et al., 1997, Goodwin et al., 1994). Wax nooc ah oo ka tirsan safka US-1 laguma aqoonsan Australia oo keliya. Sida muuqata, P. infestans go'doominta waxay Australia yimaadeen iyagoo wata hijro kale oo guuritaan ah (Goodwin, 1997).
Clone US-6 wuxuu ka soo haajiray waqooyiga Mexico una aaday Kaliforniya dabayaaqadii 70-yadii wuxuuna halkaa ka sababay cudur faafa oo ku jira baradhada iyo yaanyada 32 sano ka dib cudur la'aan. Cadaadis xumadeeda awgeed, waxay barakicisay dhagxaanta US-1 waxayna bilaabeen inay xukumaan xeebta galbeed ee Mareykanka (Goodwin et al., 1995a).
Noocyada genotypes US-7 iyo US-8 waxaa laga helay Mareykanka 1992, mar horena 1994 ayaa si weyn loogu qeybiyay Mareykanka iyo Kanada. Inta lagu jiro hal xilli beer, clone US-8 wuxuu awood u leeyahay inuu gebi ahaanba barakiciyo dhirta US-1 ee baradhada dhulalka baradhada oo markii hore ay ku dhacday labada isku midba marka loo fiirsho isku mid (Miller iyo Johnson, 2000).
Clones BC-1 to BC-4 waxaa lagu aqoonsaday British Columbia tiro yar oo ka go'doominta Goodwin et al., 1995b). Clone US-11 wuxuu si ballaadhan ugu faafay Mareykanka wuxuuna ku beddelay US-1 Taiwan. Clones JP-1 iyo EC-1, oo ay weheliso isku-xidhka US-1, ayaa ku badan Japan iyo Ecuador, siday u kala horreeyaan (Koh et al., 1994; Forbes et al., 1997).
SIB-1 waa xarig ka dhexjirta Ruushka dhul ballaaran oo ka socda gobolka Moscow illaa Sakhalin. Gobolka Moosko, waxaa laga helay 1993, qaar ka mid ah dadweynaha ku noolna waxay ka koobnaayeen noocyo kala duwan oo xariiqan, oo aad ugu adkaysta metalaxyl. Wixii ka dambeeyay 1993, baahsanaanta khariidadan ayaa si aad ah hoos ugu dhacday. Bannaanka Ural-ka, 1997 - 1998, SIB-1 waxaa laga helay meel kasta marka laga reebo Territory-ka Khabarovsk (goobta loo yaqaan 'SIB-2 clone' ayaa ku baahsan halkaas). Kala-soocidda durdurrada ee noocyada leh noocyo kala duwan oo isu geyn ah ayaa ka reebaysa habka galmada ee Siberia iyo Bariga Fog. Gobolka Moosko, marka loo eego Siberia, dadweynaha waxaa matala clones badan; ku dhowaad go'doon kastaa wuxuu leeyahay genotype multilocus gaar ah (Elansky et al., 2001, 2015). Kala duwanaanshahan laguma sharixi karo oo keliya soo dejinta noocyo ka mid ah fangaska oo laga keenay qaybo kala duwan oo adduunka ah oo leh waxyaabo abuur ah oo dibedda laga keenay. Maaddaama labada nooc ee isweheshiga ay ku dhacaan dadka, waxaa suurtagal ah in kala duwanaanshaheeda ay sidoo kale sabab u tahay dib-isu-soo-kabashada. Sidaa darteed, British Columbia, soo bixitaanka hidde-wadaha BC-2, BC-3, iyo BC-4 waxaa loo maleynayaa inay sabab u tahay isku-darka isku-dheelitirnaanta clones BC-1 iyo US-6 (Goodwin et al., 1995b). Waxaa suurtagal ah in noocyo isku dhafan laga helo dadweynaha Moscow. Tusaale ahaan, noocyada MO-4, MO-8 iyo MO-11 heterozygous ee loo yaqaan 'PEP locus' waxay noqon karaan kuwo isku dhafan oo u dhexeeya noocyada MO-12, MO-21, MO-22, oo leh nooca loo yaqaan 'A2 mating' iyo 'homozygous' oo loogu talagalay hal luuq oo ka mid ah aagga PEP iyo culeyska MO-8, oo leh nooca loo yaqaan 'A1 mating type' iyo 'homozygous' ee lidka kale ee aagga. Hadday sidaas tahayna, iyo dadka casriga ah ee P. infestans waxaa jira u janjeera in la kordhiyo doorka geedi socodka galmada, ka dib qiimaha macluumaadka ee falanqaynta xirmooyinka multilocus wuu yaraan doonaa (Elansky et al., 2001, 2015).
Kala duwanaanta khadadka isku xidhka
Ilaa 90-meeyadii qarnigii 20-aad, khadka khadadka ee US-1 wuxuu ku baahday adduunka. Inta badan goobaha iyo dadka ku nool gobollada waxay ka koobnaayeen noocyo noocyo leh nooca genotype ee US-1. Si kastaba ha noqotee, kala duwanaanshaha u dhexeeya go'doominta ayaa sidoo kale la arkay, oo ay u badan tahay inay sababtay geedi socodka isbadalka. Is-beddellada ayaa ka dhacay labada nukliyeer iyo mitochondrial DNA oo ay saameeyeen, iyo waxyaabo kale, heerka iska caabbinta daawooyinka phenylamide iyo tirada hiddo-wadayaasha feyraska. Khadadka ka duwan noocyada asalka ah ee asalka isbadalka waxaa lagu muujiyaa tirooyin dheeri ah ka dib dhibcaha ka dambeeya magaca genotype asalka ah (tusaale ahaan, US-1.1 line mutant ee xariiqa xargaha ee US-1). Khadadka DNA ee faraha faraha US-1.5 iyo US-1.6 waxay kakooban yihiin xariiqyo siyaado ah oo cabbirro kaladuwan leh (Goodwin et al., 1995a, 1995b); khadka isku xidhka US-6.3 sidoo kale wuxuu kaga duwan yahay US-6 hal xariiq oo siyaado ah (Goodwin, 1997, Shaxda 7).
Daraasadda mitochondrial DNA, waxaa la ogaaday in khadka klinikada US-1 kaliya nooca 1b ee mitochondrial DNA laga helo (Carter et al., 1990). Si kastaba ha noqotee, daraasad ku saabsan noocyada isirka ee ka soo jeeda Peru iyo Filibiin, go'doominta ayaa la helay kuwaas oo noocyada DNA ee mitochondrial ay kaga duwan yihiin 1b joogitaanka gelinta iyo tirtirka (Goodwin, 1991, Koh et al., 1994).
Jadwalka 6. Noocyada 'Multilocus genotypes' ee qaar ka mid ah P. infestans khadadka 'clonal'
Title | Nooca rimay | Isozymes | Faraha DNA | Nooca 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 |
Fiiro gaar ah: * - xog maleh.
Shaxda 7. Multilocus genotypes iyo khadadka is beddelaya
Title | Nooca rimay | | Faraha DNA (RG57) | Ogeysiis | |
GPI | PEP-1 | ||||
US-1 | A1 | 86/100 | 92/100 | 1011101011001101000110011 | Genotype asalka 1 |
US-1.1 | A1 | 86/100 | 100/100 | 1011101011001101000110011 | Is-beddel ku yimaada PEP |
US-1.2 | A1 | 86/100 | 92/100 | 1011101010001101000110011 | Is-beddelashada RG57 |
US-1.3 | A1 | 86/100 | 92/100 | 1011101001001101000110011 | Is-beddelashada RG57 |
US-1.4 | A1 | 86/100 | 100/100 | 1011101010001101000110011 | Is-beddelashada RG57 iyo PEP |
US-1.5 | A1 | 86/100 | 92/100 | 1011101011001101010110011 | Is-beddelashada RG57 |
US-6 | A1 | 100/100 | 92/100 | 1011111001001100010110011 | Genotype asalka 2 |
US-6.1 | A1 | 100/100 | 92 /92 | 1011111001001100010110011 | Is-beddel ku yimaada PEP |
US-6.2 | A1 | 100/100 | 92/100 | 1011101001001100010110011 | Is-beddelashada RG57 |
US-6.3 | A1 | 100/100 | 92/100 | 1011111001011100010110011 | Is-beddelashada RG57 |
US-6.4 | A1 | 100/100 | 100/100 | 1011011001001100010110011 | Is-beddelashada RG57 iyo PEP |
US-6.5 | A1 | 100/100 | 92/100 | 1011111001001100010010011 | Is-beddelashada RG57 |
BR-1 | A2 | 100/100 | 100/100 | 1011101000001100001111011 | Genotype asalka 3 |
BR-1.1 | A2 | 100/100 | 100/100 | 1010101000001100001110011 | Is-beddelashada RG57 |
Waxa kale oo jira isbeddelo ku saabsan muuqaalka 'isozymes'. Sida caadiga ah, waxaa sababa burburka noolaha markii horeba heterozygous ee loogu talagalay enzyme-kan loo yaqaan 'homozygous'. 1993, miraha yaanyada, waxaan ku ogaannay culeys leh astaamo lagu garto US-1: RG57 faraha, nooca mitochondrial DNA, iyo 86/100 genotype ee gulukoos-6-phosphati-isomerase, laakiin waxay ahayd homozygous (100/100) meesha ugu horeysa ee peptidase halkii laga heli lahaa a 92/100 heterozygote oo caadi u ah xariiqda 'clonal'. Waxaan ugu magac darnay nooca noocaan ah MO-17 (Shaxda 6). Khadadka mutant-ka US-1.1 iyo US-1.4 sidoo kale waxay kaga duwan yihiin US-1 isbadal ku yimaada meesha ugu horeysa ee peptidase (Sanduuqa 7).
Is-beddellada keena isbeddelka ku dhaca tirada hiddo-wadayaasha fircooniga ee noocyada baradhada iyo yaanyada waa wax caadi ah. Waxaa lagu xusay inay ka mid yihiin go'doominta khadadka isku xidhka US-1 ee dadweynaha Netherlands (Drenth et al., 1994), Peru (Goodwin et al., 1995a), Poland (Sujkowski et al., 1991), waqooyiga Ameerika (Goodwin et al., , 1995b). Kala duwanaanshaha tirada hiddo wadayaasha ee baradho ayaa sidoo kale lagu xusay ka go'doominta khadadka isku xidhka US-7 iyo US-8 ee Kanada iyo Mareykanka (Goodwin et al., 1995a), oo ka mid ah go'doominta khadka SIB-1 ee Aasiya qeybta Ruushka (Elansky et al, 2001) ).
Go'doominta kala duwanaanta adag ee heerarka iska caabbinta daawooyinka phenylamide waxaa lagu aqoonsaday dadka ku nool monoclonal, dhammaantoodna waxay ka tirsanaayeen xariiqda 'clonal line Sib-1' (Elansky et al, 2001, Table 1). Ku dhowaad dhammaan noocyada khadka dhexe ee US-1 aad ayey ugu nugul yihiin metalaxyl; si kastaba ha noqotee, go'doominnada aadka u adkaysta ee xariiqdan ayaa lagu go'doomiyay Filibiin (Koh et al., 1994) iyo Ireland (Goodwin et al., 1996).
Dadweynaha casriga ah ee P. infestans
Bartamaha Ameerika (Mexico)
Tirada dadka P. ee ku nool Mexico waxay si weyn uga duwan yihiin dadka kale ee adduunka, taas oo ugu horreyntii ay ugu wacan tahay booskeeda taariikhiga ah. Daraasado fara badan oo dadkan ah iyo kuwo laxiriira P. infestans noocyada clade Phytophthora, iyo sidoo kale noocyada maxalliga ah ee genus Solanum, ayaa horseeday gabagabada in isbeddelka cudur-sidaha qeybta dhexe ee Mexico ay wadajireen is-beddelka dhirta martida loo yahay oo lala xiriiriyay dib-isu-galmoodka (Grünwald, Flier) , 2005). Labada nooc ee isweheshiga waxaa matalaya dadweynaha, iyo si isku mid ah, iyo joogitaanka oospores ee carrada, dhirta iyo digriiga baradhada iyo noocyada la xiriira ee duurjoogta ee Solanum waxay xaqiijineysaa jiritaanka geedi socodka galmada ee dadweynaha (Fernández-Pavía et al., 2002). Daraasadihii ugu dambeeyay ee Dooxada Toluca iyo hareeraheeda (xarunta loo maleynayo inuu asal ahaan ka soo jeedo cudur-sidaha) ayaa xaqiijiyay kala duwanaanta hidde-wadaha sare ee dadka deegaanka ee P. infestans (134 multilocus genotypes ee muunad ka mid ah 176 shaybaar) iyo joogitaanka dad badan oo ku kala duwan gobolka (Wang et al., 2017). Waxyaabaha keenaya kala duwanaanshahani waa kala qaybsanaanta goobaha dadyowga lagu garto aagagga sare ee badhtamaha Mexico, kala duwanaanta xaaladaha beerashada iyo noocyada baradhada ee loo isticmaalo dooxooyinka iyo buuraha, iyo joogitaanka noocyada duurjoogta ee Solanum ee u dhaqmi kara sidii martida kale (Fry et al. , 2009).
Si kastaba ha noqotee, waa in la ogaadaa in dadweynaha P. infestans ee waqooyiga Mexico ay yihiin kuwo isku mid ah oo aad ula mid ah dadka Waqooyiga Ameerika, taas oo laga yaabo inay muujiso in kuwani ay yihiin noocyada cusub (Fry et al., 2009).
Woqooyiga Ameerika
Dadka ku nool Waqooyiga Ameerika ee P. infestans waxay had iyo jeer lahaayeen qaab dhismeed aad u fudud oo dabeecadooda isku dhafan ayaa la aasaasay muddo dheer ka hor isticmaalka falanqaynta mikrosatellite. Ilaa 1987, khadka khadadka ee US-1 ayaa ku xoog badnaa Mareykanka iyo Kanada (Goodwin et al., 1995). Bartamihii-70-yadii, markii ay muuqatay fungicides-ku saleysan metalaxyl, midabkani wuxuu bilaabay in lagu beddelo noocyo kale oo adkaysi badan oo ka soo hayaamay Mexico (Goodwin et al., 1998). Dhamaadkii 90-maadkii. genotype-ka US-8 wuxuu gabi ahaanba bedelay genotype-ka Mareykanka-1 ee Mareykanka wuxuuna noqday khadka ugu weyn ee baradhada (Fry et al., 2009; Fry et al., 2015). Xaaladda way ka duwaneyd yaanyada, oo had iyo jeer ka kooban dhowr xariiqyo klinik ah, halabuurkoodana sanadba sanadka ka dambeeya wuu is beddelaa (Fry et al., 2009).
Sannadkii 2009-kii, faafitaan baaxad weyn oo goor dambe ku dhacay yaanyada ayaa ka dilaacay Mareykanka. Muuqaalka cudurkan faafa wuxuu ahaa ku dhowaad isku mar inuu ka billowdo meelo badan oo woqooyi-bari Mareykanka ah, waxayna u noqotay inuu la xiriiro iibinta ballaaran ee geedo yaanyo qaba cudurka ee xarumaha beeraha waaweyn (Fry et al., 2013). Khasaaraha dalagyada ayaa aad u badnaa. Falanqaynta Microsatellite ee shaybaarada ay dhibaatadu saameysey ayaa muujisay in masiibada aafada ah ay leedahay khadka isku xidhka US-22 A2. In 2009, saamiga genotype this ee dadka Mareykanka ee P. infestans ayaa gaadhay 80% (Fry et al., 2013). Sannadaha soo socda, saamiga noocyada dagaal ee US-23 (badanaa yaanyada) iyo US-24 (baradhada) si isdaba joog ah ayey ugu kordheen dadka, hase yeeshe, wixii ka dambeeyay 2011, heerka ogaanshaha ee US-24 aad ayuu hoos ugu dhacay, ilaa maantana, qiyaastii 90% ee dadka cudurka keena. Mareykanka waxaa matalaya genotype US-23 (Fry et al., 2015).
Kanada gudaheeda, sida Mareykanka oo kale, dhammaadkii 90-meeyadii. genotype-kii ugu weynaa ee US-1 waxaa beddelay US-8, boosaska ugu waaweyn ee aan is beddelin illaa 2008. Kanada gudaheeda, waxaa ka jiray cudurro halis ah oo xilli dambe ah oo la xiriira iibinta geedo yaanyo qaba, laakiin waxaa sababay genotypes US-2009 iyo US-2010 (Kalischuk et al., 23). Kala duwanaanshaha juqraafi ahaaneed ee genotypes-ka ayaa ahaa mid cajiib ah: US-8 wuxuu xukumayay gobollada galbeedka Kanada (2012%), halka US-23 uu xukumayay gobollada bariga (68%). Sannadihii xigay, US-8 wuxuu ku faafay gobollada bari; si kastaba ha noqotee, guud ahaan, saamiga ay ku leeyihiin dadka ayaa xoogaa hoos uga dhacay asalka muuqaalka genotypes US-83 iyo US-23 ee dalka (Peters et al., 22) Ilaa hadda, US-24 waxay ku sii jirtaa booska xukunka Kanada oo dhan; US-2014 wuxuu joogaa British Columbia, halka US-23 iyo US-8 ay joogaan Ontario (Peters, 23).
Marka, dadweynaha Waqooyiga Ameerika ee P. infestans badanaa waa khadadka isku xidhka. 40-kii sano ee la soo dhaafay, tirada la ogaadey genotypes clonal-ka ayaa gaartey 24. In kasta oo xaqiiqda ah in noocyada labada nooc ee isutagga ay ku dhex jiraan dadka, suurtagalnimada muuqaalka hiddo-wadayaasha cusub ee ka dhalanaya isu-keenista galmada ayaa weli aad u hooseysa. Si kastaba ha noqotee, 20-kii sano ee la soo dhaafay, dhowr kiis oo ka mid ah muuqaalka dadka rephebinant recombinant ayaa la duubay (Gavino et al., 2000; Danies et al., 2014; Peters et al., 2014), iyo hal kiis, natiijada isgoyska waxay ahayd genotype US-11 , kaas oo ku xididaystay Waqooyiga Ameerika sanado badan (Gavino et al., 2000). Ilaa iyo 2009, isbeddelada ku yimid qaab dhismeedka dadka waxay la xiriireen soo bixitaan cusub, hiddo-wadayaal dagaal badan oo la socda socdaalkooda iyo barokicinta kuwii hore ee xoogga badnaa. Maxaa dhacay sanadihii 2009-2010 USA iyo Kanada, epiphytotics markii ugu horreysay waxay muujiyeen in xilligii caalamiyeynta, faafida cudurku la xiriiri karo faafitaanka firfircoon ee genotypes cusub marka la iibinayo waxyaabaha dhirtu ku jirto.
South America
Ilaa dhowaan, daraasadaha dadka ku nool Koonfurta Ameerika ee P. infestans ma ahayn kuwo joogto ah ama ballaaran. Waxaa la ogyahay in qaab dhismeedka dadkani uu yahay mid aad u fudud oo ay ku jiraan 1-5 nasab isku xidhka wadankiiba (Forbes et al., 1998). Marka, 1998, genotypes US-1 (Brazil, Chile) BR-1 (Brazil, Bolivia, Uruguay, Paraguay), EC-1 (Ecuador, Colombia, Peru iyo Venezuela), AR-1, AR waxaa laga helay baradhada -2, AR-3, AR-4 iyo AR-5 (Argentina), PE-3 iyo PE-7 (koonfurta Peru). Nooca Naas-nuujinta A2 wuxuu ku sugnaa Brazil, Bolivia iyo Argentina lagamana helin meel ka baxsan xadka Bolivian-Peruvian ee aagga harada Lake Titicaca, oo gadaashiisa ay EC-1 A1 genotype ku badan tahay Andes. Yaanyada, US-1 waxay ku sii jirtey genotype ku baahsan Koonfurta Ameerika.
Xaaladda in badan ama ka yar ayaa sii jirtey 2000s. Qodob muhiim ah ayaa ahaa helitaanka Waqooyiga Andes ehelka baradhada duurjoogta ah ee soo koraya (S. brevifolium iyo S. tetrapetalum) ee xariiqa cusub ee loo yaqaan 'clonal line EC-2' ee nooca A2 (Oliva et al., 2010). Daraasadaha cilmiga jirka waxay muujiyeen in xariiqan uusan gebi ahaanba la mid ahayn P. infestans, in kasta oo uu xiriir dhow la leeyahay, xiriir la leh kaas oo loo soo jeediyay in laga fiirsado, iyo sidoo kale xarriiq kale, EC-3, oo laga soocay geedka yaanyada S. betaceum ee ku koraya Andes, nooc cusub oo la yiraahdo P. andina; si kastaba ha noqotee, xaaladda noocyadan (nooc madax-bannaan ama isku-dhaf ah P. infestans oo leh khad aan weli la aqoon) weli ma cadda (Delgado et al., 2013).
Waqtigan xaadirka ah, dhammaan dadka Koonfurta Ameerika ee P. infestans waa isku mid. In kasta oo ay jiraan labada nooc ee isutagga, haddana lama oga tirada dadka dib isugu noqda. Yaanyada, gen-ka US-1 waa mid meel walba yaal, oo sida muuqata ka soo bara kacay baradhada noocyada maxalliga ah, asalka dhabta ah ee aan weli la aqoon. Waddamada Brazil, Bolivia iyo Uruguay, nooca loo yaqaan 'BR-1 genotype' ayaa jooga; gudaha Peru, oo ay weheliyaan US-1 iyo EC-1, waxaa jira dhowr nooc oo genotypes maxalli ah. Andes-ka, booska ugu sarreeya waxaa ku sii jira khadka xariiqa EC-1, xiriirka uu la leeyahay P. andina dhowaan la ogaaday weli lama sahamin. Meesha kaliya "aan xasilloonayn" ee loogu talagalay muddada 2003-2013. waxaa jiray isbeddello muhiim ah oo ku yimid dadka, oo noqday Chile (Acuña et al., 2012), halkaas oo 2004-2005. dadka cudurada keena waxay ku sifoobeen u adkaysiga metalaxyl iyo haptotype cusub oo mitochondrial DNA ah (Ia halkii Ib hore looga heli lahaa). 2006 ilaa 2011 Dadweynaha, genotype 21 (sida ku cad SSR) ayaa xukumay, qaybta ay gaareen 90%, intaas ka dib calaacasha waxay u gudubtay genotype 20, inta jeer ee ay dhacdo labada sano ee soo socota waxaa lagu hayaa qiyaastii 67% (Acuña, 2015).
Yurub
Taariikhda Yurub, waxaa jiray ugu yaraan laba mowjadood oo guuritaan ah oo dadka P. ay ka soo guureen Waqooyiga Ameerika: qarnigii 1aad. (HERB-1) iyo horaantii qarnigii 70aad (US-1). Qeybinta meel walba ee fangaska lagu buuxiyo ee metalaxyl XNUMXs. u horseeday barakicinta hidde-wadaha weyn ee US-XNUMX laguna beddelay genotypes cusub. Natiijo ahaan, inta badan waddamada Galbeedka Yurub, dadyowga cudur-sidaha waxaa metelayay inta badan khadado isku mid ah.
Adeegsiga falanqaynta mikrosatelite-ka falanqaynta ee dadka cudurada keena ayaa suurta galisay in la aqoonsado isbadalada halista ah ee ka dhacay Galbeedka Yurub sanadihii 2005-2008. 2005, UK gudaheeda ayaa laga helay khad cusub oo loo yaqaan 'clonal line', kaas oo loo yaqaan '13_A2 (ama "Blue 13") oo lagu garto nooca A2 ee isu geynta , gardarro sare iyo iska caabbinta phenylamides (Shaw et al., 2007). Isla genotype ayaa laga helay shaybaarro laga soo aruuriyay 2004tii Nederland iyo waqooyiga Faransiiska, taasoo soo jeedinaysa inay u guurtay Boqortooyada Midowday ee Yurub, iyadoo laga yaabo inay ku jirto baradho abuur (Cooke et al., 2007). Daraasada genome ee wakiilada khadkan cnal waxay muujisay heer sare oo polymorphism ah oo isku xigxiga (2016, tirada kala duwanaanshaheeda subclonal-ka waxay gaadhay 340) iyo darajo weyn oo kala duwanaansho heerka muujinta hidda-wadaha, incl. hiddo-wadaha waxyeellada inta lagu jiro infekshanka dhirta (Cooke et al., 2012; Cooke, 2017). Astaamahan, oo ay weheliso muddada sii kordheysa ee wajiga biotrophic, waxay sababi kartaa dagaal xoog leh oo ah 13_A2 iyo awoodda ay u leedahay in ay qaadsiiso xitaa noocyada baradhada ee u adkaysata aafada dambe.
Sanadaha soo socda, genotype wuxuu si dhaqso leh ugu faafay wadamada Waqooyiga-galbeed ee Yurub (Great Britain, Ireland, France, Belgium, Netherlands, Germany) iyadoo isla mar ahaantaana la barakicinayo noocyada genotypes ee horay u jiray 1_A1, 2_A1, 8_A1 (Montarry et al., 2010; Gisi et al. , 2011; Van den Bosch et al., 2011; Cooke, 2015; Cooke, 2017). Sida laga soo xigtay shabakada www.euroblight.net, saamiga 13_A2 ee dadweynaha dalalkan ayaa gaadhay 60-80% iyo in ka badan; jiritaanka genotype-kan ayaa sidoo kale laga diiwaan geliyay qaar ka mid ah wadamada Bariga iyo Koonfurta Yurub. Si kastaba ha noqotee, 2009-2012. 13_A2 wuxuu lumiyay booskiisii ugu weynaa ee Great Britain iyo France, isagoo udhaxeeya khadka 6_A1 (8_A1 ee Ireland), Nederland iyo Beljigana qeyb ahaan waxaa lagu badalay genotypes 1_A1, 6_A1, iyo 33_A2 (Cooke et al., 2012; Cooke, 2017; Stellingwerf, 2017).
Ilaa hadda, qiyaastii 70% dadka reer Yurubta Galbeed ee P. infestans waa monoclonal. Sida laga soo xigtay websaydhka www.euroblight.net, genotypes-ka ugu badan ee dalalka Waqooyiga-Galbeedka Yurub (Boqortooyada Ingiriiska, Faransiiska,
Nederland, Beljim) weli, qiyaas ahaan waa loo siman yahay, 13_A2 iyo 6_A1, tan dambe ficil ahaan lagama helin meel ka baxsan gobolka la cayimay (marka laga reebo Ireland), laakiin horeyba ugu yaraan 58 subclones (Cooke, 2017). Kala duwanaanshaha 13_A2 ayaa ku jira tirooyin la ogaan karo oo ku yaal Jarmalka, waxaana sidoo kale si goos goos ah loogu arkaa waddamada Bartamaha iyo Koonfurta Yurub. Genotype 1_A1 wuxuu qayb weyn ka yahay dadka reer Beljam iyo qayb ahaan Nederland iyo Faransiiska. Genotype 8_A1 wuxuu ku xasillay dadka reer Yurub heerka 3-6%, marka laga reebo Ireland, halkaasoo ay ku sii heysato booskeeda hogaamineed waxayna u qeybsan tahay laba subclones (Stellingwerf, 2017). Ugu dambeyntiina, sanadkii 2016, kororka soo noqnoqoshada dhacdooyinka genotypes cusub 36_A2 iyo 37_A2, oo markii ugu horreysay la diiwaangeliyey 2013-2014, ayaa lagu xusay; ilaa maanta, genotypes-kan waxaa laga helaa Nederland iyo Beljam iyo qayb ahaan Faransiiska iyo Jarmalka, iyo sidoo kale qaybta koonfureed ee Great Britain (Cooke, 2017). Qiyaastii 20-30% dadka reer Yurubta Galbeed waxaa matala genotypes gaar ah sanad kasta.
Si ka duwan Galbeedka Yurub, markii ay muuqatay nooca '13_A2 genotype', dadka Waqooyiga Yurub (Sweden, Norway, Denmark, Finland) waxaa matalayay khadadka isku xidhka, laakiin tiro badan oo noocyo gaar ah (Brurberg et al.,
2011). Intii lagu gudajiray muddadii faafitaanka firfircoonaa ee 13_A2 ee Galbeedka Yurub, joogitaanka genotype-kan Scandinavia lama ogaanin illaa 2011, markii ugu horreysay laga helay woqooyiga Jylland (Danmark), halkaas oo inta badan noocyada baradhada warshadaha lagu beero iyadoo si firfircoon loo isticmaalo metalaxyl-ka fungicides (Nielsen et al., 2014). Sida laga soo xigtay www.euroblight.net, genotype 13_A2 ayaa sidoo kale laga helay dhowr tijaabo oo laga keenay Norway iyo Danmark sanadkii 2014 iyo dhowr tijaabo oo norwiiji ah sanadkii 2016; waxaa intaa dheer, sanadkii 2013 gudaha Finland joogitaanka genotype 6_A1 ayaa lagu xusay qadar yar. Sababta ugu weyn ee fashilka 13_A2 iyo khadadka kale ee kiliinikada ee qabsashadii Iskandaneefiyanka ayaa loo arkaa inay tahay kala duwanaanshaha cimilada ee gobolkan ee ka imanaya waddamada Galbeedka Yurub.
Marka lagu daro xaqiiqda ah in xagaaga qabow iyo jiilaalka qabow ay kor u qaadaan jiritaanka dhirta aan aadka u badnayn ee loo yaqaan 'oospores' (Sjöholm et al., 2013), qaboojinta carrada xilliga qaboobaha (oo badanaa aan ka dhicin waddamada diirran ee Galbeedka Yurub) waxay gacan ka geysaneysaa is-waafajinta biqilka dhirta iyo abuurka. baradhada, taas oo kor u qaadeysa kaalinteeda sida ilaha cudurka aasaasiga ah (Brurberg et al., 2011). Sidoo kale waa in la ogaadaa in, xaaladaha waqooyiga, horumarka infekshinka ka yimaada oospores ay kor u dhaafayso horumarinta cudurka infekshinka, taas oo ugu dambeyntii ka hortageysa xukunka xitaa ka sii daran, laakiin markii dambe la sameeyay khadadka isku xirnaanta (Yuen, 2012). Qaab dhismeedka dadka ugu daraasadda badan ee loo yaqaan 'P. infestans' ee dalalka Bariga Yurub (Poland, gobollada Baltiga) wuxuu aad ugu eg yahay kan Scandinavia.
Labada nooc ee isweydaarsiga ayaa sidoo kale halkan jooga, iyo inta badan noocyada genotypes ee lagu go'aamiyo falanqaynta SSR waa kuwo gaar ah (Chmielarz et al., 2014; Runno-Paurson et al., 2016). Sida Waqooyiga Yurub, qaybinta khadadka isku xidhka (ugu horreyntii 13_A2 genotype) ficil ahaan saameyn kuma yeelan dadka maxalliga ah ee cudur-sidaha, kuwaas oo xajiya kala-duwanaansho aad u sarreeya iyadoo maqnaanshaha khadadka lagu dhawaaqo.
Joogitaanka 13_A2 ayaa mararka qaarkood lagu arkaa beeraha leh noocyada baradhada ganacsiga. Ruushka dhexdiisa, xaaladdu si la mid ah ayey u horumaraysaa. Falanqaynta Microsatellite ee P. infestans go'doominta la uruuriyay 2008-2011 10 gobol oo kala duwan oo ka tirsan qaybta Yurub ee Ruushka, waxay muujiyeen heer sare oo kala duwanaanshaha hidde-wadaha ah iyo la'aanta dhammaystirnaanta isku-xidhka khadadka midowga Yurub (Statsyuk et al., 2014). Dhowr sano ka dib, daraasad ku saabsan shaybaarada P. infestans ee laga soo ururiyay gobolka Leningrad ee 2013-2014 ayaa muujisay farqi weyn oo u dhexeeya iyaga iyo genotypes-ka gobolkan lagu aqoonsaday daraasaddii hore. Labada daraasadoodba, looma helin genotypes Yurubta galbeed (Beketova et al., 2014; Kuznetsova et al., 2016).
Kala duwanaanta hidde-wadaha sare ee dadweynaha Bariga Yurub ee P. infestans iyo maqnaanshaha khadadka cnal ee ugu weyn iyaga ayaa sababi kara dhowr sababood. Marka hore, sida Waqooyiga Yurub, xaaladaha cimilada ee waddammada la tixgeliyo waxay gacan ka geystaan sameynta oospores oo ah isha koowaad ee cudurka (Ulanova et al., 2010; Chmielarz et al., 2014). Marka labaad, qayb muhiim ah oo baradho ah oo laga soo saaray dalalkan ayaa lagu beeray beeraha yaryar ee gaarka loo leeyahay, oo badanaa ku hareeraysan kaymo ama caqabado kale oo dhaqdhaqaaqa xorta ah ee walxaha faafa (Chmielarz et al., 2014). Sida caadiga ah, baradhada lagu beero xaaladaha noocaas ah ficil ahaan looma daaweeyo kiimikooyin, xulashada noocyada waxay ku saleysan tahay iska caabbintooda goor dambe ee cudurka, i.e. ma jiro cadaadis la xushay oo ku saabsan gardarrada iyo iska caabinta metalaxyl, taas oo xannibaysa hidde-wadayaasha adkaysi u leh, sida 13_A2, faa'iidooyinka ay ka leeyihiin genotypes kale (Chmielarz et al., 2014). Ugu dambeyntiina, cabirka yar ee dhulal dhul awgood, milkiilayaashooda sida caadiga ah kuma dhaqmaan wareegga dalagga, baradhada ku koraya isla goobtaas muddo sannado ah, taas oo gacan ka geysaneysa isku soo aruurinta inoculum hiddo ahaan kala duwan (Runno-Paurson et al., 2016; Elansky, 2015; Elansky et al. , 2015).
Aasiya
Ilaa dhowaan, qaab dhismeedka P. infestans ee ku nool Aasiya weli si hoose looma fahmin. Waxaa la ogaa in inta badan ay matalaan khadadka loo yaqaan 'clonal lines', iyo saameynta isku soo noqoshada galmada ee soo bixitaanka hiddo-wadaha cusubi waa mid aad u yar. Marka, tusaale ahaan, 1997-1998. Qeybta Aasiya ee Ruushka (Siberia iyo Bariga Fog), dadka cudurada keena waxaa matalayay kaliya seddex nooc oo genotypes ah oo u badan SIB-1 genotype (Elansky et al., 2001). Joogitaanka khadadka cudurada wadnaha ayaa lagu muujiyay wadamada sida Shiinaha, Japan, Korea, Filibiin, iyo Taiwan (Koh et al., 1994; Chen et al., 2009). Khadka isku xidhka US-1 wuxuu ku xoog badnaa dhul ballaaran oo Aasiya ah dabayaaqadii 90-meeyadii - horraantii 2000-meeyadii. ku dhowaad meel kasta waxay bilaabeen in lagu beddelo hiddo-wadaha kale, kuwaas oo, iyagana, u beddelay kuwa cusub. Xaaladaha badankood, isbeddelada qaabdhismeedka iyo qaab dhismeedka dadka ku nool waddamada Aasiya waxay la xiriireen u guuritaanka genotypes cusub oo ka yimid dibedda. Marka, Japan, marka laga reebo genotype JP-3, dhammaan noocyada kale ee Jabaan ee soo muuqday US-1 kadib (JP-1, JP-2, JP-3) waxay leeyihiin asal dibadeed oo ka badan ama ka yar (Akino et al., 2011) ... Waqtigan xaadirka ah Shiinaha waxaa ku nool saddex qof oo ugu badan ee cudurada keena, kuwaas oo leh kala qaybsanaan juquraafi ahaan; Ma jiro ama qulqulka hidde-wadaha aadka u daciifka ah ee u dhexeeya dadkan (Guo et al., 2010; Li et al., 2013b). Genotype 13_A2 ayaa ka soo muuqday dhulka Shiinaha ee gobollada koonfureed (Yunnan iyo Sichuan) 2005-2007, iyo 2012-1014. sidoo kale waxaa lagu arkay waqooyi-bari dalka (Li et al., 2013b). Hindiya gudaheeda, 13_A2 waxay u muuqatay inay macquul tahay isla waqtigaas Shiinaha, waxayna u badan tahay baradhada iniinyaha qaba (Chowdappa et al., 2015), iyo 2009-2010. wuxuu sababay epiphytosis daran oo ah duqeyn goor dambe ku dhacday yaanyada koonfurta dalka, kadib markuu ku faafay baradhada sanadkii 2014 wuxuu sababay dillaac soo daahay West Bengal, taasoo horseedday burburka iyo isdilka beeraley badan oo deegaanka ah (Fry, 2016).
Afrika
Ilaa 2008-2010 daraasad nidaamsan oo ku saabsan P. infestans ee dalalka Afrika lama fulin. Xilligan la joogo, dadka Afrikaanka ah ee loo yaqaan 'P. infestans' waxaa loo qaybin karaa laba kooxood, qaybtani waxay si cad ula xiriirtaa xaqiiqda soo dejinta baradhada abuurka ee Yurub.
Waqooyiga Afrika, oo si firfircoon uga soo dhoofisa baradhada abuurka Yurub, nooca A2 nooca mating ayaa si ballaaran loogu metelaa ku dhowaad dhammaan gobollada, taas oo bixisa suurtagal aragti ahaaneed oo ah soo bixitaanka hiddo-wadaha cusub ee ka yimaada dib-isu-galmoodka galmada (Corbière et al., 2010; Rekad et al., 2017). Intaa waxaa dheer, Algeria, joogitaanka genotypes 13_A2, 2_A1, iyo 23_A1 waxaa lagu xusay xukun xoog leh oo kii ugu horreeyay ka mid ah, iyo sidoo kale hoos u dhac tartiib tartiib ah oo ku yimid saamiga genotypes gaar ah illaa dhammaad la waayo (Rekad et al., 2017). Marka la barbardhigo gobolka intiisa kale, Tuunis (marka laga reebo woqooyi-bari ee waddanka), dadka cudurada keena waxaa inta badan matala nooca A1 mating (Harbaoui et al., 2014).
Khadadka isku xidhka NA-01 ayaa halkan ku xoog badan. Guud ahaan, saamiga khadadka isku-xidhka 'clonal' ee dadweynaha waa kaliya 43%. Bariga iyo Koonfurta Afrika, halkaasoo mugga soo dejinta abuurka ay aad u yar yihiin (Fry et al., 2009), P. infestans waxaa matalaya kaliya laba xariiq oo nooca A1 ah, US-1 iyo KE-1, kan dambena wuxuu si firfircoon u barakicinayaa kii hore ee baradhada ( Pule et al., 2012; Njoroge et al., 2016). Ilaa hadda, labada nooc ee genotypes waxay leeyihiin tiro muuqata oo kala duwanaansho subclonal ah.
Australia
Warbixintii ugu horreysay ee ku saabsan baradhada soo daahay ee baradhada ee Australiya waxay soo jirtay illaa 1907, iyo epiphytotia-kii ugu horreeyay, oo loo maleynayo inay sababeen roobab culus bilaha xagaaga, waxay dhaceen 1909-1911. (Drenth et al., 2002). Guud ahaan, si kastaba ha ahaatee, duqeynta goor dambe wax macno leh oo dhaqaale kuma leh dalka. Dilaac goos goos ah oo goor dambe ah, oo ay kiciso xaaladaha cimilada ee bixiya huurka badan, ma dhacaan in ka badan hal jeer 5-7-dii sanoba mar waxaana laga helaa inta badan waqooyiga Tasmania iyo bartamaha Victoria. Iyadoo lala xiriirinayo kor ku xusan, daabacaadaha loo qoondeeyay barashada qaab-dhismeedka dadweynaha Australiyaanka ah ee P. infestans ficil ahaan ma maqna. Macluumaadka ugu dambeeya ee la heli karo waa 1998-2000. (Drenth et al., 2002). Sida ay qoreen qorayaashu, dadweynaha gobolka Victoria waxay ahaayeen nasab isku xidha US-1.3, oo si dadban u xaqiijinaya u guuritaanka genotype-kani Maraykanka. Tijaabooyinka Tasmania waxaa loo kala saaray inay yihiin AU-3, oo ka duwan noocyada genotypes ee xilligaas ka jiray qaybaha kale ee adduunka.
Noocyada horumarinta duqayn dambe ee Russia
Yurub gudaheeda, infekshinka waxaa lagu soo qaadaa digriiga abuurka buka, oospores oo hareeyay carrada, iyo sidoo kale zoosporangia oo ay dabaysha ka keento dhir ka soo baxday digriiga hareeraha ah ee beeraha sanadkii la soo dhaafay ("iskaa wax u qabso" dhirta), ama tuulmooyin la soo saaray calaamadee kaydinta digriiga. Kuwaas oo ka mid ah, dhirta ku koray tuulbooyinkii digriiga la tuuray ayaa loo tixgeliyaa inay yihiin isha ugu halista badan ee infekshanka. tirada digriiga magooshay waxaa jira inta badan mid muhiim ah, iyo zoosporangia ayaa looga qaadi karaa iyaga masaafo dheer. Inta kale ee ilaha (oospores, "iskaa wax u qabso" dhirta) ma aha kuwo khatar badan, maxaa yeelay caado uma aha in dhir lagu beero isla beeraha in ka badan marar badan 3-4 sanoba mar. Caabuqa ka soo baxa digriiga abuurka buka ayaa sidoo kale ugu yar sababtoo ah nidaamka xakamaynta tayada abuurka oo wanaagsan.
Guud ahaan, qaddarka tallaalka ee dadka reer Yurub way xadidan yihiin, sidaa darteedna koritaanka faafa waa mid gaabis ah oo si guul leh ayaa loo xakameyn karaa iyadoo la adeegsanayo diyaarinta fungicidal kiimikada. Hawsha ugu weyn ee xaaladaha yurubiyanku waa la dagaallanka infekshinka wejiga markii kala-firdhinta ballaaran ee zoosporangia ee dhirta ay saamaysay bilaabmayso.
Ruushka, xaaladdu si aad ah ayey uga duwan tahay. Inta badan dalagga baradhada iyo yaanyada waxaa lagu beero jardiinooyin yaryar oo gaar loo leeyahay; talaabooyinka kahortaga ah ama laguma fulin iyaga gebi ahaanba, ama daaweynta fungicidal-ka waxaa lagu sameeyaa tiro aan ku filnayn waxayna bilaabaan ka dib marka ay soo ifbaxaan nabar soo daahay dushooda. Natiija ahaan, jardiinooyinka khudradda ee gaarka loo leeyahay waxay u dhaqmaan sida ilaha ugu weyn ee infekshanka, kaas oo xayawaannada loo yaqaan 'zoosporangia' ay dabayshu u qaaddo ilaa ay ku beero ganacsi. Tan waxaa lagu xaqiijiyay kormeerkeena tooska ah ee gobolada Moscow, Bryansk, Kostroma, Ryazan: waxyeelada soo gaadhay dhirta jardiinooyinka gaarka loo leeyahay ayaa la arkay xitaa kahor bilawga daaweynta fungicide ee beerashada ganacsiga. Intaas ka dib, cudurka faafa ee beeraha waaweyn waxaa lagu xakameynayaa adeegsiga diyaargarowga fungicidal, halka jardiinooyinka gaarka loo leeyahay ay si deg deg ah uga soo baxayaan goor dambe.
Xaaladda daaweynta khaldan ama "miisaaniyadeed" ee beerashada ganacsiga, ujeeddooyinka duqeynta goor dambe ayaa sidoo kale ka soo muuqanaya beeraha; ka dib waxay si firfircoon u horumarinayaan, iyagoo daboolaya aagag weligood ka ballaaran (Elansky, 2015). Infekshanka ku dhaca jardiinooyinka gaarka loo leeyahay ayaa saameyn weyn ku leh cudurrada faafa ee goobaha ganacsiga. Dhammaan gobollada baradhada ka baxa ee Ruushka, aagga ay baradho ku leedahay jardiinooyinka gaarka loo leeyahay waxay dhowr jeer ka weyn yihiin aagga guud ee beeraha wax soo saarayaasha waaweyn. Jawiga noocan oo kale ah, jardiinooyinka khudradda ee gaarka loo leeyahay ayaa loo arki karaa inay yihiin ilaha waxbarasho ee caalamiga ah ee goobaha ganacsiga. Aynu isku dayno inaan aqoonsanno astaamaha astaamaha lagu garto noocyada kala duwan ee jardiinooyinka gaarka loo leeyahay.
Beerista abuurka iyo xakameynta karantiilida baradhada, miraha yaanyada ee laga helo soosaarayaasha ajaanibta ah ee shakiga leh, beerashada mudada dheer ee baradhada iyo yaanyada isla aagagga, daaweynta fangaska ee aan habooneyn ama maqnaashahooda oo dhameystiran waxay keeneysaa epiphytoties daran oo ku saabsan waaxda gaarka loo leeyahay, natiijadeeduna waa lacag isgoysyada, iskudhafka iyo samaynta oospore ee jardiinooyinka gaarka loo leeyahay. Natiijo ahaan, kala duwanaansho hiddo-wadaha oo aad u sarreeya oo loo yaqaan 'pathogen' ayaa la arkaa, marka ku dhowaad culeys kasta uu gaar u yahay genotype (Elansky et al., 2001, 2015). Abuuritaanka baradho abuur ah asal ahaan hidde-sideyaal kala duwan waxay ka dhigeysaa wax aan macquul aheyn in khadadka isku-dhafan ee ku takhasusay inay weeraraan noocyo gaar ah ay soo baxaan. Noocyada lagu xushay xaaladdan oo kale waxaa lagu kala soocayaa kala duwanaanshahooda la xiriira noocyada ay dhibaatadu saameysey, badankood waxay ku dhowyihiin tirada ugu badan ee hiddo-wadayaasha feyraska. Tani aad ayey uga duwan tahay nidaamka "khadadka kiliinikada" ee caadiga u ah beeraha waaweyn ee shirkadaha beeraha oo leh nidaam si sax ah loo rakibay oo ka hortag ah duqeynta goor dambe. Khadadka loo yaqaan 'Clonal khadadka' (marka dhammaan noocyada cudurka dabaysha dambe ee aagga ku jira ay metelaan hal ama in ka badan hiddo-wadaha) ayaa ku baahsan dalal ay ka baxayaan baradhada gaar ahaan beeraha waaweyn: USA, Netherlands, Denmark, iwm. England, Ireland, Poland, halkaas oo guryooyinka guryuhu sidoo kale dhaqan ahaan ku faafaan. baradhada sii kordheysa, sidoo kale waxaa jira kala duwanaansho hiddo-wadaha sare ee jardiinooyinka gaarka loo leeyahay. Dhamaadkii qarnigii 20-aad, "khadadka loo yaqaan" clonal lines "waxay ku faafeen qeybaha Aasiya iyo Bariga Fog ee Ruushka (Elansky et al., 2001), taas oo sida muuqata sabab u ah isticmaalka noocyo isku mid ah oo baradho ah oo si gaar ah loo beero. Dhawaan, xaaladda gobolladaan sidoo kale waxay bilaabeen inay u beddelaan dhinaca kororka noocyada kala duwan ee genotypic ee dadweynaha.
La'aanta daaweyn deg deg ah oo lala yeesho diyaarinta fungicidal ayaa leh cawaaqib kale oo toos ah - ma jiraan uruurinta noocyo adkaysi u leh jardiinooyinka. Runtii, natiijooyinkayagu waxay muujinayaan in noocyada u adkaysta metalaxyl laga helo meelo aad uga yar inta badan beeraha gaarka loo leeyahay marka loo eego beeraha ganacsiga.
U dhawaanshaha baradhada iyo yaanyada la beero, ee caadiga u ah beeraha gaarka loo leeyahay, waxay fududeeysaa socdaalka noocyada u dhexeeya dalagyadan, taas oo ka dhalatay, tobankii sano ee la soo dhaafay, oo ka mid ah noocyada laga soocay baradhada, saamiga noocyada noocyada ee qaada hidda-wadaha iska caabbinta noocyada yaanyada cherry (T1), oo hore loogu yaqaanay kaliya " yaanyo "nooc. Dhibaatooyinka hiddo-wadaha T1 xaaladaha badankood waxay aad ugu dagaallamaan baradhada iyo yaanyada labadaba.
Sanadihii la soo dhaafay, duqeynta goor dambe ee yaanyada waxay bilaabatay inay ka muuqato xaalado badan oo ka horreeyay baradhada. Geedo yaanyo waxaa ku dhici kara oogyada carrada ku jirta, ama hoormoonka ay ku jiraan miraha yaanyada ama u hoggaansamaya iyaga (Rubin et al., 2001). 15-kii sano ee la soo dhaafay, tiro fara badan oo ah miraha la soo baakadeeyay oo qaali ah, oo inta badan la soo dhoofiyo, ayaa ka soo muuqday dukaamada, badankoodna soo saarayaasha yar yar ayaa u wareegay isticmaalkooda. Abuurku wuxuu ka koobnaan karaa noocyo noocyo hidde ah oo ka mid ah gobollada ay wax ku beertaan. Mustaqbalka, genotypes-kan waxaa lagu daraa habka galmada ee jardiinooyinka gaarka loo leeyahay, taas oo horseedaysa soo bixitaanka genotypes gabi ahaanba cusub.
Sidaa darteed, waa la sheegi karaa in jardiinooyinka gaarka loo leeyahay ay yihiin "dunid dhalaasho" oo adduunka ah, taas oo ay sabab u tahay is-weydaarsiga maaddada hidde-wadaha, hiddo-wadaha jira ayaa la farsameeyaa oo kuwa cusub ayaa si buuxda u muuqda Xitaa, xulashadoodu waxay ku dhacdaa xaalado aad uga duwan kuwa loo abuuray baradhada beeraha waaweyn: maqnaanshaha saxaafadda fungicidal, isku mid ahaanshaha dhirta, u badnaanta dhirta ay saameeyeen noocyada kala duwan ee infekshanka fayraska iyo bakteeriyada, u dhowaanshaha yaanyada iyo habeynnada duurjoogta ah, isgoyska firfircoon iyo samaynta oospore, suurtagalnimada sanka oo dhan si uu ugu dhaqmo il caabuq sanadka soo socda.
Waxaas oo dhami waxay u horseedaan kala duwanaansho hidde-wadareed aad u sarreeya oo dadka daaradda deggan. Xaaladaha epiphytotic, duufaanta goor dambe waxay si dhakhso leh ugu faaftaa beeraha khudradda tiro aad u tiro badanna waa la siidaayaa, kuwaasoo u duulaya beero ganacsi oo u dhow. Si kastaba ha noqotee, markay la soo galeen goobaha ganacsiga nidaamka saxda ah ee tiknoolajiyada beeraha iyo ilaalinta kiimikada, qulqulatooyinka yimid ma jiraan wax fursad ah oo ay ku bilaabaan epiphytotics berrinka, taas oo ay ugu wacan tahay maqnaanshaha khadadka cnal ee u adkaysta fungicides iyo ku takhasusay noocyada beeraha.
Ilaha kale ee tallaalka aasaasiga ah wuxuu noqon karaa digriiga buka ee ku xayiran geedo ganacsi. Digriigaan waxaa lagu koray, sida caadiga ah, beeraha leh farsamada beeraha oo wanaagsan iyo ilaalinta kiimikada degdega ah. Noocyada hidde-wadayaasha ah ee go'doomiya infekshanka digriiga waxaa loo waafajiyay horumarinta noocyadooda kala duwan. Noocyadan ayaa si aad ah uga khatar badan abuuritaanka ganacsiga marka loo eego tallaalka ka imanaya jardiinooyinka gaarka loo leeyahay. Natiijooyinka baaritaankayagu sidoo kale waxay taageerayaan fikirkaan. Dadweynaha ka go'doonsan beeraha waaweyn ee leh ilaalinta kiimikada ee si habboon loo qabtay iyo tikniyoolajiyadda beeraha oo wanaagsan kuma kala duwanaan kala duwanaanta hidde-wadayaasha sare Badanaa kuwani waa dhowr khadad isku mid ah oo si xoog leh u dagaal badan.
Dhibaatooyinka ka imanaya walxaha ganacsiga ayaa geli kara dadyowga ku jira beero khudradda ah waxayna ku lug yeelan karaan geeddi-socodka ka socda iyaga. Si kastaba ha noqotee, beerta khudradda, tartankooda ayaa aad uga hooseeya kan ganacsiga, mar dhowna way joogsan doonaan jiritaanka qaab khad cnal ah, laakiin hidde-sideyaashooda ayaa loo isticmaali karaa dadka "beerta".
Infekshinka ku soo baxa dhirta "iskaa wax u qabso" iyo tuulbooyinkii la jaray intii lagu guda jiray goosashada kuma xirna Ruushka, maxaa yeelay Gobollada ugu waaweyn ee baradhada ka baxa ee Ruushka, barafowga ciidda xilliga qaboobaha ayaa lagu arkaa, dhirta ka soo baxda digriiga xilliga qaboobaha carradu ku yartahay. Intaa waxaa sii dheer, sida tijaabooyinkeennu muujinayaan, cudur-sidaha xilli-dambe ee cudurku kuma noolaado heerkul xun xitaa digriiga sii xajistay jiritaankooda. Aagga oomanaha ah, oo ah meesha lagu beero baradhada hore, duqeynta goor dambe waa mid naadir ah sababo la xiriira xilliga qalalan iyo kuleylka.
Marka, waxaan hadda ku guda jirnaa u kuurgalidda kala-soocidda tirada dadka ku dhaca P. ee ku nool "berrin" iyo "beero". Si kastaba ha noqotee, sanadihii la soo dhaafay, geeddi-socodyada ayaa la arkay taasoo horseedaysa isku-xirnaanta iyo dhexgalka genotypes ee dadkan.
Kuwaas waxaa ka mid ah, mid ayaa xusi kara kororka guud ee akhriska iyo qoraalka ee soo saarayaasha yaryar, soo ifbaxa xirmooyinka yaryar ee la awoodi karo ee baradhada abuurka, faafitaanka u diyaargarowga fungicidal ee baakadaha yaryar, iyo luminta cabsida "kiimikada" ee dadweynaha.
Xaaladaha ayaa soo ifbaxa marka, mahadnaqa firfircoonida hal alaab-qeybiye, tuulooyinka oo dhan lagu beeray digrii abuur ah oo noocyo kala duwan ah isla markaana la siinayo baakado yaryar oo isku sun ah. Waxaa loo qaadan karaa in baradho isku nooc ah laga heli doono beeraha ganacsiga ee u dhow.
Dhinaca kale, qaar ka mid ah shirkadaha ka ganacsada sunta cayayaanka ayaa kor u qaadaya qorshooyinka daaweynta kiimikada "miisaaniyad". Xaaladdan oo kale, tirada daaweynta lagu taliyay waa la dhayalsaday waxaana loo bixiyaa fungicides-ka ugu raqiisan, xooga lasaarana maaha kahortaga horumarinta cudurka soo daaha ilaa iyo in la jaro dusha sare, laakiin dib udhac gaar ah ayaa kudhaca epiphytoty si loo kordhiyo waxsoosaarka. Qorshayaasha noocan oo kale ah waa kuwo dhaqaale ahaan qiil u leh marka laga beerinaayo baradhada laga helo alaab abuurka heerka hoose ah, marka mabda 'ahaan aysan jirin wax su'aal ah oo ku saabsan helitaanka dhalidda sare Si kastaba ha noqotee, xaaladdan, marka loo eego dadka beeraha, asalka hidde-wadaha ee baradhada waxay gacan ka geysaneysaa xulashada jinsiyadaha jireed ee gaarka ah, kuwaas oo aad khatar ugu ah noocyadan kala duwan.
Guud ahaan, u janjeedha isu imaatinka "beerta" iyo hababka "wax-soo-saarka" ee wax soo saarka baradhada ayaa noo muuqda kuwo halisteeda leh. Si looga hortago cawaaqib xumadooda, labadaba guriga iyo qaybaha ganacsiga, waxay noqon doontaa lagama maarmaan in la xakameeyo noocyada kala duwan ee baradhada abuurka iyo noocyada kala duwan ee sunta loo adeegsado milkiileyaasha gaarka loo leeyahay ee ku jira baakadaha yaryar, iyo sidoo kale la socoshada qorshayaasha ilaalinta baradhada iyo adeegsiga diyaarinta fungicidal ee qaybta ganacsiga.
Meelaha ganacsiga gaarka loo leeyahay, waxaa ka jira horumar deg deg ah oo aan ku ekaan oo keliya goor dambe, laakiin sidoo kale Alternaria. Inta badan milkiileyaasha qorshooyinka gaarka loo leeyahay ma qaadaan talaabooyin gaar ah oo ay kaga hortagayaan Alternaria, iyaga oo ku qaldaya horumarka Alternaria waxyeellada dabiiciga ah ee dushooda ama horumarka duqowga soo daahay. Sidaa darteed, horumarka ballaaran ee 'Alternaria' ee noocyada nugul ee u nugul, qorshayaasha guryuhu waxay u adeegi karaan inay noqdaan isha tallaalka beerashada ganacsiga.
Farsamooyinka kala duwanaanta
Habka isbedelka
Maadaama dhacdooyinka isbadalku ay yihiin geedi socod aan kala sooc lahayn oo ku socda mowjado hooseeya, dhacdooyinka isbeddellada ee goob kasta waxay kuxirantahay inta jeer ee isku beddelidda goobtan iyo baaxadda dadka. Markaad baraneyso inta jeer ee isku-beddelka noocyada 'inf infansans', tirada gumeysiga ee ku koray warbaahinta nafaqada leh ee xulashada ka dib daaweynta kemikal ama isbadal jireed ayaa badanaa la go'aamiyaa. Sida laga arki karo xogta lagu soo bandhigay Jadwalka 8, inta jeer ee isbeddelka isbeddelka ee isla culeyska meesha kala duwan ayaa ku kala duwanaan kara dhowr amarro oo baaxad leh. Soo noqnoqoshada isbeddelka isbeddelka ee iska caabbinta metalaxyl waxay noqon kartaa mid ka mid ah sababaha isku soo ururinta noocyada noocyada u adkaysta dabeecadda.
Soo noqnoqoshada is-beddelka is-beddelka ama is-beddelada, oo lagu xisaabiyo iyadoo lagu saleynayo tijaabooyinka shaybaarka, had iyo jeer uma dhigna hababka ka dhacaya dadka dabiiciga ah, sababaha soo socda awgood:
1. Kala-baxa nukliyeerka asynchronous, waa wax aan macquul aheyn in lagu qiyaaso inta jeer ee isbadalku ku yimaado hal qarni oo nukliyeer ah. Sidaa darteed, tijaabooyinka badankood waxay si toos ah u bixiyaan macluumaadka kaliya ee ku saabsan inta jeer ee is-beddellada, iyada oo aan la kala soocin labada dhacdo ee is-beddelashada iyo hal dhacdo oo ka dambeysa mitosis
2. Isbeddelada hal-tallaabada ah waxay caadi ahaan yareeyaan dheelitirka hidde-wadaha, sidaas darteed, oo ay weheliso helitaanka guri cusub, jir ahaanta guud ee jirku wuu yaraanayaa. Inta badan isbeddellada tijaabada ah ee la helay waxay leeyihiin dagaal-yari hoos u dhacday oo laguma qoro dadka dabiiciga ah. Sidaa darteed, isku-dheelitirka isku-xirnaanta u dhexeeya heerka iska caabinta P. infestans mutant in fenylamide fungicides iyo heerka kobaca ee bey'ada macmalka ah wuxuu ahaa celcelis ahaan (-0,62), iyo iska caabbinta fungicides iyo dagaal ku caleenta baradhada (-0,65) (Derevyagina et al. , 1993), oo tilmaamaysa jirdhiska hooseeya ee mutantaska. Is-beddellada iska caabinta dimethomorph waxaa sidoo kale weheliyay hoos u dhac xoog leh oo ku yimid jiritaanka (Bagirova et al., 2001).
3. Badanaa isbeddelada is-beddelka ah ee is-beddelka iyo is-beddelka sababa ayaa ah kuwo aan waxtar lahayn oo aan si muuqata ugu muuqan tijaabooyin, laakiin waxay ka dhigan yihiin keyd qarsoon oo ay ku kala duwan yihiin dadka dabiiciga ah. Noocyada isbeddelka ah ee ku go'doonsan tijaabooyinka shaybaarka waxay wataan isbeddello awood leh ama nugul (Kulish iyo Dyakov, 1979). Sida muuqata, diblumaasiyadda nukliyeerka ayaa sharraxaysa iskudayada aan lagu guuleysan ee lagu doonayo in lagu helo mutantada iyada oo ay saameyn ku yeelanayso shucaaca UV-da ee ku kacsan noocyo horay u adkaysan jiray (McKee, 1969). Marka loo eego xisaabinta qoraaga, is-beddelka noocaas ah wuxuu ku dhici karaa ku-celcelis ka yar 1: 500000. U guuritaanka isbadal ku yimaada isbadalka recessive-ka ee loo yaqaan 'homozygous', xaalad ahaan si muuqata loo muujiyey ayaa dhici karta sababo la xiriira galmo ama isku soocelin galmo la'aan (hoos eeg). Si kastaba ha noqotee, xitaa kiiskan, isbeddelku wuxuu qarin karaa astaamaha ugu waaweyn ee nooca duurjoogta ah ee 'mycelium' cenotic (multinucleated) mycelium oo si muuqata loo hagaajiyay oo keliya inta lagu guda jiro sameynta xayawaannada mononuclear.
Jadwalka 8. Soo noqnoqoshada P. infestans waxay isku badashaa walxaha ka hortaga koritaanka iyadoo loo marayo tallaabada nitrosomethylurea (Dolgova, Dyakov, 1986; Bagirova et al., 2001)
Iskuxirka | Isbedelka isbeddelka |
Oxytetracycline | 6,9 10 x-8 |
Blasticidin S | 7,2 x 10-8 |
Streptomycin | 8,3 x10-8 |
Trichothecin | 1,8 10 x-8 |
Cycloheximide | 2,1 10 x-8 |
Daaconil | <4 x 10-8 |
Dimethomorph | 6,3 10 x-7 |
Metalaxil | 6,9 10 x-6 |
Tirada dadka ayaa sidoo kale door muhiim ah ka ciyaareysa dhacdooyinka isbeddellada lama filaanka ah. Dad aad u tiro badan, oo ay ku jiraan tirada unugyada N> 1 / a, halkaasoo ah heerka isbeddelka isbeddellada, isbadalku wuxuu joogsadaa inuu noqdo dhacdo aan kala sooc lahayn (Kvitko, 1974).
Xisaabinta ayaa muujineysa in celcelis ahaan ku faafidda beed baradho ah (35 dhibic halkii geed), 8x1012 kudka ayaa maalin walba lagu sameeyaa hal hektar (Dyakov iyo Suprun, 1984). Sida muuqata, dadka noocan oo kale ah waxay ka kooban yihiin dhammaan isbeddellada loo oggol yahay nooca is-weydaarsiga goob kasta. Xitaa isbeddel naadir ah, oo ku dhaca inta u dhexeysa 10-9, waxaa heli doona kun qof oo ka mid ah malaayiin ku nool hal hektar oo beer baradho ah. Is-beddelka ku dhaca soo noqnoqoshada sarreeya (tusaale ahaan, 10-6), dadkan oo kale, is-beddello isku-dhafan oo kala duwan ayaa ku dhici kara maalin kasta (isku mar laba loci) nidaamka isbadalku wuxuu badali doonaa dib uqabashada.
Tahriibka
Dadka loo yaqaan 'inf infansans', laba nooc oo waaweyn oo tahriib ah ayaa loo yaqaan: in la xiro masaafada (gudaha baradho ama beero deriska la ah) iyadoo lagu faafinayo zoosporangia hawo hawada ama buufin roob, iyo masaafo dheer - iyadoo la beero digriiga ama miraha yaanyada la soo qaado. Habka koowaad wuxuu bixiyaa fiditaanka diiradda cudurka, tan labaad - abuuritaanka ujeeddooyin cusub meelaha fog fog ee aasaasiga ah.
Faafitaanka infekshinka digriiga yaanyada iyo miraha kaliya kuma darsato soo ifbixida cudurka meelo cusub, laakiin sidoo kale waa isha ugu weyn ee kala duwanaanta hidaha ee dadka. Gobolka Moosko, baradho ayaa la beeray, oo laga keenay gobolada kala duwan ee Ruushka iyo Galbeedka Yurub. Miraha yaanyada waxaa laga keenaa gobollada koonfureed ee Ruushka (gobolka Astrakhan, gobolka Krasnodar, Waqooyiga Caucasus). Abuurka yaanyada, oo waliba u adeegi kara ilaha cudurka (Rubin et al., 2001), sidoo kale waxaa laga keenaa gobollada koonfureed ee Ruushka, Shiinaha, waddamada Yurub iyo dalal kale.
Marka loo eego xisaabinta E. Mayr (1974), isbeddelada hidaha ee dadka maxalliga ah ee ay sababtay isbadalku marar dhif ah ayuu ka badanayaa 10-5 halkii goobba, halka dadka furan, isweydaarsiga ay sabab u tahay socodka hiddo-wadaha ugu yaraan 10-3 - 10-4.
U guuritaanka digriiga cudurka qaba ayaa mas'uul ka ah soo gelitaanka P. infestans ee Yurub, kuna faafaya dhammaan gobollada adduunka ee baradhada laga beero; waxay keeneen isbadalkii ugu darnaa ee dadka. Dillaacii ugu dambeeyay ee baradhada ayaa ka soo muuqday dhulkii Boqortooyada Ruushka ku dhowaad isku mar muuqaalkiisa Galbeedka Yurub.
Tan iyo markii cudurka lagu arkay 1846-1847 ee wadamada Baltiga oo kaliya sanadihii ku xigay ku faafay Belarus iyo gobolada waqooyi-galbeed ee Ruushka, asalkeeda Galbeedka Yurub waa iska cadahay. Isha ugu horeysa ee duqeynta goor dambe ka dhacda Dunida Duugga ah si muuqata uma muuqato. Qiyaasta ay soo saartay Fry et al. (Fry et al., 1992; Fry, Goodwin, 1995, Goodwin et al., 1994) waxay soo jeedinaysaa in dulinku markii ugu horreysay ka yimid Mexico ilaa Waqooyiga Ameerika, halkaas oo uu ku faafay dalagyada, ka dibna loo qaaday Galbeedka Yurub. (fig. 7).
Dariiqii soo noqnoqday awgiis (saamaynta laba-geesoodka ah ee "dhalada"), hal isku-dhegyo ayaa la gaadhay Yurub, oo dhasha ay ka dhasheen faafa oo ku baahday dhulkii Hore ee Adduunku lahaa ee ay baradho ku korayso. Caddayn u ah male-awaalkan, qorayaashu waxay caddeeyeen, marka hore, baahinta kaliya ee hal nooc oo isweydaarsi ah (A1) iyo, tan labaad, isu-ekaanshaha hidde-wadaha noocyada cilmiga laga bartay ee ka kala yimid gobollada kala duwan (dhammaantood waxay ku saleysan yihiin astaamo molikal ah, oo ay ku jiraan 2 isozyme loci, qaababka faraha DNA, iyo Qaab dhismeedka mitochondrial DNA waa isku mid, waxayna u dhigantaa midka loo yaqaan 'clone US-1' ee lagu sharaxay USA). Si kastaba ha noqotee, xogta qaar waxay keeneysaa shaki ku saabsan ugu yaraan qaar ka mid ah qodobbada qiyaasta la sheegay. Falanqaynta P. infestans DNA mitochondrial DNA oo laga soocay shaybaarka baradhada herbarium ee cudurka ku dhacay mudadii ugu horeysay ee epiphytotic ee 40s ayaa muujisay inay ku kala duwan yihiin qaabdhismeedka mitochondrial DNA ee loo yaqaan 'clone US-1', taas oo, sidaa darteed, uguyaraan ahayd ma aha isha kaliya ee cudurka laga helo Yurub (Ristaino et al, 2001).
Xaaladda duqeyntu goor dambe way ka sii xumaatay 80-meeyadii qarnigii XX. Isbedelada soo socda ayaa dhacay:
1) Celceliska gardarrada dadweynuhu wuu kordhay, taas oo horseeday, gaar ahaan, faafitaanka baahsan ee nooca ugu waxyeellada badan ee goor dambe - waxyeelo soo gaadhay geedaha yaryar iyo jirida.
2) Waxaa jiray isbedel ku yimid waqtigii duqeynta soo daahay ee baradhada - laga soo bilaabo dabayaaqadii July ilaa horraantii July iyo xitaa illaa dhammaadka Juun.
3) Nooca isweheshiga ee A2, oo horay uga maqnaa Adduunyadii Hore, wuxuu noqday mid meel walba yaal.
Isbadalada waxaa kahoreeyay laba dhacdo: isticmaalka baaxada leh ee maadada cusub ee loo yaqaan "metalaxyl" (Schwinn iyo Staub, 1980) iyo soo bixitaanka Mexico oo ah dhoofiyaha aduunka ee baradhada (Niederhauser, 1993). Iyadoo la raacayo tan, laba sababood oo ku saabsan isbeddelka dadweynaha ayaa la soo saaray - beddelaadda nooca isutagga iyadoo la adeegsanayo metalaxyl (Ko, 1994) iyo soosaarid ballaaran oo noocyo cusub ah oo leh digriiga cudurka laga keenay ee Mexico (Fry iyo Goodwin, 1995). In kasta oo iskudhafka noocyada isutagga ee saameynta metalaxyl la helay aysan ahayn oo keliya Ko, laakiin sidoo kale shaqooyinka laga fuliyay shaybaarka Jaamacadda Moscow (Savenkova, Chherepennicova-Anikina, 2002), mala-awaalka labaad ayaa la door bidaa. Iyada oo ay weheliso muuqaalka nooca labaad ee isugeynta, isbadalo halis ah ayaa ka dhacay genotypes-ka noocyada jilicsan ee Ruushka P., oo ay ku jiraan hiddo-wadayaasha dhexdhexaadka ah (isozyme iyo RFLP loci), iyo sidoo kale qaab dhismeedka mitochondrial DNA. Isku-dhafka isbeddeladan laguma sharxi karo ficilka metalaxyl; halkii, waxaa jiray soo dejin ballaaran oo noocyo cusub ah oo ka yimid Mexico, taas oo, iyada oo ka sii dagaal badan (Kato et al., 1997), ay barakicisay noocyadii hore (US-1), oo ku adkaatay dadka. Isbeddelka ku yimid halabuurka dadka reer Yurub wuxuu dhacay waqti aad u yar - laga soo bilaabo 1980 ilaa 1985 (Fry et al., 1992). Dhulkii USSR-dii hore, "noocyo cusub" ayaa laga helay aruurinta Estonia sanadkii 1985, taasi waa, horaantii Poland iyo Jarmalka (Goodwin et al., 1994). Markii ugu dambeysay ee "culeys duug ah US-1" ee Ruushka laga go'doomiyey yaanyo qaba gobolka Moskow 1993 (Dolgova et al., 1997). Sidoo kale Faransiiska, noocyo "duug ah" ayaa laga helay beero yaanyo ilaa horraantii 90-meeyadii, taasi waa, ka dib markii ay muddo dheer ku baaba'een baradho (Leberton iyo Andrivon, 1998). Isbedelada ku dhaca P. infestans noocyada kala duwan waxay saameeyeen astaamo badan, oo ay ku jiraan kuwa muhiimad weyn leh, waxayna kordhisay waxyeelada cudurka goor dambe.
Dib isu soo celinta galmada
Si dib loogu soo celiyo galmada si ay gacan uga geysato kala duwanaanta, waxaa lagama maarmaan ah, marka hore, joogitaanka laba nooc oo isweydaarsi ah oo ku yimaada dadweynaha marka loo eego saamiga ku dhow 1: 1, iyo, tan labaad, joogitaanka kala duwanaanta bilowga ah.
Qiyaasta noocyada isutagga ayaa si weyn ugu kala duwan dadyowga kala duwan iyo xitaa sannado kala duwan oo hal dad ah (Sanduuqa 9,10, 90). Sababaha isbeddelada noocaas ah ee isdaba joogga ah ee isweydaarsiga noocyada isweheshiga ee dadweynaha (sida, tusaale ahaan, Ruushka ama Israel horaantii 2002-meeyadii qarnigii la soo dhaafay) lama garanayo, laakiin waxaa la rumeysan yahay in tani ay ugu wacan tahay soo-dejinta midabbo tartan badan (Cohen, XNUMX).
Qaar ka mid ah xogta aan tooska ahayn waxay muujinayaan habka habka galmada sanadaha qaarkood iyo gobollada qaarkood:
1) Daraasadaha dadweynaha ee ka yimid gobolka Moscow waxay muujiyeen in 13 dadweyne ah oo saamiga A2 nooca isugeynta uu ka yaraa 10%, wadarta guud ee kaladuwanaanta hidde-wadaha loo xisaabiyay saddex isozyme loci waxay ahayd 0,08, iyo 14 dad ah oo qaybta A2 ay dhaaftay 30%, kala duwanaanta hidde-wadaha ayaa laba jeer ka sarreysay (0,15) (Elansky et al., 1999). Marka, sare u kaca ixtimaalka galmada, waxaa sii weynaanaya kala duwanaanshaha hidaha ee dadka.
2) Xidhiidhka ka dhexeeya saamiga noocyada isugeynta ee dadweynaha iyo xoojinta samaynta oospore ayaa lagu arkay Israel (Cohen et al., 1997) iyo Holland
(Flier et al., 2004). Daraasaddeennu waxay muujisay in dadka ku go'doonsan nooca A2 ee isugeynta ay yihiin 62, 17, 9, iyo 6%, oospores ayaa laga helay 78, 50, 30, iyo 15% ee caleemaha baradhada la falanqeeyay (oo leh 2 ama dhibco ka badan).
Muunado 2 ama dhibco ka badan leh ayay aad ugu dhowdahay inay ku jiraan oospore ka badan shaybaarro leh 1 bar (32 iyo 14% baarka, siday u kala horreeyaan) (Apryshko et al., 2004).
Oospores waxay aad ugu badnaayeen caleemaha qaybta dhexe iyo hoose ee geedka baradhada (Mytsa et al., 2015; Elansky et al., 2016).
3) Gobollada qaarkood, genotypes gaar ah ayaa laga helay, dhacdadooda oo lala xiriirinayo isu galmoodka galmada. Sidaa darteed, Poland 1989 iyo Faransiiska 1990, noocyo isku mid ah oo loo yaqaan 'glucose-6-'
fosfat isomerase (GPI 90/90). Tan iyo markii hore oo keliya 10/90 heterozygotes ayaa lala kulmay 100 sano, homozygosity waxaa loo aaneeyaa dib-isu-galmoodka galmada (Sujkowski et al., 1994). Gudaha Kolombiya (USA), go'doominta isku-darka A2 ee GPI 100/110 iyo A1 iyo GPI 100/100 waa wax iska caadi ah, laakiin dhammaadka xilli-1994kii (Agoosto 16 iyo Sebtember 9), noocyo leh genotypes recombinant (A1 GPI 100/110) iyo A2 GPI 100/100) (Miller et al., 1997).
4) Qaar ka mid ah dadweynaha ka yimid Poland (Sujkowski et al., 1994) iyo Waqooyiga Caucasus (Amatkhanova et al., 2004), qaybinta sawirka DNA-ga faraha iyo borotiinka allozyme loci waxay u dhigantaa Hardy-Weinberg qaybinta, taas oo muujinaysa
ku saabsan saamiga sare ee ka qayb qaadashada isu geynta galmada ee kala duwanaanta dadka. Gobollada kale ee Ruushka, looma helin wax waraaqo ah oo ku saabsan qaybinta Hardy-Weinberg ee dadka, laakiin joogitaanka isku xirnaanta cudurka ayaa la muujiyey, taas oo muujinaysa sida ugu badan ee taranka isku-darka (Elansky et al., 1999).
5) Kala duwanaanta hidde-wadaha (GST) ee udhaxeeya muruqyada noocyadooda kala duwan (A1 iyo A2) way ka hooseeyeen inta udhaxeysa dadyowga kala duwan (Sujkowski et al., 1994), oo si aan toos ahayn u tusaysa iskutallaabta galmada.
Isla mar ahaantaana, ka-qaybgalka isku-darka galmada ee kala-duwanaanta dadka ma noqon karo mid aad u sarreeya. Qaadhaankan waxaa loo xisaabiyay dadka ku nool gobolka Moscow (Elansky et al., 1999). Marka la eego xisaabinta Lewontin (1979) "dib-u-isu-soo-celinta, oo soo saari kara noocyo cusub oo ka socda laba loci oo leh soo noqnoqosho aan ka badnayn wax soo saarka heterozygosities-kooda, waxay noqoneysaa mid waxtar leh oo keliya haddii qiimaha heterozygosity ee labada allele ay horeba u sarreeyeen."
Marka la eego saamiga labada nooc ee lammaanaha ah, ee caadiga u ah gobolka Moosko, oo u dhigma 4: 1, soo noqnoqoshada isugeynta waxay noqon doontaa 0,25. Suurtagalnimada in laga gudbo noocyo ayaa noqon doona heterozygous laba ka mid ah saddexda daraasadood ee loo yaqaan 'isozygous loci' ee dadka daraasadda lagu sameeyay waxay ahayd 0,01 (2 nooc oo ka baxsan 177). Sidaas awgeed, suurtagalnimada in ay dhacdo heterozygotes labalaab ah oo ka dhalanaya dib-isu-soo-celinta waa inaysan ka badnayn badeecadooda oo lagu dhuftay itimaalka isgoyska (0,25x0,02x0,02) = 10-4, i.e. dib u soo kabashada galmada badanaa kuma dhacdo shaybaarka daraasadda ee noocyada. Xisaabintaan waxaa loo sameeyay dadka ka yimid gobolka Moscow, kuwaas oo lagu garto kala duwanaansho xad dhaaf ah. Dadka monomorphic sida kuwa Siberia, habka galmada, xitaa haddii ay ku dhacdo dadka shaqsiyaadka ah, saameyn kuma yeelan karaan kala duwanaanshahooda hidde.
Intaa waxaa dheer, P. infestans waxaa lagu gartaa isku-dheelitirnaan la'aanta chromosome ee meiosis, taas oo keenta aneuploidy (Carter et al., 1999). Xadgudubyada noocan ahi waxay yareeyaan bacriminta isku-dhafka.
Dib-u-cusbooneysiinta jinsiga, isu-beddelka hidda-wadaha mitotic
Tijaabooyin ku saabsan kala-goynta P. infestans noocyo kala duwan oo leh isbeddello iska caabin ah ka-hortaggayaasha koritaanka kala duwan, soo bixitaanka misolates u adkaysta labada xakameeyayaasha ayaa la helay (Shattock iyo Shaw, 1975; Dyakov, Kuzovnikova, 1974; Kulish, Dyakov,
1979). Muruqyo adkaysi u leh laba kahortagayaasha kobcitaanka ayaa ka dhashay natiijada heterokaryotization ee mycelium, kiiskanna waxay ku dhegeen inta lagu gudajiray taranka mononuclear zoospores (Judelson, Ge Yang, 1998), ama ma aysan kala bixin farcanka monozoosporous, maxaa yeelay waxay lahaayeen tetraploid (tan iyo markii ay go'doominta hore yihiin diploid) nuclei (K , 1979). Dibloomaatada Heterozygous oo loo kala soocay si aad u hooseeya sababtoo ah haploidization, chromosome nondisjunction, iyo mitotic gudbinta (Poedinok et al., 1982). Inta jeer ee hawlgalladaan waa la kordhin karaa iyadoo la kaashanayo saameyno gaar ah oo ku saabsan dibloomaatada heterozygous (tusaale ahaan, shucaaca UV ee biqilka dhirtu ku jirto).
In kasta oo sameysanka iskudhafka daaqsinka leh iska caabin laba-geesood ah aysan ku dhicin fitamiinnada oo keliya, laakiin sidoo kale digriiga baradhada ee uu ku dhacay isku-dhafka mutant (Kulish et al., 1978), way adag tahay in la qiimeeyo doorka dib-u-isu-beddelka jinsiga ee jiilka genotypes cusub ee dadweynaha. Inta jeer ee sameynta segregan ay sabab u tahay haploidization, isku xirnaansho la'aanta koromosoomyada iyo iskutallaab la'aanta mitotic iyada oo aan la helin saameyn gaar ah waa wax la iska indhatiri karo (in ka yar 10-3).
Soo bixitaanka segregants homozygous ee noocyada heterozygous waxaa laga yaabaa inay ku saleysnaato marinka mitotic-ka iyo beddelidda hiddo-wadaha, taas oo P. sojae ku dhacda inta jeer ee 3 x 10-2 ilaa 5 x 10-5 halkii goobba, iyadoo kuxiran culeyska (Chamnanpunt et al. , 2001).
In kasta oo inta jeer ee ay dhacaan heterokaryons iyo heterozygous dibloomaasiyiin ay noqdeen kuwo si lama filaan ah u sarreeya (gaaraya tobaneeyo boqolkiiba), howshani waxay dhacdaa oo keliya marka dhaqamada mutantiga ee laga helo isla caynkaas la kala qaado. Markaad isticmaaleyso noocyo kala duwan oo laga soocay dabeecadda, heterokaryotization ma dhacdo (ama waxay ku dhacdaa soo noqnoqosho aad u hooseysa) iyadoo ay ugu wacan tahay jiritaanka is-waafajin la'aanta daaqsinka (Poedinok iyo Dyakov, 1981; Anikina et al., 1997b; Cherepennikova-Anikina et al., 2002). Sidaa awgeed, doorka dib-u-soo-kabashada jinsiga ah ayaa loo yareyn karaa oo keliya dib-u-soo-celinta intraclonal ee heterozygous nuclei iyo u gudubka hiddo-wadaha shaqsiyeed ee udhaxeeya gobolka 'homozygous' oo aan lahayn geedi socod galmo. Nidaamkani wuxuu noqon karaa mid muhiimad gaar ah u leh cudurrada faafa ee noocyada isku-dhafka caabbinta ama ka-hortagga fangaska. U guuritaankeeda dowlad khaniis ah sababta oo ah habka loo yaqaan 'parasexual' ayaa kordhin doonta iska caabbinta sideha isbedelka (Dolgova, Dyakov, 1986).
Soo-kicinta hiddo-wadaha
Noocyada 'Heterothallic Phytophthora' waxay awood u leeyihiin inay ku dhex milmaan sameynta oospores isku dhafan (eeg Vorob'eva iyo Gridnev, 1983; Sansome et al., 1991; Veld et al., 1998). Isugeynta dabiiciga ah ee labada nooc ee 'Phytophthora' waxay ahayd mid aad u dagaal badan oo dishay kumanaan alder ah oo ku nool Boqortooyada Midowday (Brasier et al., 1999). P. infestans waxay ku dhici karaan noocyada kale ee hiddo-wadaha (P. erythroseptica, P. nicotianae, P. Cactorum, iwm.) Dhirta martida loo yahay ee caadiga ah iyo carrada, laakiin waxaa jira macluumaad yar oo ku saabsan suugaanta oo ku saabsan suurtagalnimada isku-darka isku-dhafan. Xaaladaha shaybaarka hoostiisa, iskudhaf ayaa laga helay inta udhaxeysa P. infestans iyo P. Mirabilis (Goodwin iyo Fry, 1994).
Jadwalka 9. Qiyaasta P. infestans noocyada nooca A2 ee mating ee dalal kala duwan oo adduunka ah muddadii u dhexaysay 1990 ilaa 2000 (sida ku cad xogta ilaha suugaanta furan iyo bogagga www.euroblight.net, www.eucablight.org)
dalka | 1990 | 1991 | 1992 | 1993 | 1994 | 1995 | 1996 | 1997 | 1998 | 1999 | 2000 |
---|---|---|---|---|---|---|---|---|---|---|---|
Belarus | 33 (12) | 34 (29) | |||||||||
Belgium | 15 (49 *) | 6 (66) | 20 (86) | ||||||||
Ecuador | 0 (13) | 0 (12) | 0 (19) | 0 (21) | 12 (41) | 25 (39) | 15 (75) | 22 (73) | 25 (68) | 0 (35) | |
Estonia | 8 (12) | ||||||||||
England | 4 (26) | 3 (630) | 9 (336) | ||||||||
Finland | 0 (15) | 19 (117) | 12 (16) | 21 (447) | 6 (509) | 9 (432) | 43 (550) | ||||
France | 0 (35) | 0 (56) | 0 (83) | 0 (67) | 0 (86) | 2 (135) | 7 (156) | 6 (123) | 0 (73) | 0 (285) | 0 (135) |
Hungary | 72 (32) | ||||||||||
Ireland | 4 (145) | ||||||||||
Waqooyiga. Ireland | 10 (41) | 9 (58) | 1 (106) | 0 (185) | 0 (18) | 0 (56) | 0 (35) | 0 (26) | |||
Netherlands | 7 (41) | 5 (276) | 24 (377) | 44 (353) | 23 (185) | ||||||
Norway | 25 (446) | 28 (156) | 8 (39) | 18 (257) | 38 (197) | ||||||
Peru | 0 (34, 1984 -86) | 0 (287, 1997-98) | 0 (112) | 0 (66) | |||||||
Poland | 19 (180) | 21 (142) | 33 (256) | 26 (149) | 35 (70) | ||||||
Iskodland | 25 (147) | 11 (163) | 22 (189) | 5 (22) | |||||||
Sweden | 25 (263) | 62 (258) | 49 (163) | ||||||||
Wales | 0 (16) | 7 (97) | 0 (48) | 0 (25) | |||||||
Kuuriya | 36 (42) | 10 (130) | 15 (98) | ||||||||
Shiinaha | 20 (142, 1995-98) | 0 (6) | 0 (8) | 0 (35) | |||||||
Colombia | 0 (40, 1994-2000) | ||||||||||
Urugay | 100 (25, 1998-99) | ||||||||||
Morocco | 60 (108, 1997-2000) | 52 (25) | 42 (40) | ||||||||
Serbia | 76 (37) | ||||||||||
Mexico (Toluca) | 28 (292, 1988-89) | 50 (389, 1997-98) |
Shaxda 10. Qiyaasta P. infestans muruqyada leh nooca A2 ee mating-ka dalal kala duwan oo adduunka ah muddadii u dhexaysay 2000 ilaa 2011
dalka | 2001 | 2002 | 2003 | 2004 | 2005 | 2006 | 2007 | 2008 | 2009 | 2010 | 2011 |
---|---|---|---|---|---|---|---|---|---|---|---|
Austria | 65 (83) | ||||||||||
Belarus | 42 (78) | ||||||||||
Belgium | 20 (102 *) | 4 (32) | 50 (14) | 25 (16) | 62 (13) | 54 (26) | 70 (54) | 30 (23) | 29 (35) | 62 (71) | 45 (49) |
Switzerland | 89 (19) | ||||||||||
Czech Republic | 35 (31) | 54 (64) | 38 (174) | 12 (80) | |||||||
Germany | 95 (53) | ||||||||||
Denmark | 48 (52) | ||||||||||
Ecuador | 5 (178) | 6 (108) | 9 (121) | 18 (94) | 2 (44) | 0 (66) | 5 (47) | ||||
Estonia | 54 (25) | 0 (24) | 33 (62) | 45 (140) | 25 (100) | 12 (103) | |||||
England | 4 (47) | 10 (96) | 31 (55) | 55 (790) | 68 (862) | 70 (552) | 68 (299) | ||||
Finland | 47 (162) | 12 (218) | 42 | ||||||||
France | 0 (186) | 4 (108) | 8 (61) | 22 (103) | 33 (303) | 65 (378) | 74 (331) | 75 (125) | 75 (12) | ||
Hungary | 48 (27) | 48 (90) | 9 | 7 | |||||||
Waqooyiga. Ireland | 0 (38) | 0 (58) | 0 (40) | 0 (24) | 5 (54) | 0 (18) | 27 (578) | 45 (239) | 36 (213) | 82 (60) | 10 (80) |
Netherlands | 66 (24) | 93 (15) | 91 (11) | ||||||||
Norway | 39 (328) | 3 (115) | 12 (19) | ||||||||
Peru | 0 (36) | ||||||||||
Poland | 25 (46) | 10 (30) | 85 (20) | 38 (44) | 75 (66) | 55 (56) | 65 (35) | 72 (81) | 85 (21) | ||
Iskodland | 3 (213) | 2 (474) | 24 (135) | 86 (337) | 88 (386) | 74 (172) | |||||
Sweden | 60 (277) | 39 (87) | |||||||||
Slovakia | 0 (36) | 14 (26) | 62 (26) | 0 (26) | |||||||
Wales | 25 (12) | 68 (106) | 80 (88) | 92 (143) | 75 (45) | ||||||
Kuuriya | 46 (26) | ||||||||||
Brazil | 0 (49) | 0 (30) | |||||||||
Shiinaha | 10 (30) | 0 (6) | 0 (6) | ||||||||
Vietnam | 0 (294, 2003-04) | ||||||||||
Ugaandha | 0 (8) |
Dhaqdhaqaaqa isku dhafka genotypic ee dadweynaha
Isbeddelada ku dhaca isir ahaanta hidde-wadaha ee P. dadka ku dhaca infestans dadku waxay ku dhici karaan saameynta hijrada ee xirmooyinka cusub ee gobollada kale, dhaqamada beeraha (isbeddelka noocyada, isticmaalka fungicides), iyo xaaladaha cimilada. Saameynta dibedda waxay saameyn ku yeelataa jimicsiyo kala duwan heerarka kala duwan ee wareegga nolosha; Sidaa darteed, dadweynuhu sannad walba waxay la kulmaan isbeddelo meerto ah oo ku yimaada mowjadaha hidda-wadaha ee xulashada, sababo la xiriira isbeddelka doorka ugu weyn ee hidda-wadaha iyo xulashada.
Saamaynta noocyada kala duwan
Noocyada cusub ee leh hiddo-wadaha wax ku oolka ah ee iska caabbinta toosan (R-genes) ayaa ah cunsur xulasho xoog leh leh oo lagu xusho midabyo leh hiddo-wadayaal dhammaystiran oo ku dhaca tirada dadka inf. Maqnaanshaha caabbinta aan caadiga ahayn ee noocyada baradhada ee hor istaagaya koritaanka dadka cudurada keena, geeddi-socodka lagu beddelayo isku-darka ugu weyn ee dadka ayaa si dhakhso leh u dhaca. Marka, ka dib markii uu ku faafay gobolka Moscow ee kaladuwan ee Domodedovsky, oo leh hiddo-wadaha R3 iska caabbinta, inta jeer ee isku xigxiga ee u nugul noocyadan kala duwan ayaa ka kordhay 0,2 ilaa 0,82 hal sano (Dyakov, Derevjagina, 2000).
Si kastaba ha noqotee, isbeddelka ku yimaada inta jeer ee hiddo wadayaasha (cudur-sidaha) ee dadka ku dhaca ma aha oo kaliya saameynta noocyada baradhada la beeray. Tusaale ahaan, Belarus illaa 1977, clones oo leh hiddo-wadayaasha 1 iyo 4 ayaa ku badan, taas oo ay sababtay beerista noocyo baradho ah oo leh hiddo-wadayaal iska caabin ah R1 iyo R4 (Dorozhkin, Belskaya, 1979). Si kastaba ha noqotee, dhammaadkii 70s ee qarnigii 2002aad, clones waxay u muuqdeen hiddo-wadayaal kaladuwan oo kala duwan iyo iskudhafkooda, iyo hiddo-wadayaasha iska caabinta waligood looma adeegsan taranka baradhada (hiddo-wadayaasha dheeraadka ah ee fiiruska) (Ivanyuk et al., XNUMX). Sababta muuqaalka dhirta noocaas ah, waxay u eg tahay socdaalka Yurub ee walxaha faafa ee ka yimaada Mexico oo leh digriiga baradhada. Guriga dhexdiisa, xirmooyinkaani waxay ku kobceen oo keliya baradhada la beeray, laakiin sidoo kale noocyada duurjoogta ah ee sidda hiddo-wadayaasha iska caabinta kala duwan; Sidaa darteed, isku-darka hiddo-wadayaasha fara badan ee hidde-wadaha ayaa lagama maarmaan u ahaa badbaadada xaaladahaas.
Sida noocyada leh iska caabin aan qeexanayn, iyagu, iyaga oo yareeya xaddiga dhalmada ee cudur-sidaha, waxay dib u dhigaan isbeddelka dadkooda, taas oo, sidaan horeba u sheegnay, ay tahay shaqo tiro. Maaddaama gardarradu ay tahay mid polygenic ah, klootooyin ay ku jiraan tiro aad u tiro badan oo hidde-sideyaal ah "gardarro" ayaa isku soo uruurinaya sida ugu dhaqsaha badan tirada dadka. Sidaa darteed, jinsiyadaha gardarrada aadka u lihi maaha wax soo saar la-qabsi ah oo loogu talagalay noocyo la beeray oo leh iska caabin aan qeexnayn, laakiin, liddi ku ah, waxay u dhowdahay in lagu ogaado abuurka noocyo aad u nugul oo ah kuwa soo urursada kudka dulinka
Marka, Ruushka dadka ugu dagaal badan ee P. Infestans waxaa laga helay aagagga sanadlaha ah ee loo yaqaan 'epiphytoties' (dadweynaha ka soo jeeda gobollada Sakhalin, Leningrad, iyo Bryansk). Rabshadaha dadkan waxay noqdeen kuwo ka sarreeya kuwa reer Mexico (Filippov et al., 2004).
Intaas waxaa sii dheer, oospore ka yar ayaa lagu sameeyay caleemaha noocyo adkaysi leh marka loo eego kuwa u nugul (Hanson iyo Shattock, 1998), taasi waa, iska caabinta aan kala sooca lahayn ee noocyada kala duwan waxay sidoo kale yareysaa awoodaha isu-soo-nooleynta dulinka iyo suurtagalnimada habab kale oo qabow.
Saamaynta sunta sunta
Fungicides kaliya ma yareynayso tirada fungi phytopathogenic, i.e. waxay saameyn ku yeelanayaan astaamaha tirada dadkooda, laakiin waxay sidoo kale beddeli karaan inta jeer ee isir ahaanta shaqsiyeed, ie saamayn ku yeesho tayada bulshada. Tilmaamayaasha ugu muhiimsan ee dadweynaha ee isbeddelaya saameynta sunta sunta ayaa ah kuwa soo socda: isbeddelada ku dhaca iska caabbinta fungicides, isbeddelada kufsiga iyo kacsanaanta, iyo isbeddelada nidaamka taranka.
Saamaynta sunta loo adeegsado ee ku saabsan caabbinta iyo dagaal dad-weynaha
Heerka saameynta noocaas ah waxaa lagu go'aamiyaa, ugu horreyntii, nooca fungicide-ka loo isticmaalo, oo xaalad ahaan loogu qaybin karo polysite, oligosite iyo monosite.
Midka hore waxa ku jira sunta lagu buuxiyo ee lala xiriiriyo. Iska caabinta iyaga (haddii ay suurtagal tahayba) waxaa gacanta ku haya tiro aad u fara badan oo hiddo-wadayaal aad u liita. Waxyaabahani waxay go'aamiyaan maqnaanshaha isbeddelada muuqda ee caabbinta dadweynaha ka dib daaweynta sunta loo yaqaan (inkastoo tijaabooyinka qaarkood xoogaa iska caabin ah la helay). Dadka fangaska ku jira ee la ilaaliyo ka dib ku buufinta sunta lagu buufiyo waxay ka kooban tahay laba nooc oo noocyo kala duwan ah:
1) Dhibaatooyin ku haray meelaha dhirta aan lagu daweynin daroogada. Maaddaama uusan jirin xiriir lala sameeyay sunta fangaska, gardarrada iyo iska caabinta noocyadan isma beddelo.
2) Muruqyada taabanaya sunta fangaska, isku urursanaanta meelaha lagala xiriiro ayaa ka hooseysa tan dilaaga ah. Sida aan kor ku soo sheegnay, iska caabinta qaybtan dadka ka mid ah sidoo kale isma beddelayso, hase yeeshe, saamaynta waxyeellada ee qayb ka mid ah fangaska xitaa marka la eego isku-darka sublethal ee ku saabsan dheef-shiid kiimikaadka fungal, jimicsiga guud iyo qayb ka mid ah dulinkiisa, gardarrada, hoos u dhaca (Derevyagina iyo Dyakov, 1990).
Sidaa darteed, xitaa qayb ka mid ah dadka aan dhiman, u soo gaadhin inay la xiriiraan fungicide, waxay leedahay dulmi daciif ah mana noqon karto ilaha epiphytotics. Sidaa darteed, daaweynta taxaddarka leh ee yareyneysa inta jeer ee saamiga dadka ee aan la xiriirin fungicide-ka waa xaalad lagu guuleysto tallaabooyinka ilaalinta. Diidmada oligosite fungicides waxaa xukuma dhowr hiddo-wade oo lagu daro.
Is-beddelidda hiddo-wade kasta wuxuu keenaa xoogaa xoogaa iska caabin ah, iyo heerka guud ee iska caabbinta waxaa sabab u ah is-beddelka noocaas ah. Sidaa darteed, kororka iska caabbintu waxay u dhacdaa si tallaabo tallaabo ah. Tusaalaha kororka tallaabada tallaalka ah ayaa ah isbeddello iska caabiya dimethomorph fungicide, oo si ballaadhan loogu adeegsado in baradhada laga ilaaliyo duqowga goor dambe. Iska caabinta Dimethomorph waa polygenic iyo daro. Isbeddel hal-tallaabo ah ayaa waxoogaa kordhiya iska caabinta.
Isbeddel kasta oo dambe wuxuu yareeyaa cabbirka bartilmaameedka iyo, sidaas awgeed, inta jeer ee isbeddelada ku xiga (Bagirova et al., 2001). Kordhinta celceliska iska caabinta dadweynaha ka dib daaweyn badan oo lala yeesho fungicide oligosite waxay u dhacdaa si tartiib tartiib ah iyo si tartiib tartiib ah. Qiyaasta geeddi-socodkan waxaa lagu go'aamiyaa ugu yaraan saddex arrimood: inta jeer ee isku-beddelidda hiddo-wadaha iska caabbinta, isku dheelitirnaanta iska caabbinta (saamiga qiyaasta halista ah ee culeys iska caabin ah ee la xiriirta mid xasaasi ah) iyo saameynta is-beddelka hiddo-wadaha iska caabinta.
Inta jeer ee ay dhacdo isbeddel kasta oo soo socda ayaa ka hooseeya kii hore, sidaa darteed geeddi-socodku wuxuu leeyahay dabeecad qoyan (Bagirova et al., 2001). Si kastaba ha noqotee, haddii hababka dib-u-qaadista (galmada ama galmada) ay ku dhacdo dadka, markaa waxaa suurtagal ah in la isku daro isbeddelada kala duwan ee waalidiinta ee isku-darka isku-darka iyo dardar-gelinta hawsha. Sidaa darteed, dadka panmix waxay helaan iska caabin ka dhaqso badan dadka agamic, iyo tan dambe, dadka aan haysan caqabadaha is-waafajinta daaqsinka ayaa ka dhakhso badan dadka ay kala sooceen caqabadahaas. Marka la eego, joogitaanka noocyada kala duwan ee dadweynaha ee ku kala duwan noocyada isugeynta waxay dardargelineysaa geeddi-socodka helitaanka iska caabbinta fungicides oligosite.
Qodobbada labaad iyo saddexaad ayaan wax ku darsanayn isku-uruurinta degdegga ah ee noocyada u adkaysta dimethomorph ee dadka. Isbeddel kasta oo dambe wuxuu ku dhowaad laba jibbaaraa iska caabbinta, taas oo ah mid aan muhiimad lahayn, isla mar ahaantaana hoos u dhigaysa heerka koritaanka ee deegaan dabiici ah iyo gacan ka hadal (Bagirova et al., 2001; Stem, Kirk, 2004). Waxaa laga yaabaa in tani ay tahay sababta ficil ahaan u jirin noocyo iska caabin ah oo ka mid ah noocyada dabiiciga ah ee P. infestans, xitaa kuwa laga soo ururiyay dhirta baradho ee lagu daaweeyay dimethomorph.
Dad lagu daweeyey fungicide oligosite ayaa sidoo kale ka koobnaan doona laba kooxood oo noocyo ah: kuwa aan la xiriirin fungicide, oo sidaas darteed ma aysan bedelin astaamihii hore (haddii noocyo iska caabin ah laga helo kooxdan, ma ururi doonaan sababtoo ah gardarrada sare iyo tartan ee noocyo xasaasi ah), iyo noocyo taabasho leh suugaanta halista ah ee fungicide. Waxay ka mid tahay kuwa dambe in uruurinta noocyo adkaysi leh ay suurtagal tahay, maxaa yeelay halkan waxay kaga faa'iidaystaan kuwa xasaasiga ah.
Sidaa darteed, marka la isticmaalayo fungicides oligosite, maahan daaweyn dhameystiran oo muhiim u ah uruurinta sare ee daroogada, dhowr jeer ka sarreysa qiyaasta halista ah, sababtoo ah mutagenesis tallaabo tallaabo ah, iska caabinta bilowga ah ee noocyada is beddelay ayaa hooseeya.
Ugu dambeyntiina, is-beddelka iska caabbinta funaangooyinka monosite-ka ah ayaa ah mid aad u muujiya, taas oo ah, hal isbadal ayaa soo sheegi kara heer sare oo iska caabin ah, illaa iyo gabi ahaanba luminta dareenka. Sidaa darteed, kororka caabbinta dadka waxay u dhacdaa si dhakhso leh.
Tusaalaha sunta noocaas ah waa fenylamides, oo ay kujirto sunta ugu caansan, metalaxyl. Is-beddelka iska-caabbinta ayaa ku soo noqnoqonaya iyadoo aad u sarreeya, iyo heerka iska caabbinta mutantiyayaashu aad u sarreeyaan - waxay ka sarreysaa culeyska xasaasiga ah ee gaaraya kun ama in ka badan (Derevyagina et al., 1993). In kasta oo heerka koritaanka iyo gardarrada mutantada u adkaysata ay hoos u dhacayso marka loo eego asalka dhimashada noocyo nugul oo ka yimaad masaaji nidaam, tirada dadka iska caabbinta ah ayaa si xawli ah u koraysa, marka la barbar dhigo, gardarradeedu way sii kordheysaa. Sidaa darteed, ka dib dhowr sano oo la isticmaalayey fungicide, gardarrada noocyada iska caabinta kaliya ma sinnaan karto gardarrada kuwa xasaasiga ah, laakiin sidoo kale wey dhaafi kartaa (Derevyagina iyo Dyakov, 1992).
Saameynta ku-soo noqoshada galmada
Maaddaama dhacdooyinka soo noqnoqda ee nooca A2 ee isu-tagga ah ee ku dhaca P. dadka ku dhex nool ay ku soo beegantay isticmaalka xoogga leh ee metalaxyl ee looga soo horjeedo nabarrada soo daahay, waxaa la soo jeediyay in metalaxyl ay keento u-beddelashada nooca isku-darka. P. parasitica, beddelaad noocan oo kale ah oo hoos timaada ficilka chloroneb iyo metalaxyl ayaa si tijaabo ah loo cadeeyay (Ko, 1994). Hal marin oo dhexdhexaad ah oo leh feejignaan hoose oo metalaxyl ah ayaa horseeday soo bixitaanka go'doominta homothallic ee culeyska P. infestans xasaasi u ah metalaxyl oo leh nooca isku-xirnaanta A1 (Savenkova iyo Cherepnikova-Anikina, 2002). Intii lagu jiray marinnada soo socda ee warbaahinta ee leh metalaxyl badan, ma jirin hal gooni ah oo nooca A2 lammaanaha ah la ogaaday, hase yeeshe, inta badan go'doominnada, markii laga tallaabayo go'doommada A2, halkii ayspore-ku ahaayeen, waxay sameeyeen isku-uruursi mycelium fool xun oo nadiif ahaa. Marinnada culeyska adkeysiga leh ee leh nooca isku-darka A2 ee leh miisaanka leh metalaxyl badan ayaa noo oggolaatay inaan ogaanno saddex nooc oo isbeddellada nooca isku-dhafka ah: 1) nadiifnimo dhammaystiran markii laga gudbo A1 iyo A2 go'doominno; 2) homotallism (sameynta oospores ee monoculture); 3) u-beddelashada nooca A2 nooca mating-ka oo loo beddelay A1. Marka, metalaxyl wuxuu sababi karaa isbeddelo ku dhaca noocyada isutagga ee P. dadka ku dhaca iyo sidaas awgeed, soo noqoshada galmada iyaga dhexdeeda.
Saameyn ku yeelashada isku soo noqoshada daaqsinka
Qaar ka mid ah hiddo-wadahayaasha iska caabbinta antibiyootigga ayaa kordhiyay inta jeer ee heterokaryotization hyphal iyo diblumaasiyadda nukliyeerka (Poedinok iyo Dyakov, 1981). Sidii hore loo xusay, heterokaryotization of hyphae inta lagu guda jiro isku dhafka noocyada kala duwan ee P. infestans waxay dhacdaa marar aad u yar sababtoo ah ifafaale la'aanta daaqsinka ee funguskan. Si kastaba ha noqotee, hiddo-wadaha iska caabbinta antibiyootikada qaarkood wuxuu yeelan karaa waxyeelo, oo lagu muujiyo ka-adkaanta is-waafajinta daaqsinka. Hantidan waxaa lahaan jiray hiddo-wadaha iska caabinta 'StStomycin gene' 1S-1 mutant. Joogitaanka mutantanada noocan oo kale ah ee ku nool beeroweynta phytophthora waxay kordhin kartaa socodka hiddo-wadaha u dhexeeya noocyada waxayna dardargelinaysaa la qabsiga dadka oo dhan noocyada cusub ama fungicides.
Qaar ka mid ah fungic-yada iyo antibiyootigyada ayaa saameyn ku yeelan kara inta jeer ee isku soo noqoshada mitotic, taas oo sidoo kale wax ka beddeli karta jaangooyooyinyada caadiga ah ee dadweynaha. Benomyl-ka loo adeegsado fungicide-ka ee sida ballaaran loo adeegsado waxay ku xidhan tahay beta-tubulin, oo ah borotiin ay ka samaysato microtubules-ka cytoskeleton, halkaasna ay ku carqaladeynayso geeddi-socodka kala-soocidda koromosoomka ee ku dhaca xasaasiyadda 'mitosis', taasoo sii kordheysa soo noqnoqoshada mitotic (Hastie, 1970).
The fungicide para-fluorophenylalanine, oo loogu talagalay in lagu daaweeyo cudurka Dutch ee elms, ayaa leh isla hantida. Para-fluorophenylalanine waxay kordhisay soo noqnoqoshada dib-isu-soo-celinta heterozygous diploids P. infestans (Poedinok et al., 1982).
Isbeddelada isbeddelka ee ku dhaca halabuurka genotypic ee dadweynaha ee wareegga nolosha ee P. infestans
Wareegga koritaanka caadiga ah ee P. infestans aagga kuleylka leh wuxuu ka kooban yahay 4 weji.
1) Wejiga koritaanka jibbaaranaha ee dadka (polycyclic phase) oo leh jiillo gaagaaban. Wejigani wuxuu badanaa bilaabmaa bisha Luulyo wuxuuna socdaa 1,5-2 bilood.
2) Wajiga joojinta koritaanka dadka iyadoo ay ugu wacan tahay hoos u dhac fara badan oo ku yimid saamiga unugyada taabanaya ama bilawga xaaladaha cimilada ee aan fiicnayn. Marxaladdaan beeraha fulisa hore u saarista caleen horay u goosashada waxay ka dhacdaa wareegga sanadlaha.
3) Wajiga jiilaalka ee digriiga, oo ay weheliso hoos u dhac weyn oo ku yimid tirada dadka sababtoo ah infekshin kadis ah oo ku dhaca digriiga, horumarka gaabiska ah ee infekshanka ku jiro, maqnaanshaha dib-u-qaadista cudurka ee digriiga, qurminta iyo qaboojinta digriiga ay saameysay xaaladaha caadiga ah ee keydinta.
4) Wajiga horumarka gaabiska ah ee carrada iyo geedaha (wajiga monocyclic), kaas oo muddada jiilku gaari karo bil ama ka badan (dabayaaqadii Maajo - horraantii Julaay). Caadi ahaan waqtigan, caleemaha jirran way adag tahay in la ogaado, xitaa iyadoo fiiro gaar ah leh.
Wejiga koritaanka dadweynaha ee jaban (polycyclic phase)
Indha indheyn fara badan (Pshedetskaya, Kozubova, 1969; Borisenok, 1969; Osh, 1969; Dyakov, Suprun, 1984; Rybakova, Dyakov, 1990) waxay muujiyeen in bilowga epiphytoty, noocyo jilicsan oo xoogaa yar oo gardarro u badan, kuwaas oo markii dambe lagu beddelay kuwa ka sii daran oo dagaal badan. xaddiga koritaanka ee gardarrada dadweynuhu waa mid sareysa, oo u adkeysi yar noocyada kala duwan ee dhirta martida loo yahay.
Maaddaama dadku sii kordhayo, isku-darka labada hiddo-wadaha muhiimka ah ee loo soo bandhigay noocyo ganacsi (R1-R4) iyo xulasho dhexdhexaad ah (R5-R11) ayaa kordha. Sidaa darteed, dadka ku dhow magaalada Moskow 1993, celceliska feyraska laga soo bilaabo dabayaaqadii bishii Luulyo ilaa bartamihii Agoosto wuxuu ka kordhay 8,2 ilaa 9,4, kororka ugu weyna waxaa lagu arkay hiddo wadayaasha dhexdhexaadka ah ee Virulence R5 (laga bilaabo 31 ilaa 86% ee isku darka culeyska) (Smirnov, 1996) ).
Hoos u dhaca heerka koritaanka dadweynaha waxaa weheliya hoos u dhac ku yimaada waxqabadka dulin ee dadweynaha. Sidaa darteed, sanadaha murugada leh, tirada guud ee jinsiyadaha iyo saamiga jinsiyadaha aadka u daran waxay ka hooseeyaan kuwa epiphytotic-ka (Borisenok, 1969). Hadday sare u kacdo xaaladaha cimilada epiphytotic-ka waxay isu beddelaysaa mid aan fiicnayn oo loogu talagalay burburka goor dambe iyo cayayaanka baradhada ayaa hoos u dhacaya, isku-soo-ururinta isku-dhafan ee isku-dhafan oo isku-dhafan ayaa sidoo kale hoos u dhacaya (Rybakova et al., 1987).
Kordhinta soo noqnoqoshada hiddo-wadaha saameynaya qallafsanaanta iyo gardarrada dadweynaha ayaa laga yaabaa inay sabab u tahay xulashada dheellitirro dheellitiran oo xoog badan oo isku-dhafan oo ku dhaca dadka isku-dhafan. Si loo muujiyo xulashada, hab loogu talagalay falanqaynta is-beddelka dhexdhexaadka ah ayaa la soo saaray, kaas oo si guul leh loogu adeegsaday dadka ugu badan ee khamiirka (Adams et al., 1985) iyo Fusarium graminearum (Wiebe et al., 1995).
Soo noqnoqoshada mutantooyinka u adkaysta blasticidin S ee ku jira dadka ku nool P. infestans ayaa hoos u dhacay barbar socda koritaanka gardarrada dadweynaha, taas oo muujinaysa isbeddel ku yimaadda xirmooyinka ugu badan inta lagu jiro koritaanka dadka (Rybakova et al., 1987).
Wajiga Wintering ee digriiga
Xilliga qaboobaha ee digriiga baradhada, kacsanaanta iyo gacan ka hadalka ee P. infestans muruqyada ayaa yaraada, hoos u dhaca firirku wuxuu u dhacaa si tartiib tartiib ah gardarada (Rybakova iyo Dyakov, 1990). Sida muuqata, xaaladaha ku habboon koritaanka degdegga ah ee dadweynaha (xulashada), hiddo-wadayaasha "dheeraadka ah" iyo fal-xumada sare ayaa faa'iido leh, sidaa darteed horumarinta epiphytotics waxaa weheliya xulashada kuwa ugu halista badan uguna xanaaqa badan. Xaaladaha cufnaanta deegaanka, marka aan ahayn heerka taranka, laakiin ku adkaysiga jiritaanka xaaladaha aan fiicnayn (xulashada) ayaa door muhiim ah ka ciyaara, "hiddo-wadeyaasha" dheeraadka ah ee qallafsanaanta iyo dagaalku waxay yareeyaan jirdhiska, xirmooyinka hiddo-wadahaani waa kuwa ugu horreeya ee dhinta, si celceliska gardarrada iyo kacsanaanta dadka ayaa hoos u dhacaysa.
Wajiga dhirtu ku jirto ciida
Marxaladani waa tan ugu yaabka badan ee wareegga nolosha (Andrivon, 1995). Jiritaankeeda waxaa lagu dhajiyay si mala awaal ah - sababtoo ah xog la'aanta ku saabsan waxa ku dhaca cudur-sidaha muddo dheer (mararka qaarkood in ka badan hal bil) - laga soo bilaabo soo bixitaanka geedo baradho illaa iyo muuqaalka dhibcaha ugu horreeya cudurka iyaga ku dhaca. Iyada oo ku saleysan fiiro gaar ah iyo tijaabooyin, habdhaqanka fungus ee muddadan nolosha ayaa dib loo dhisay (Hirst iyo Stedman, 1960; Boguslavskaya, Filippov, 1976).
Sporulation of fungus waxay ku samaysmi kartaa digriiga cudurka ku haya ciidda. Isboortiga ka dhashay waxay ku dhalaalaan hiddo, taas oo ku noolaan kara muddo dheer ciidda. Aasaasiga ah (oo lagu sameeyay digriiga) iyo kan labaad (mycelium-ka carrada) kudka ayaa kor u kaca dusha sare ee ciidda iyadoo la adeegsanayo qulqulo qulqulaya, laakiin waxay helayaan awood ay ku kala qaadaan baradhada kaliya markay caleemaha hoose hoos u soo dhacaan oo ay taabtaan dusha ciidda. Caleemaha noocan oo kale ah (kuwaas oo ah, dhibcaha ugu horreeya ee cudurka ayaa laga helaa iyaga) isla markiiba ma sameysma, laakiin ka dib koritaanka daba dheeraaday iyo horumarka dusha baradhada.
Sidaa darteed, wareegga nolosha ee P. infestans, wejiga dhirta saprotrophic sidoo kale wuu jiri karaa. Haddii marxaladda dulin ee wareegga nolosha ee dagaalku yahay qaybta ugu muhiimsan ee jirdhiska, markaa xulashada wejiga saprotrophic waxaa loogu talagalay in lagu yareeyo guryaha dulinka ah, sida tijaabada loogu muujiyey fangaska phytopathogenic qaar ka mid ah (eeg Carson, 1993). Sidaa darteed, marxaladan wareegga, guryaha gardarrada ah waa in si xoog leh loo lumiyaa. Laakiin illaa iyo hadda wax tijaabooyin toos ah lama samayn si loo xaqiijiyo fikradaha kor ku xusan.
Isbeddelada xilliyeed ma saameeyaan oo keliya sifooyinka cudurada ee cudurka 'P. infestans', laakiin sidoo kale waxay iska caabiyaan fungicides, oo ku soo baxda wajiga polycyclic (inta lagu jiro epiphytoties), iyo hoos u dhaca inta lagu jiro keydinta xilliga qaboobaha (Derevyagina et al., 1991; Kadish iyo Cohen, 1992). Hoos u dhac gaar ah oo iska caabin ah metalaxyl ayaa lagu arkay inta udhaxeysa abuurka digriiga ay saameysey iyo muuqaalka dhibcaha ugu horeeya ee cudurka beerta.
Takhasus gaar ah iyo isbeddelkiisa
P. infestans wuxuu cudurrada faafa ku keenayaa laba dalag oo ganacsi muhiim u ah, baradhada iyo yaanyada. Epiphytoties-ka baradhada waxay bilaabatay waxyar kadib markii fangaska soo gashay aagag cusub. Guuldarada yaanyada ayaa sidoo kale la xusay waxyar uun ka dib muuqaalka caabuqa ee baradhada, laakiin epiphytoties on yaanyo ayaa la ogaaday kaliya boqol sano ka dib - bartamihii qarnigii XNUMXaad. Waa tan waxa Hallegli iyo Niederhauser ka qorayaan jabkii yaanyada ee USA
(1962): “Ilaa 100 sano kadib epiphytoty-kii daran ee 1845, in yar ama kudhowaad iskudayo lama sameyn si loo helo noocyo yaanyo adkaysi leh. In kasta oo goor dambe duqeynta lagu duubay yaanyada horaantii sanadkii 1848, hadana ma aysan noqonin wax si weyn looga fiirsado oo ay ka soo baxaan dhirta ilaa iyo markii uu cudurku ka dillaacay 1946. Dhulkii Ruushka duqayn dambe oo yaanyo ayaa laga diiwaan galiyay qarnigii 60aad. “Muddo dheer, cilmi baarayaashu dhag jalaq uma siinin cudurkan, maadaama uusan keenin waxyeello dhaqaale oo weyn. Laakiin 70-meeyadii iyo 1979-meeyadii. Qarnigii XNUMX-aad ee epiphytoties ee duqeynta goor dambe ee yaanyada ayaa sidoo kale lagu arkay Midowgii Soofiyeeti, gaar ahaan gobolka Volga Hoose, Ukraine, Waqooyiga Caucasus, Moldova ... ”(Balashova, XNUMX).
Tan iyo waagaas, duqeynta yaanyada ee duqeynta goor dambe waxay noqotey mid sanadle ah, oo ku faafta dhamaan dhulka warshadaha iyo beerashada guryaha waxayna u keentaa waxyeello dhaqaale oo baaxad weyn dalaggan. Maxaa dhacay? Muxuu muuqaalka koowaad ee dulinka baradhada iyo nabarada epiphytotic-ka ee dhaqankan kudhacay isku mar, halka ay qarnigii ku qaadatay inuu epiphytotic-ka ka muuqdo yaanyada? Kala duwanaanshahani waxay taageeraan reer Mexico halkii laga heli lahaa ilaha Koonfurta Ameerika ee infekshanka. Haddii noocyada 'Phytophthora infestans' ay u samaysteen sidii dulin ka mid ah noocyada tuber-ka ee meksiko ee hiddo-wadaha Solanum, markaa waa la fahmi karaa sababta baradhada la beeray ee ka tirsan isla qeyb ka mid ah genus sida noocyada reer Mexico ay si aad ah u saameeyeen, laakiin ay ugu wacan tahay maqnaanshaha is-beddel la'aanta dulinka, oo aan soo saarin farsamooyin iska caabin gaar ah oo aan macno lahayn
Yaanyada waxay ka tirsan tahay qeyb ka duwan genus, nooca sarrifkiisa wuxuu ku kala duwan yahay noocyada tuubbada, sidaas darteed, in kasta oo ay xaqiiqda tahay in yaanyada aysan ka baxsaneyn takhasuska cuntada ee P. infestans, xoojinta guuldaradiisu kuma filneyn khasaaro dhaqaale oo daran.
Soo bixitaanka epiphytoties ee yaanyada waxaa sabab u ah isbedelada hidaha ee halista ah ee dulinka, taas oo kordhisay la qabsiga (cudurada) markii la daweeyo. Waxaan aaminsanahay in foomka cusub ee ku takhasusay parasiticinta tamaandhada uu yahay T1-ka oo uu sharaxay M. Gallegly, oo saameynaya noocyo ka mid ah yaanyada cherry (Red Cherry, Ottawa), adkaysi u leh tartanka T0 ee ku baahsan baradhada (Gallegly, 1952). Sida muuqata, isbadal ku yimid (ama isbadal isdaba joog ah) oo u rogay T0-ga tartanka T1 una horseeday muuqaalka clones aad loogu habeeyay jabkii yaanyada. Sida had iyo jeer dhacda, kororka cudur ku dhaca hal marti soor waxaa weheliya hoos u dhac ku yimaada mid kale, taasi waa, bilow gaar ah, oo aan weli dhammaystirnayn takhasus gaar ah - baradhada (T0 tartanka) iyo tamaandhada (tartanka T1).
Maxaa caddeyn u ah male-awaalkan?
- Ku dhaca baradhada iyo yaanyada. Caleemaha yaanyada, tartanka T1 ayaa ku badan, halka caleemaha baradhada ay dhif tahay. Sida laga soo xigtay S.F.Bagirova iyo T.A. Oreshonkova (aan la daabicin) gobolka Moskow 1991-1992, dhacdada tartanka T1 ee beero baradho wuxuu ahaa 0%, iyo in yaanyo lagu beero - 100%; 1993-1995 - 33% iyo 90%, siday u kala horreeyaan; 2001 - 0% iyo 67%. Xog la mid ah ayaa laga helay Israel (Cohen, 2002). Tijaabooyin ku saabsan infekshinka digriiga baradhada oo leh go'doominta jinsiyada T1 iyo isku dhafka go'doominta T0 iyo T1 ayaa muujisay in go'doominta jinsiyada T1 ay si liidata ugu keydsan tahay digriiga waxaana lagu beddelay go'doominta jinsiyadda T0 (Dyakov et al., 1975; Rybakova, 1988).
2) Dynamics ee tartanka T1 ee beerashada yaanyada. Cudurka asaasiga ah ee caleemaha yaanyada waxaa lagu fuliyaa go'doominta jinsiga T0, oo ku badan falanqaynta cudurka dhibcaha ugu horreeya ee lagu sameeyay caleemaha. Tani waxay xaqiijineysaa qorshaha guud ee la oggol yahay ee socdaalka dulinka: Cudurka ugu weyn ee infekshanka ka yimaada baradhada wuxuu ka kooban yahay tartanka T0, si kastaba ha noqotee, in yar oo ka mid ah T1 clones oo lagu keydiyo baradhada, marna yaanyada, ayaa barakiciya tartanka T0 oo ku urursada dhammaadka muddada epiphytotic. Waxa kale oo suurtagal ah inay jirto ilo kale oo caabuqa caleemaha yaanyo leh tartanka T1, oo aan u xoog badnayn sida digriiga iyo caleemaha baradhada, laakiin joogto ah. Sidaa darteed, ilahaani waxay saameyn daciif ah ku leeyihiin qaab dhismeedka hidde ee dadka qaadsiinaya yaanyada, laakiin markii dambe ayaa go'aamisa uruurinta tartanka T1 (Rybakova, 1988; Dyakov et al., 1994).
3) Ku dirir baradho iyo yaanyo. Cudurka macmalka ah ee yaanyada iyo caleemaha baradhada oo ay kujiraan noocyada jinsiyadaha T0 iyo T1 waxay muujiyeen in kuwii hore ay uga bareeran yihiin baradhada marka loo eego ta yaanyada, tan dambena ay uga yaryihiin yaanyo marka loo eego baradhada. Kala duwanaanshahan waxaa lagu muujiyey barakicinta go'doominta jinsiyad aan "gaar ahayn" oo ka timid dad isku dhafan inta lagu jiro marinnada caleemaha ee aqalka dhirta lagu koriyo (Dyakov et al., 1975) iyo goos gooska (Leberton et al., 1999); kala duwanaanshaha culeyska faafa ee ugu yar, waqtiga daahitaanka, cabirka dhibcaha faafa iyo wax soo saarka cayayaanka (Rybakova, 1988; Dyakov et al., 1994; Legard et al., 1995; Forbes et al., 1997; Oyarzun et al., 1998; Leberton et al., 1999; Vega-Sanchez et al., 2000; Knapova, Gisi, 2002; Sussuna et al., 2004).
Ku xadgudubka go'doominta jinsiyada T1 ee beeraha yaanyada oo aan lahayn hiddo-wadayaal iska caabin ah ayaa aad u sarreeya in go'doominnadaasi ay ku caleema baxaan caleemaha sida nafaqada leh iyada oo aan la nadiifin unugyada cudurka qaba (Dyakov et al., 1975; Vega-Sanchez et al., 2000).
4) Firfircoonida baradhada iyo yaanyada. Tartanka T1 wuxuu ku dhacaa noocyo yaanyo cherry ah oo leh hidde-wadaha caabbinta Ph1, halka tartanka T0 uusan awood u lahayn inuu ku dhaco noocyadan, i.e. wuxuu leeyahay firir cidhiidhi ah. Marka la eego kala duwanaanshaha
R-hiddo-wadaha baradhada ayaa isku khilaafsan, i.e. noocyada laga soocay caleemaha yaanyada way ka dhib yar yihiin noocyada "baradhada" (Shaxda 11).
5) Calaamadaha dhexdhexaadka ah. Falanqaynta astaamaha dhexdhexaadka ah ee dadka P. infestans parasitizing on baradhada iyo yaanyada ayaa sidoo kale ka markhaati furaya xulashada intraspecific intrapection. Dadka reer Brazil ee P. infestans, go'doominta caleenta yaanyada waxaa iska lahaa safka loo yaqaan 'clonal line US-1', kuwa caleenta baradhada ka tirsanna waxay ahaayeen laynka BR-1 (Suassuna et al., 2004). Magaalada Florida (USA), laga soo bilaabo 1994, clone US-90 wuxuu bilaabay inuu ku adkaado baradhada (oo ay dhacdo in ka badan 8%), iyo isku darka US-11 iyo US-17 ee yaanyada, iyo go'doominta dambe waxay uga sii daran yihiin yaanyada tan baradhada (Weingartner , Tombolato, 2004). Kala duwanaansho weyn oo ku saabsan isdabajoogga genotype (faraha faraha DNA) ee baradhada iyo yaanyada go'doomin ayaa loo sameeyay 1200 P. noocyada infestans ee laga soo ururiyay Mareykanka laga bilaabo 1989 ilaa 1995 (Deahl et al., 1995).
Adeegsiga habka loo yaqaan 'AFLP' ayaa suurta galisay in lakala saaro 74 nooc oo laga soo ururiyey caleemaha baradhada iyo yaanyada sanadihii 1996-1997. Faransiiska iyo Switzerland, oo ka kooban 7 koox. Noocyada baradhada iyo yaanyada si cad ugama duwanaan, laakiin noocyada "baradhada" waxay ahaayeen kuwo hidde ahaan aad uga duwan kuwii "yaanyo". Kuwii hore waxaa laga helay dhamaan toddobada qaybood, tan dambena, kaliya afar, taas oo muujinaysa hiddo-wadaha takhasuska leh ee kan dambe (Knapova iyo Gisi, 2002).
6) Farsamooyinka go'doominta. Haddii dadka ku dul noolaha laba nooc oo dhirta martida loo yahay ay u xuubsiibtaan cidhiidhiga takhasuska gaar u ah martigaliyahooda, markaa farsamooyin kala duwan oo ka hor iyo postmeiotic ah ayaa soo baxa oo ka hortaga isdhaafsiga hidda-wadaha (Dyakov iyo Lekomtseva, 1984).
Daraasado dhowr ah ayaa baaray saameynta isha ilaha waalidku ku leeyihiin waxtarnimada isku-darka. Markii noocyo ka go'doonsanaa noocyo kala duwan oo hiddo-wadaha ah loo gudbiyo Ecuador (Oliva et al., 2002), waxaa la ogaaday in noocyada nooca A2 ee isku-dhafka ah ee ka imanaya habeennada duurjoogta ah (khadka dhexe ee EC-2) ay uga gudbeen kuwa ugu xun noocyada tamaandhada (khadka EC -3), oo sida ugu wax ku oolsan looga gudbey culeyska baradhada (EC-1).
Dhammaan iskudhafyadii la helay ayaa la ogaaday inay yihiin kuwo aan cudur lahayn. Qorayaashu waxay rumeysan yihiin in boqolkiiba hooseeya ee isugeynta iyo yareynta cudurada ku dhaca isugeynta ay sabab u tahay farsamooyinka postmeiotic ee go'doominta taranka ee dadka.
Tijaabooyinka Bagirova et al. (1998), tiro badan oo ah baradhada iyo noocyada yaanyada ayaa lagu tallaalay astaamaha jinsiyadaha T0 iyo T1. Iskutallaabyada ugu waxsoosaarka wanaagsan ee noocyada T1xT1 ee laga soocay yaanyada (36 oospor oo ku jira mekroskoobka aragtida, 44% ee biqilka oospore), kuwa ugu wax ku oolka ah waxay ahaayeen iskutallaabyada T0xT1 jinsiyadaha laga soocay martida kala duwan (tiro yar oo soosaarayaasha soo koraya iyo kuwa soobaxay, tiro aad u tiro badan oo ah ubaxyo ilmo soo ridid ah iyo kuwa aan horumarin) ... Waxtarka iskutallaabta udhaxeysa go'doominta jinsiyada T0 ee laga soocay baradhada waxay ahayd dhexdhexaad. Maaddaama jirka ugu muhiimsan ee noocyada T0 uu saameyn ku yeesho baradhada, wuxuu leeyahay ilo lagu kalsoonaan karo oo xilliga qaboobaha - digriiga baradhada, taas oo ka dhalatay taas oo ahmiyadda ay daaqadu u leedahay sida qabowga unugyada faafa ee dadka ka yimaada baradhada. Foomka la yareeyay ee "yaanyada" wuxuu awoodaa inuu xilliga jiilaalka ku jiito yaanyo qaab loo yaqaan 'oospores' (hoos fiiri) sidaas darteedna wuxuu hayaa wax soo saar sare oo habka galmada ah. Bacriminta sare awgeed, T1 waxay leedahay awood madax-bannaan oo loogu talagalay infekshanka asaasiga ah ee yaanyada. Natiijooyinka ay heleen Knapova et al. (Knapova et al., 2002) ayaa si isku mid ah loogu fasiri karaa. Iskutallaabyada noocyada laga soocay baradhada oo leh noocyo laga soo qaaday yaanyo ayaa bixiyay tirada ugu badan oospore - 13,8 halkii sq.mm. dhexdhexaad ah (oo ku faafaya 5-19) iyo boqolkiiba dhexdhexaad ee biqilka oospores (6,3 oo ku faafaya 0-24). Isgoysyada noocyada ah ee laga soocay yaanyada ayaa bixiyay boqolkiiba ugu hooseeya oospore (7,6 oo ku faafaya 4-12) oo leh boqolkiiba ugu badan ee biqilkooda (10,8). Isgoysyada udhaxeeya muruqyada laga soocay baradhada waxay siisay tiro dhexdhexaad ah oospore ah (8,6 oo leh firdhin badan oo xog ah - 0-30) iyo boqolkiiba ugu hooseeya ee biqilka oospores (2,7). Sidaa awgeed, noocyada ka soo baxa baradhada ayaa ka wax-tar yar marka loo eego kuwa laga helo yaanyada, laakiin iskutallaabyada is-dhexgalka dadku ma keenin natiijooyin ka xun kuwa dadka ku dhex jira. Waa suurtagal in kala duwanaanshaha xogta kor ku xusan ee Bagirova et al. waxaa lagu sharaxay xaqiiqda ah in cilmi baarayaal Ruush ah ay la shaqeeyeen noocyo go'doonsan horaantii 90-meeyadii qarnigii labaatanaad, iyo cilmi baarayaasha Switzerland - oo ay lajireen noocyo dhammaadkii 90-maadkii.
Saldhigga taranka hooseeya wuxuu noqon karaa heteroploidy ee noocyada. Haddii dadka reer Mexico, halkaasoo habka galmada iyo infekshanka asaasiga ah ee loo yaqaan 'oospore progeny' ay yihiin kuwo joogto ah, inta badan noocyada daraasadda ee P. Infestans waa diploid, markaa waddamada Old World intrapopulation polymorphism of ploidy ayaa lagu arkay (di-, tri- iyo tetraploid muruqyada, iyo sidoo kale noocyada heterokaryotic oo leh heteroploid nuclei) , iyo noocyo leh noocyo kala duwan oo isugeyn, i.e. si wada jir ah u bacriminaya, kuna kala duwan khiyaanada nukliyeerka (Therrien et al., 1989, 1990; Whittaker et al., 1992; Ritch, Daggett, 1995). Kala duwanaanta nuclei ee antheridia iyo oogonia waxay sababi kartaa taranka hooseeya.
Isweydaarsiga nukliyeerka ee udhaxeeya hyphae inta lagu jiro anastomoses, tan waxaa kahortaga iswaafaq la'aanta daaqsinka, taas oo uqeybisa dadka asexual noocyo badan oo hidde ahaan go'doonsan (Poedinok iyo Dyakov, 1987; Gorbunova et al., 1989; Anikina et al., 1997b).
7) Isku dhafka dadka. Xogta kor ku xusan waxay muujineysaa isku-dhafid u dhexeeya "baradho" iyo "yaanyo" noocyada infestans ee P. waa suurtagal. Dib-u-caabuq dib-u-soocid ah ee marti-geliyeyaal kala duwan ayaa sidoo kale suurtagal ah, in kasta oo ay jirto dagaal-yari hoos u dhacday.
Daraasad ku saabsan astaamaha dadka ee ku go'doonsan meelaha baradhada iyo yaanyada ku dhow ee 1993 waxay muujisay in ku dhowaad rubuc ka mid ah go'doonnada laga go'doomiyey caleemaha yaanyo laga soo wareejiyey beero baradho deris la ah (Dolgova et al., 1997). Aragti ahaan, waxaa loo qaadan karaa in kala duwanaanta dadka ee laba marti-qaade ay kordhin doonto oo ay horseedi doonto soo bixitaanka qaabab khaas ah oo iskudhaf ah (f.sp. baradho iyo yaanyo f.sp.), gaar ahaan maadaama oospore ku sii jiri karo qashinka dhirta (Drenth et al., 1995) ; Bagirova, Dyakov, 1998) iyo iniin yaanyo ah (Rubin et al., 2001). Sidaa awgeed, yaanyada xilligan waxay leeyihiin il dib-u-cusboonaysiinta gu'ga oo ka madax bannaan digriiga baradhada.
Si kastaba ha noqotee, wax walba waxay u dhaceen si kala duwan. Kufsiga oospore-ka ayaa u oggolaaday dulinku inuu ka fogaado marxaladda ugu yar ee wareegga noloshiisa - marxaladda monocyclic ee dhirta carrada, xilligaas oo ay hoos u dhacayaan sifooyinka dulinku, kuwaas oo si tartiib tartiib ah loogu soo celiyo wejiga polycyclic-ka xagaaga.
Jadwalka 11. Soo noqnoqdayaasha hiddo-wadayaasha noocyada kala duwan ee baradhada noocyada kala duwan ee P. infestans
dalka | Sannad | Celceliska tirada hiddo-wadayaasha firirrada ku dhaca noocyada | qoraaga | |
ka soo baradho | laga bilaabo yaanyo | |||
France | 1995 | 4.4 | 3.3 | Leberton et al., 1999 |
1996 | 4.8 | 3.6 | Leberton, Andrivon, 1998 | |
France, Switzerland | 1996-97 | 6.8 | 2.9 | Knapova, Gisi, 2002 |
United States | 1989-94 | 5 | 4.8 | Goodwin et al., 1995 |
USA, Zap. Washington DC | 1996 | 4.6 | 5 | Dorrance et al., 1999 |
1997 | 6.3 | 3.5 | " | |
Ecuador | 1993-95 | 7.1 | 1.3 | Oyarzun iyo al., 1998 |
Israa'iil | 1998 | 7 | 4.8 | Cohen, 2002 |
1999 | 6 | 5.7 | " | |
2000 | 6.7 | 6.1 | " | |
Ruushka, Mosk. gobolka | 1993 | 8.9 | 6.7 | Smirnov, 1996 |
Ruushka, gobollo kala duwan | 1995 | 9.4 | 8 | Kozlovskaya iyo kuwa kale. |
1997 | 9.2 | 9.2 | " | |
2000 | 8.7 | 4.8 | " |
Meelaha loo yaqaan 'zoosporangia Primary' iyo zoospores, oo biqiya oospores, waxay leeyihiin heer sare oo dhaqdhaqaaq dulin ah, gaar ahaan haddii oospores loo sameeyay si habsami leh iyada oo ay saameyn ku yeelaneyso pheromones-ka culeyska oo leh nooca ka soo horjeedka mataanka. Sidaa darteed, walxaha faafa ee ku saabsan geedo yaanyo ah oo ka soo baxay abuurka ay ku dhacdey oospore ayaa si aad ah u cudur u leh yaanyada iyo baradhada labadaba.
Isbeddeladani waxay keeneen dib-u-habeyn dadweyne oo kale, oo lagu muujiyey isbeddelada soo socda ee muhiimka ah marka laga eego aragtida faafa:
- Geedo yaanyo jeermis yeeshay waxay noqdeen isha muhiimka ah ee caabuqa asaasiga ah ee baradhada (Filippov, Ivanyuk, fariimo shaqsiyeed).
- Epiphytoties-ka baradhada waxay bilaabeen in la arko horaantii Juun, qiyaastii bil ka hor sidii caadiga ahayd.
- Beerista baradhada, boqolkiiba tartanka T1 ayaa kordhay, taas oo horey loogula kulmay halkaas qadar aan qiimo lahayn (Ulanova et al., 2003).
- Dhibaatooyinka laga soocay caleemaha yaanyada waxay joojiyeen inay ku kala duwanaadaan noocyada baradhada ee ku dhaca kaladuwanaanta kala duwanaanta hiddo wadayaasha waxayna bilaabeen inay dhaafeen noocyada “baradhada” ee ku xadgudubka kaliya yaanyada, laakiin sidoo kale baradhada (Lavrova et al., 2003; Ulanova et al. , 2003).
Sidaa darteed, halkii lagu kala duwanaan lahaa, waxaa jiray isku soo dhowaansho dadweyne, soo ifbaxa hal dad ah oo ku saabsan laba dhir oo martida loo yahay oo leh dabadhilifnimo sare iyo dagaal ku dhaca labada nooc.
gunaanad
Sidaa darteed, inkasta oo in ka badan 150 sano oo daraasad degdeg ah lagu sameynayay P. infestans, cilmiga bayoolajiga, oo ay ku jiraan bayoolajiga dadweynaha ee wakiilkan sababa ee cudurrada ugu muhiimsan ee dhirta solanaceous, haddana weli wax badan lama oga. Ma cadda sida marinka marxaladaha shaqsiyeed ee meertada nolosha ay u saameyneyso qaab dhismeedka dadka, waa maxay farsamooyinka hidde-wadaha ee kala duwanaanshaha kala duwanaanta rabshadaha iyo xanaaqa, waa maxay saamiga taranka iyo taranka taranka ee nidaamyada dadka dabiiciga ah, sidee isu-dheelitir la'aanta daaqsinka, maxay tahay doorka baradhada iyo yaanyada ee cudurka asaasiga ah ee dalagyadan iyo waa maxay saamaynta ay ku leeyihiin qaab dhismeedka dadka dulinka ah. Illaa iyo hadda, arrimaha muhiimka ah ee muhiimka ah sida farsamooyinka hidda-socodka ee beddelka gardarrada dulin ama nabaad-guurka caabbinta baradho aan macno lahayn lama xallin. Iyada oo la sii xoojinayo lana ballaadhinayo cilmi baarista baradhada goor dambe, dulinku wuxuu caqabado hor leh ku abuuraa cilmi baarayaasha. Si kastaba ha noqotee, hagaajinta awoodaha tijaabada ah, soo ifbaxa habab cusub oo habab loo adeegsado khalkhal gelinta hiddo-wadaha iyo borotiinnada ayaa noo oggolaanaya inaan rajo ka qabno xalka guul ee su'aalaha la soo saaray.
Maqaalka waxaa lagu daabacay joornaalka "Difaaca Baradhada" (Lambarka 3, 2017)