S.N. Elansky, L.Yu. Kokaeva, N.V. Statsyuk, Yu.T. Dyakov
Gabatarwar
Oomycete Phytophthora infestans (Mont.) De Bary - wakili mai haddasa cutar sankara, cuta mafi muhimmanci ta tattalin arziki dankali da tumatir - ya ja hankalin masu bincike daga kasashe daban daban fiye da karni da rabi. Ba zato ba tsammani ya bayyana a cikin Turai a tsakiyar karni na XNUMX, ya haifar da annobar dankalin turawa wanda ya kasance cikin ƙwaƙwalwar ajiyar ƙarni da yawa.
Har zuwa yanzu, ana kiransa sau da yawa "naman kaza na yunwar Irish". Kusan shekaru ɗari bayan annoba ta farko, an gano nau'ikan dankalin turawa na Mexico da ke jure wa cutar bushewa, an ci gaba da hanyoyin ƙetare su da dankalin da aka noma (Muller, 1935), kuma an sami nau'ikan farko da ke saurin kamuwa da cutar (Pushkarev, 1937). Koyaya, ba da daɗewa ba bayan fara kasuwancinsu na kasuwanci, jinsi na ƙarshen cututtukan cututtukan ƙwayoyin cuta waɗanda ke da lahani ga nau'o'in jurewa da aka tara. da kuma gabatar da sabbin kwayoyin halittar juriya daga dankalin Mexico na dangi zuwa iri sun fara rasa tasiri cikin sauri.
Kasawa tare da yin amfani da juriya mai tsauri (a tsaye) ya tilasta wa masu shayarwa neman hanyoyin da ke da rikitarwa na amfani da juriya ta rashin daidaituwar polygenic (kwance). A cikin 'yan shekarun nan, jinsi masu saurin tashin hankali sun fara taruwa a cikin daidaikun mutanen dake fama da cutar, wanda ke haifar da zagon kasa na mawuyacin yanayin gwagwarmaya. Zuwan nau'ikan da ke jure wa kayan gwari ya haifar da matsaloli game da amfani da sinadaran kare dankalin turawa.
Saboda manyan bambance-bambance tsakanin oomycetes da fungi a cikin abubuwan da ke cikin sinadarai, mafi kyawu, da kuma ciwan jiki, kayan gwari, musamman wadanda ake amfani dasu don kare tsirrai daga cututtukan fungal da yawa, basu da tasiri kan oomycetes.
Sabili da haka, a cikin kariya ta sinadarai daga ƙarshen buguwa, an yi amfani da yawa (har sau 12 a kowane lokaci ko sama da haka) ana amfani da fesawa tare da shirye-shiryen tuntuɓar fage mai fa'ida. Matakin juyi shine amfani da phenylamides, waɗanda suke da guba ga oomycetes kuma suna yaɗa cikin tsarin shuke-shuke. Koyaya, yawan amfani da su da sauri ya haifar da tarin ƙwayoyi masu juriya a cikin yawan fungal (Davidse et al., 1981), wanda ke da mahimmancin kariyar tsire-tsire. P. infestans kusan shine kawai kwayar cutar ta yanki mai laushi, cutar da ita wanda ba a iya raba shi da aikin gona ba tare da amfani da hanyoyin kariya na kariya ba (Van Bruggen, 1995).
Abubuwan da ke sama suna bayanin babban hankalin da masu bincike daga ƙasashe daban-daban ke bayarwa don nazarin yawan 'yan ƙanƙancin P., tasirin tasirin yalwar su da ƙwayoyin halittar su, gami da hanyoyin halittar canjin yanayi.
Rayuwar rayuwar R. INFESTANS
Oomycete Phytophthora infestans suna haɓaka ƙananan ƙwayoyin cuta tare da haustoria a cikin ganyen dankalin turawa. Ciyar da kyallen takarda na ganye, yana haifar da samuwar tabo mai duhu, wanda ya zama baƙi kuma ya ruɓe a cikin yanayin ruwa. Tare da shan kashi mai karfi, dukkan ganyen sun mutu. Bayan wani lokaci na ciyarwa, ana yin ɓarna a cikin mycelium - sporangiophores - wanda ke girma zuwa waje ta cikin stomata. A cikin yanayin ruwa, suna yin farar fure a kewayen aibobi a ƙasan ganyen. A ƙarshen sporangiophores, an kafa zoosporangia mai siffar lemun tsami, wanda ke ɓarkewa kuma ana ɗaukar shi ta hanyar watsa ruwan sama (Fig. 1). Samun saukad da ruwa a saman ganyen dankalin turawa, sporangia yayi girma tare da zoospores 6-8, wanda, bayan wani lokaci na motsi, an zagaye su, an rufe su da harsashi kuma sun tsiro da bututun tsiro. Ciyawar ta tsiro ta cikin stomata a cikin kayan ganye. A ƙarƙashin wasu yanayi, ɓarnatarwar na iya girma a cikin bututun girma kai tsaye zuwa cikin kayan ganye. A karkashin yanayi mai kyau, lokaci daga kamuwa da cuta zuwa samuwar sabon sporulation kwana 3-4 ne kawai.
Da zarar a ƙasa kuma an tace ta cikin ƙasa, sporangia na iya kamuwa da tubers. Tumani mai tsanani ya ruɓe yayin ajiya; a cikin rauni mai rauni, kamuwa da cuta na iya ci gaba har zuwa kakar ta gaba. Bugu da kari, wakili mai haddasa cutar sanyin lokaci na iya ci gaba a lokacin sanyi a cikin sifar feshin (zina mai dauke da walwala a lokacin hutu) a cikin ƙasa a kan tarkacen tsire-tsire da kuma yayan tumatir. Oospores yana samuwa ne akan gabobi masu rai na shuke-shuke lokacin da nau'ikan nau'ikan hanyoyin saduwa suka hadu da danshi mai yawa. A lokacin bazara, ana yin sporulation wanda ba na jinsi ba akan tubers da ke dauke da cutar da kuma ragowar tsire-tsire tare da oospores; zoospores suna shiga cikin ƙasa suna haifar da kamuwa da ƙananan ganyen tsire-tsire. A wasu lokuta, mycelium na iya girma daga tuber mai cutar tare da koren ɓangaren shukar kuma yawanci yakan bayyana a ɓangaren sama na tushe.
Babban banbanci tsakanin oomycetes da mafi yawan fungi sun ta'allaka ne akan fifikon yawan diplophase a cikin tsarin rayuwarsu tare da meiosis na gametic da germination na zygotes (oospores) ba tare da raguwar makaman nukiliya ba. Wannan fasalin, tare da maye gurbin heterotallism, maye gurbin bisexuality, zai zama kamar zai ba da damar yin amfani da hanyoyin ne don samar da hanyoyin da ake bi don nazarin yawan yawan eukaryotes (nazarin panmixia da rarrabuwa na yawan jama'a, intra- da yaduwar kwayar halittar mutane, da sauransu). Koyaya, abubuwa uku basu bada izinin canja waɗannan hanyoyin gabaɗaya a cikin binciken yawan mutanen inf infans.
1. Tare da wadatattun ospores, ana samarda wadataccen kayan abinci da kuma na parthenogenetic oospores a cikin jama'a (Fife da Shaw, 1992; Anikina et al., 1997a; Savenkova, Cherepnikoba-Anirina, 2002; Smirnov, 2003), kuma yawan samuwar su na iya isa ya yi tasiri. akan sakamakon gwaji.
2. Tsarin jima'i a cikin P. infestans yana bada muhimmiyar gudummawa ga mahimmancin girman yawan mutane, saboda naman gwari yana haifuwa akasari ta hanyar tsire-tsire masu tsire-tsire, yana samar da fiye da kashi 90% na sakamakon nazarin nau'in nau'in jima'i ta hanyar gargajiyar gargajiya akan matsakaiciyar matsakaiciyar ... Lokacin girma girma ƙarni da yawa na zafin jini na zamani (ci gaban cutar polycyclic). Oospores suna taka muhimmiyar rawa wajen kiyaye kwayar halitta a lokacin da babu kore shuke-shuke (a cikin hunturu) kuma a cikin kamuwa da cuta ta farko na tsire-tsire. Bayan haka, a lokacin bazara, haifuwa cikin ɗabi'a da ƙaruwa ko, akasin haka, raguwar adadin kwafin jikin mutum wanda ya tashi sakamakon sake haɗuwa da jima'i yana faruwa, wanda yawanci ya ƙaddara ta zaɓin wanda yafi dacewa. Sabili da haka, yawan adadin kwayoyi a cikin yawan mutane a farkon da ƙarshen epiphytotics na iya zama daban.
3. Tsarin da aka bayyana shine halayyar 'yan asalin garin P. mazauna ƙasarsu, Amurka ta tsakiya. A wasu yankuna na duniya, tsarin jima’i ba a san shi sama da shekaru 100 ba; mycelium mai cin ganyayyaki a cikin tubers dankalin turawa shi ne matakin sanyi. Tsarin rayuwa gabaɗaya ya kasance mai rikitarwa, kuma yaduwar ta kasance mai mahimmancin yanayi: kamuwa da cuta daga ƙwaya ɗaya da aka dasa shuki ya wuce zuwa ganye, wanda ya zama tushen cutar na farko, wanda zai iya haɗuwa da haɓakar cutar.
Don haka, a wasu yankuna akwai yuwuwar sauyawa tsakanin jima'i da zagayowar jima'i, yayin da a wasu - kawai zagayowar asexual.
Asalin P. INFESTANS
P. infestans sun bayyana a Turai a ƙarshen rabin farkon karni na 1991. Tun da dankalin turawa dan asalin yankin arewa maso gabashin Kudancin Amurka ne, an ɗauka cewa an kawo cutar daga nan zuwa Turai yayin haɓakar gishirin Chile. Koyaya, karatun da aka gudanar a tashar dankalin Turawa ta Rockefeller a cikin kwarin Toluca, Mexico ta tilasta sake yin wannan ra'ayi (Niederhauser, 1993, XNUMX).
1. A cikin kwarin Toluca, nau'ikan dankalin turawa masu tsire-tsire (Solanum demissum, S. bulbocastanum, da sauransu) suna da nau'ikan kwayoyin halitta daban-daban don juriya a tsaye hade da babban matakin rashin jituwa mara ma'ana, wanda ke nuna doguwar juyin halitta tare da cutar. Jinsi na Kudancin Amurka, gami da dankalin turawa, ba su da kwayoyin halitta.
2. A cikin kwarin Toluca, ana samun keɓaɓɓe tare da nau'ikan A1 da A2 masu auratayya, sakamakon haka yawan mutanen da ke shiga tsakanin P. infestans ya bazu; yayin da a cikin asalin ƙasar dankalin turawa, Kudancin Amurka, cutar ta yadu gaba ɗaya.
3. A cikin kwarin Toluca, ana samun mummunar annoba ta shekara-shekara na ƙarshen cutar ƙaiƙayi. Saboda haka, tsakanin masu binciken Arewacin Amurka (Jami'ar Cornell), ra'ayi game da Mesoamerica (Amurka ta Tsakiya) a matsayin wurin haifuwar dankalin turawa mai suna phytophthora an kafa (Goodwin et al., 1994).
Masu binciken Kudancin Amurka ba su da wannan ra'ayin. Sun yi imanin cewa dankalin turawa da ƙarancin ƙwayar cutar ta P. suna da ƙasarsu ta asali - Andes ta Kudu ta Kudu. Sun goyi bayan ra'ayinsu ta hanyar nazarin kwayoyin akan nazarin DNA polymorphisms na mitochondrial genome (mtDNA) da kwayoyin nukiliya RAS da β-tubulin (Gomez-Alpizar et al., 2007). Sun nuna cewa nau'ikan da aka tattara daga sassa daban-daban na duniya sun samo asali ne daga layukan magabata uku da suka bambanta (duka ukun) ana samun su ne a Kudancin Amurka ta Andes. Eananan haruffan Andean zuriyar layuka biyu ne: keɓaɓɓun layin mtDNA ana samun su ne a kan Solanaceae mai tsire-tsire daga ɓangaren Anarrhicomenum a Ecuador, yayin da keɓaɓɓen layin na biyu ya zama ruwan dare a kan dankali, tumatir da kuma hasken dare. A cikin Toluca, hatta nau'ikan halittu masu saurin yaduwa sun samo asali ne daga jinsi daya kawai, tare da bambancin kwayar halittar kwayar Toluca (mabuɗin mabuɗan wasu shafuka masu canzawa) yana nuna tasirin mai kafa ƙarfi saboda guguwar kwanan nan.
Bugu da kari, an sami wani sabon nau'in P. andina a cikin Andes, ta fuskar dabi'a da kuma kwayar halitta iri daya da P. infestans, wanda, a cewar marubutan, ya nuna Andes a matsayin wuri mai zafi na musamman na jinsi na Phytophthora. Aƙarshe, a cikin Turai da Amurka, P. 'Yan Infestans sun haɗa da layin Andean, yayin da a Toluca ɗaya kawai.
Wannan littafin ya haifar da martani daga ƙungiyar masu bincike daga ƙasashe daban-daban waɗanda suka yi aikin gwaji da yawa don sake nazarin binciken da ya gabata (Goss et al., 2014). A cikin wannan aikin, da farko, an yi amfani da jerin halittun DNA masu karin bayani don nazarin DNA polymorphisms; abu na biyu, don nazarin tarin, hanyoyin ƙaura, lokacin rarrabuwar jama'a, da dai sauransu. an yi amfani da samfuran ci gaba (F-statistics, kimantawar Bayesian, da sauransu) kuma, na uku, ana amfani da kwatancen ba kawai tare da nau'ikan Andean na P. andina ba, wanda a ciki ne aka sami yanayin haɗaka (P. infestans x Phytophthora sp.) amma har ila yau tare da jinsunan Mexico masu girma P. mirabilis, P. Ipomoeae, da Phytophthora phaseoli, waɗanda ke da kusanci da asalin halittar P. waɗanda aka haɗa a cikin wannan zancen (Kroon et al., 2012). A sakamakon wadannan nazarin, an nuna babu shakka cewa asalin bangaren phylogenetic bishiyar dukkanin jinsin halittar Phytophthora da aka dauke su a cikin binciken, sai dai na matattarar P. andina, na cikin matsalolin Mexico ne, kuma tafiyar kwararar tana da alkiblar Meziko - Andes, kuma ba akasin haka ba, kuma farkonta yayi daidai da Turai mulkin mallaka na Sabuwar Duniya (shekaru 300-600 da suka gabata). Don haka, fitowar jinsunan P. infestans, na musamman don kayar da dankali, ya faru ne a cikin kwayar halittar sakandare ta biyu na samuwar tumfafiyar dare, watau a Amurka ta Tsakiya.
Genome na P. INFESTANS
A cikin 2009, ƙungiyar masana kimiyya na duniya sun tsara cikakkiyar kwayar halittar P (Haas et al, 2009), wanda girman sa ya kai 240 MB. Wannan ya fi sau da yawa fiye da na cikin alaƙar da ke da alaƙa da P. sojae (95 Mb), wanda ke haifar da tushen ruwar waken soya, da P. Ramorum (65 Mb), wanda ke shafar irin nau'ikan bishiyar masu daraja kamar itacen oak, beech da sauransu. Bayanan da aka samo sun nuna cewa kwayar halittar tana dauke da adadi masu yawa na jerin abubuwa - 74%. Kwayar halitta ta ƙunshi kwayoyin 17797 masu hada-hadar sunadarai, galibinsu kwayoyi ne da ke cikin tsarin salula, gami da kwayar halittar DNA, kwafi da fassarar sunadarai.
Kwatanta kwayoyin halittar jinsin Phytophthora ya bayyana wata kungiyar baututuwa ta jinsi, wacce ta kunshi bangarori na jerin kwayoyin halittar da aka kiyaye, wanda yawan kwayar halittar yake da yawa, kuma abubuwanda aka maimaita basu da yawa, kuma yankuna daban-daban wadanda ba a kiyaye su ba, tare da karamin kwayar halitta da kuma babban abun da ke maimaitattun yankuna. Tubalan masu ra'ayin mazan jiya suna da kashi 70% (12440) na dukkanin kwayoyin halittar kwayar halitta masu dauke da sunadarai. A cikin tubalan masu ra'ayin mazan jiya, kwayoyin halitta galibi suna tazarar tazara tare da matsakaiciyar matsakaiciyar zance na 604 bp. A cikin yankuna tsakanin tubalan masu ra'ayin mazan jiya, tazarar tazarar ta fi yawa (3700 bp) saboda ƙaruwar yawaitar abubuwa masu maimaitawa. Saurin saurin tasirin kwayar halittar sirri suna cikin yankuna masu fama da talauci.
Binciken jerin halittu na P. Infestans genome ya nuna cewa kusan kashi daya bisa uku na kwayoyin yana cikin abubuwan da za'a iya yada su. Kwayar halittar halittar P. infestans ya ƙunshi iyalai daban daban na transposons fiye da sauran sanannun kwayoyin halittar. Mafi yawa daga cikin P. infestans transposons na dangin Gypsy ne.
Yawancin adadin takamaiman dangin dangin da ke cikin kwayar cutar an gano su a cikin kwayar cutar ta inf. Wani ɓangare mai mahimmanci daga cikinsu yana sanya ƙwayoyin sunadarai masu tasiri wanda ke canza tsarin ilimin lissafi na tsire-tsire da kuma ba da gudummawa ga kamuwa da cuta. Sun fada cikin fannoni biyu masu fadi: masu bada karfi, wadanda suke aiki a sararin samaniya (apoplasts), da masu tasirin cytoplasmic, wadanda suke shiga sel ta hanyar haustoria. Apoplastic Effectors sun haɗa da enzymes na hydrolytic kamar su proteases, lipases da glycosylases waɗanda ke lalata ƙwayoyin shuka; masu hana ƙwayoyin enzymes masu kariya; da kuma toxrotizing abubuwa masu guba kamar su sunadarai masu kama da Nep1 (NPLs) da kuma Pcf-kamar ƙananan sunadarai masu arzikin cysteine (SCRs).
P. infestans effector genes suna da yawa kuma galibi sun fi ƙwayoyin cuta marasa alaƙa. Mafi shahararrun sune tasirin tasirin cytoplasmic RXLR da Crinkler (CNR). Abubuwan da ke haifar da kwayar halitta ta otopismic na oomycetes sune sunadaran RXLR. Dukkanin kwayoyin tasirin tasirin RXLR da aka gano har yanzu suna dauke da amino-terminal group Arg-XLeu-Arg, inda X shine amino acid. Sakamakon binciken, an ba da shawarar cewa akwai 563 RXLR genes a cikin P. infestans genome, wanda ya ninka 60% fiye da na P. sojae da P. ramorum. Kusan rabin kwayoyin RXLR a cikin kwayar halittar halittar jini ta P. infestans genome sune takamaiman jinsi. RXLR masu tasiri suna da nau'ikan nau'ikan jerin. Daga cikinsu, an gano babba da ƙananan iyalai 150. Ba kamar babban proteome ba, yawancin sakamako masu tasiri na RXLR yawanci ana samunsu ne a cikin yankuna matalauta kuma yankuna masu tarin yawa na kwayar halittar. Abubuwan wayar hannu wadanda suke tabbatar da tasirin wadannan yankuna suna inganta hadewa a cikin wadannan kwayoyin halittar.
An gano asalin masu tasiri na Cytoplasmic CRN a cikin rubutun P. infestans wanda ke sanya kayan tsirrai necrosis peptides. Tun lokacin da suka gano, ba a san komai game da dangin waɗannan tasirin ba. Bincike game da kwayoyin halittar P. Infestans ya bayyana wata babbar iyali ta kwayoyin halittu ta 196 CRN, wanda ya fi girma girma fiye da na P. sojae (100 CRN) da P. ramorum (19 CRN). Kamar RXLRs, CRNs sunadarai ne masu daidaitaccen tsari kuma sun kunshi yanki mai lamba N-terminal LFLAK (amino acid 50) da yankin DWL da ke kusa da shi wanda ke dauke da kwayoyin halitta daban-daban. Yawancin CRNs (60%) suna da peptide na sigina.
Anyi nazarin yiwuwar CRNs da yawa don rikitar da tsarin salon salula na tsirar rundunar. A cikin binciken necrosis na tsire-tsire, cire sunadaran CRN2 ya ba da damar gano yankin C-terminal wanda ya ƙunshi 234 amino acid (matsayi 173-407, yankin DXG) da haifar da mutuwar kwayar halitta. Tattaunawa game da kwayoyin halittar P. na CRN sun bayyana yankuna C-m huɗu daban-daban, wanda kuma yana haifar da mutuwar kwayar halitta a cikin tsiron. Wadannan sun hada da sababbin wuraren da aka gano na DC (P. Infestans yana da kwayoyin 18 da 49 pseudogenes), da kuma D2 (14 da 43) da kuma DBF (2 da 1) yankuna waɗanda suke kama da furotin kinases. Sunadaran rukunin CRN da aka bayyana a cikin shuka ana kiyaye su (in babu peptides na sigina) a cikin kwayar tsire-tsire kuma tana motsa mutuwar kwayar halitta ta hanyar hanyar intracellular. Wani jerin 255 da ke dauke da yankuna CRN bazai yuwu aiki azaman kwayoyin halitta ba.
Inara yawan lamba da girman RXLR da dangin mai tasiri na CRN mai yiwuwa ne saboda sake haɗaɗɗiyar haɗin gwiwar ba da haɗin kai da kuma kwafin halitta. Duk da cewa kwayar halittar tana dauke da adadi mai yawa na abubuwan wayar hannu, har yanzu babu wata hujja kai tsaye game da canzawar kwayoyin halitta masu tasiri.
Hanyoyin da aka yi amfani da su wajen nazarin tsarin yawan jama'a
Nazarin tsarin halittar mutane a halin yanzu ya ta'allaka ne akan nazarin kyawawan al'adu na abubuwan da ke tattare da shi. Hakanan ana yin nazarin yawan jama'a ba tare da kebe albarkatun gona mai tsabta ba don takamaiman dalilai, kamar, misali, yin nazarin ta'addancin jama'a ko kasancewar nau'ikan nau'ikan da ke jure wa kayan gwari a ciki (Filippov et al., 2004; Derevyagina et al., 1999). Irin wannan binciken ya shafi amfani da hanyoyi na musamman, wanda bayaninsa ya wuce girman wannan bita. Don nazarin kwatancen damuwa, ana amfani da hanyoyi da yawa bisa duka akan tsarin DNA da kuma nazarin abubuwan da suka shafi halittu. Nazarin kwatankwacin yawan jama'a dole ne ya yi aiki tare da adadi mai yawa na ware, wanda ke sanya wasu buƙatu akan hanyoyin da aka yi amfani da su. Ainihin, ya kamata su cika waɗannan buƙatun (Cooke, Lees, 2004, Mueller, Wolfenbarger, 1999):
- zama mai arha, mai sauƙin aiwatarwa, baya buƙatar kashe kuɗaɗe mai mahimmanci, bisa ga wadatar wadatattun fasahohi (misali, PCR);
- dole ne ya samar da adadi mai yawa na alamomin alamun alama;
- sami babban haɓakawa;
- yi amfani da mafi ƙarancin ƙwayar nama don bincika;
- zama takamaimai ga mai tushe (gurbatarwar da ke cikin al'adun bai kamata ta shafi sakamakon ba);
- baya buƙatar amfani da hanyoyin haɗari da sunadarai masu haɗari sosai.
Abin takaici, babu wasu hanyoyin da suka dace da duk abubuwan da ke sama. Don nazarin kwatancin damuwa a zamaninmu, ana amfani da hanyoyi dangane da nazarin halayen ƙirar ƙira: ƙwayoyin cuta zuwa ga dankalin turawa da tumatir (tseren dankalin turawa da tumatir), nau'in mating, zane na peptidase isoenzymes da glucose-6-phosphate isomerase, kuma akan nazarin tsarin DNA: tsawon polymorphism restricuntataccen yanki (RFLP), wanda yawanci ana haɗa shi tare da binciken bincike na RG 57, nazarin maimaita microsatellite (SSR da InterSSR), haɓakawa tare da abubuwan share fage (RAPD), haɓaka guntun gutsuri (AFLP), haɓakawa tare da share fage masu kama da jerin abubuwan wayar hannu (misali, Inter SINE PCR), ƙaddarar tasirin halittun DNA.
Takaitattun bayanai game da hanyoyin don nazarin kwatancen matsalolin da aka yi amfani da su tare da P. Infestans
Alamar alama ta Phenotypic
Tseren "Dankali"
Tseren "Dankali" galibi bincike ne da amfani da alama. Tseren "Sauƙin dankalin turawa" suna da kwayar halitta guda ɗaya don ƙwayoyin cuta na dankalin turawa, waɗanda ke da “rikitarwa” - aƙalla biyu. Black et al. (1953), da ke taƙaita duk bayanan da suke a gare su, sun gano cewa tseren phytophthora na iya kamuwa da tsire-tsire tare da kwayar halittar jini / kwayar halitta daidai da P. infestans virulence gene / genes, kuma sun sami jinsi 1, 2, 3, da 4 waɗanda ke cutar shuke-shuke tare da kwayoyin R1, R2, R3 da R4, bi da bi, watau hulɗar tsakanin mahaukaci da mai gida tana faruwa ne gwargwadon kwayar halittar kwayar halitta. Bugu da ari, Baki, tare da haɗin Gallegly da Malcolmson, sun gano ƙwayoyin halittar R5, R6, R7, R8, R9, R10 da R11, da kuma jinsin da suka dace (Black, 1954; Black & Gallegly, 1957; Malcolmson & Black, 1966; Malcolmson, 1970).
Akwai wadatattun bayanai game da jinsin abin da ke haifar da cutar daga yankuna daban-daban. Ba tare da nazarin waɗannan bayanan dalla-dalla ba, za mu nuna kawai yanayin gaba ɗaya: inda aka yi amfani da nau'ikan da ke da sabbin ƙwayoyin halitta na juriya ko haɗuwarsu, da farko akwai rauni na ƙarshen cutar, amma sai ga jinsi tare da ƙwayoyin halittar da ke kama da juna sun bayyana kuma an zaɓa kuma an sake barkewar cutar ta ƙarshe. Specific virulence on the first 4 resistance genes (R1-R4) ba safai ake lura da shi ba a cikin tarin da aka tattara kafin gabatarwar cikin noman ire-iren wadannan kwayoyin halittar, amma yawan nau'in kwayar cuta ya karu sosai lokacin da aka shawo kan kwayar cutar kan irin dake dauke da wadannan kwayoyin halittar. Kwayoyin halitta 5-11, a gefe guda, sun kasance gama-gari a cikin tarin (Shaw, 1991).
Wani bincike game da rabe-raben jinsi daban-daban a lokacin noman, wanda aka gudanar a ƙarshen shekarun 1980, ya nuna cewa a farkon ɓullar cutar, kwayoyi masu ƙarancin ƙarfi da ƙwayoyin cuta na 1-2 sun fi yawa a cikin jama'a.
Bugu da ari, tare da ci gaban ƙarshen ɓarkewa, ƙwanƙolin ƙwayoyin cuta na asali yana raguwa kuma adadin tsere "mai rikitarwa" tare da tsananin tashin hankali yana ƙaruwa. Abinda ya faru a karshen ƙarshen kaka ya kai 100%. Lokacin adana tubers, akwai raguwa a cikin tashin hankali da asarar ƙwayoyin halittar mutum. Dynamarfafawa na maye gurbin clone na iya faruwa a cikin nau'ikan daban-daban ta hanyoyi daban-daban (Rybakova & Dyakov, 1990). Koyaya, karatunmu a shekara ta 2000-2010 ya nuna cewa ana samun rikice-rikice masu rikitarwa tun daga farkon epiphytotics tsakanin matsalolin da aka ware daga duka dankali da tumatir. Wannan yana yiwuwa saboda canje-canje a cikin yawan P. Infestans a Rasha.
Zuwa 1988-1995, faruwar “superraces” tare da duka ko kusan dukkanin kwayoyin cuta na kwayar cuta a yankuna daban-daban sun kai 70-100%. An lura da wannan yanayin, alal misali, a Belarus, a cikin yankunan Leningrad da Moscow, a Arewacin Ossetia da kuma a Jamus (Ivanyuk et al., 2002a, 2002b; Polityko, 1994; Schober-Butin et al., 1995).
Tseren "Tumatir"
A cikin kayan noman tumatir, an samu kwayoyin 2 na juriya da saurin tashin hankali - Ph1 (Gallegly & Marvell, 1955) da Ph2 (Al-Kherb, 1988). Kamar yadda yake a cikin yanayin tseren dankalin turawa, hulɗar tsakanin tumatir da 'yan iska na P. suna faruwa ne a kan tsarin kwayar halitta. Gasar T0 tana haifar da nau'ikan da ba su da ƙwayoyin halitta na juriya (yawancin nau'ikan da ake amfani da su a masana'antu), tseren T1 yana haifar da nau'ikan tare da kwayar Ph1 (Ottawa), kuma tseren T2 yana haifar da nau'ikan tare da kwayar Ph2.
A cikin Rasha, kusan T0 kawai aka samo akan dankali; T0 ya mamaye tumatir a farkon kaka, amma daga baya aka maye gurbinsa da jinsin T1 (Dyakov et al., 1975, 1994). Bayan shekara ta 2000, T1 akan dankali a yawancin jama'a ya fara faruwa a farkon farkon epiphytotics. A Amurka, matsalolin dankalin turawa ba su da cuta ga tumatir, kazalika da jinsi T0, T1, da T2, yayin da T1 da T2 suka mamaye tumatir (Vartanian & Endo, 1985; Goodwin et al., 1995).
Nau'in ma'aurata
Don gudanar da binciken, ana buƙatar nau'ikan gwaji (tunani) tare da sanannun nau'ikan saduwa - A1 da A2 - ana buƙata. An yi wa gwajin gwajin tare da su nau'i-nau'i a cikin jita-jita na Petri tare da oat agar matsakaici. Bayan shiryawa na tsawon kwanaki 10, ana bincika faranti don kasancewar ko babu warkarwa a cikin matsakaici a yankin tuntuɓar damuwa. Akwai hanyoyi guda 4: damuwar tana da nau'ikan A1 na mating, idan ta samar da maganadisu tare da mai gwajin A2, zuwa A2, idan ta samar da maganadisuwa tare da mai gwajin A1, zuwa A1A2, idan ta samar da iska tare da masu gwajin guda biyu, ko kuma bakararre ne (00), idan ba ta samar da komai ba ba tare da mai gwaji ba (ƙungiyoyi biyu na ƙarshe ba su da yawa).
Don ƙarin saurin ƙayyade nau'ikan saduwa, an yi ƙoƙari don gano yankuna na ƙwayoyin halittar da ke haɗuwa da nau'in saduwa, da nufin ƙarin amfani da su don tantance nau'in ma'aurata ta PCR. Researchersaya daga cikin gwaje-gwajen nasara na farko don gano irin wannan rukunin yanar gizon shine masu binciken Amurka (Judelson et al., 1995) suka gudanar. Ta amfani da hanyar RAPD, sun sami damar gano yankin W16 da ke hade da nau'in mating a cikin zuriyar bangarorin biyu da suka tsallake, da kuma tsara zane-zane na 24-bp don haɓaka shi (W16-1 (5'-AACACGCACAAGGCATATAAATGTA-3 ') da W16-2 (5' -GCGTAATGTAGCGTAACAGCTCTC-3 ') Bayan ƙuntata samfurin PCR tare da enzyme mai ƙuntatawa HaeIII, yana yiwuwa a raba keɓaɓɓu tare da nau'ikan haɗin A1 da A2.
Wani yunƙurin samun alamomin PCR don ƙayyade nau'ikan saduwa da masu binciken Koriya suka yi (Kim, Lee, 2002). Sun gano takamaiman samfuran amfani da hanyar AFLP. A sakamakon haka, an haɓaka nau'ikan share fage PHYB-1 (gaba) (5'-GATCGGTTAGTCAGACGAG-3 ') da PHYB-2 (5'-GCGTCTGCAAGGCCCCTTTT-3'), wanda ya ba da damar haɓakar zaɓin yanayi wanda ke da alaƙa da nau'in A2. Bayan haka, sun ci gaba da wannan aikin kuma sun tsara zane-zane na 5 'AAGCTATACTGGGACAGGGT-3' (INF-1, gaba) da 5'-GCGTTCTTTCGTATTACCAC-3 '(INF-2), yana ba da damar haɓaka zaɓin yanayin Mat-A1 na yanayin damuwa da nau'in mating A1. Yin amfani da bincike na PCR na nau'ikan ma'aurata ya nuna kyakkyawan sakamako a cikin nazarin yawan mutanen da ke cin abinci a Jamhuriyar Czech (Mazakova et al., 2006), Tunisia (Jmour, Hamada, 2006), da sauran yankuna. A cikin dakin gwaje-gwajenmu (Mytsa, Elansky, ba a buga shi ba), an yi nazari kan nau'ikan 34 P. wadanda aka kebe daga cututtukan dankalin Turawa da gabobin tumatir a yankuna daban-daban na Rasha (Kostroma, Ryazan, Astrakhan, Moscow oblasts). Sakamakon bincike na PCR ta amfani da takamaiman share fage sama da 90% ya yi daidai da sakamakon bincike na nau’in saduwa ta hanyar gargajiya bisa tsarin abinci mai gina jiki.
Tebur 1. Bambancin juriya tsakanin Sib 1 clone (Elansky et al., 2001)
Samfurin wurin tattarawa | Adadin keɓance da aka bincika | Yawan m (S), mai rauni mai ƙarfi (SR) da ƙwayoyin cuta masu ƙarfi (R), inji mai kwakwalwa (%) | ||
S | SR | R | ||
G. Vladivostok | 10 | 1 (10) | 4 (40) | 5 (50) |
G. Chita | 5 | 0 | 0 | 5 (100) |
Irkutsk | 9 | 9 (100) | 0 | 0 |
G. Krasnoyarsk | 13 | 12 (92) | 1 (8) | 0 |
Yekaterinburg birni | 15 | 8 (53) | 1 (7) | 6 (40) |
O. Sakhalin | 66 | 0 | 0 | 66 (100) |
Yankin Omsk | 18 | 0 | 0 | 18 (100) |
Tsarin Metalaxyl azaman alamar yawan jama'a
A farkon 1980s, an sami barkewar annoba mai karfi ta sanadiyyar cutar sanadiyyar cutar sanadin jiki na metalaxyl-P. Gonakin dankalin turawa a kasashe da dama sun yi asara mai yawa (Dowley & O'Sullivan, 1981; Davidse et al., 1983; Derevyagina, 1991). Tun daga wannan lokacin, a cikin ƙasashe da yawa na duniya, ana ci gaba da lura da abin da ya faru na ƙwayoyin phenylamide masu jurewa a cikin al'ummomin P. infestans. Baya ga kimantawa mai amfani da ake da ita game da amfani da kwayoyi masu dauke da phenylamide, gina tsarin matakan kariya da tsinkayen epiphytoties, juriya ga wadannan kwayoyi ya zama daya daga cikin alamun alamun da ake amfani dasu sosai don kwatankwacin yawan mutanen wannan cuta. Koyaya, yin amfani da juriya ga metalaxyl a cikin nazarin yawan jama'a yakamata ayi la'akari da gaskiyar cewa: 1 - asalin kwayar halitta ta juriya ba a riga an ƙaddara ta daidai ba, 2 - juriya ga metalaxyl shine halin zaɓaɓɓe wanda zai iya canzawa dangane da amfani da phenylamides, 3 - daban mataki na ƙwarewa ga ɓarna na metalaxyl tsakanin layi ɗaya na layi (tebur. 1).
Spectra na isozymes
Alamomin Isozyme galibi basa cin gashin kansu ne daga yanayin waje, suna nuna gadon Mendelian kuma suna da yawa, suna ba da damar rarrabewa tsakanin homo da heterozygotes. Amfani da sunadarai azaman alamomin jinsi yana ba da damar gano manyan sifofin sake tsarin halittar gado, gami da chromosomal da maye gurbi, da maye gurbin amino acid daya.
Karatun Electrophoretic na sunadarai ya nuna cewa yawancin enzymes sun wanzu a cikin kwayoyin halitta a cikin nau'ikan bangarori da suka banbanta a cikin motsi na electrophoretic. Wadannan bangarorin sakamakon sakamako ne na sanya nau'ikan enzymes masu yawa ta hanyar daban-daban (isozymes ko isozymes) ko kuma ta hanyar daban-daban na wuri guda (allozymes ko alloenzymes). Wato, isozymes nau'ikan daban daban ne na enzyme daya. Hanyoyi daban-daban suna da aiki iri ɗaya, amma sun ɗan bambanta a cikin maye gurbin amino acid ɗaya a cikin peptide da caji. Irin waɗannan bambance-bambance ake bayyana yayin electrophoresis.
Yayin da ake nazarin kwayar cutar ta P. infestans, ana amfani da yanayin isoenzymes na sunadarai guda biyu, peptidase da glucose-6-phosphate isomerase (wannan enzyme din din din din din din din din din din din din din din din din din din din din din ne, saboda haka, ba a gabatar da hanyoyin bincikensa a wannan aikin ba) Don raba su cikin isozymes a cikin filin lantarki, shirye-shiryen furotin da aka keɓe daga ƙwayoyin nazarin ana amfani da su a kan farantin gel da aka sanya a cikin filin lantarki. Yawan yaduwar kwayoyin sunadarai a cikin gel ya dogara da caji da nauyin kwayoyin; sabili da haka, a cikin filin lantarki, cakudadden sunadarai sun kasu kashi-kashi, wanda za'a iya gani ta amfani da dyes na musamman.
Nazarin peptidase isoenzymes ana aiwatar dashi akan cellulose acetate, sitaci ko mala'ikan polyacrylamide. Mafi dacewa shine hanyar da aka dogara da amfani da gels acetate gel wanda Helena Laboratories Inc. Baya buƙatar adadi mai yawa na kayan gwaji, yana ba mutum damar samun ƙungiyoyi masu banbanci akan gel bayan electrophoresis don duka enzyme loci, aiwatarwar sa baya buƙatar babban lokaci da farashin kayan aiki (Fig. 2).
An canza karamin yanki na mycelium a cikin microtube milimita 1,5, an kara digo 1-2 na ruɓaɓɓen ruwa a ciki. Bayan haka, samfurin ya zama mai kama da juna (misali, tare da rawar lantarki tare da abin da aka makala na filastik wanda ya dace da microtube) kuma an share shi tsawon dakika 25 a kan centrifuge a 13000 rpm. 8 froml daga kowane microtube. an canza supernatant ɗin zuwa faranti mai nema.
An cire gel acetate acetate daga kwandon ajiyar, an goge tsakanin zannuwan takarda biyu da kuma sanya su tare da aikin aiki a saman filastik na mai nema. Maganin daga farantin ana tura shi ta mai amfani zuwa gel sau 2-4. An canza gel zuwa ɗakin electrophoresis,
Tebur na 2. Maganin maganin da aka yi amfani da shi don shafa gel acetate gel a cikin binciken peptidase isoenzymes, an sanya digo na fenti (bromophenol blue) a gefen gel.
TRIS HCl, 0,05M, Ph 8,0 2 ml
Peroxidase, 1000 U / ml 5 saukad da
o-dianisidine, 4 mg / ml 8 saukad da
MgCl2, 20 mg / ml 2 saukad da
Gly-Leu, 15 mg / ml 10 saukad da
L-amino-acid oxidase, 20 u / ml 2 saukad da
Ana amfani da wutar lantarki na mintina 20. a 200 V. Bayan electrophoresis, ana canja gel ɗin zuwa teburin zane kuma a goge shi da maganin zane na musamman (Tebur 2). 10 mil na 1,6% DIFCO agar an narke shi na farko a cikin tanda na microwave, sanyaya zuwa 60 ° C, bayan haka an gauraya agar 2 ml tare da cakuda fenti kuma an zuba shi a kan gel. Rauni yana bayyana tsakanin mintuna 15-20. An kara reagent na L-amino-acid oxidase nan da nan kafin a hada maganin da narkakken agar.
A cikin jama'ar Rasha, ,ungiyar Pep 1 tana wakiltar nau'ikan jinsin 100/100 da 92/100. Homozygote 92/92 ba safai ake samun sa ba (kusan 0,1%). Locus Rehr 2 yana da wakiltar nau'ikan halittu guda uku 100/100, 100/112, da 112/112, kuma dukkan nau'ikan 3 suna gama gari (Elanky da Smirnov, 2003, Hoto 2).
Bincike na Genome
Muntataccen ɓangaren polymorphism tare da haɗuwa mai zuwa (RFLP-RG 57)
Ana amfani da jimlar DNA tare da Eco R1 enzyme na ƙuntatawa, an raba sassan DNA ta hanyar electrophoresis a cikin gel na agarose. DNA na Nukiliya yana da girma sosai kuma yana da jerin maimaitawa da yawa, wanda hakan ke da wuya kai tsaye a bincika yawancin gutsutsuren da aka samu ta hanyar aikin enzymes na ƙuntatawa. Sabili da haka, ana jujjuya sassan DNA da aka rabu a cikin gel zuwa membrane na musamman kuma ana amfani dasu don haɗuwa tare da binciken RG 57, wanda ya haɗa da nucleotides da aka lakafta tare da tasirin rediyo ko alamun haske. Wannan binciken yana haɗuwa tare da maimaita jigilar kwayoyin halitta (Goodwin et al., 1992, Forbes et al., 1998). Bayan gani na sakamakon hadewa akan abu mai haske- ko radiyo, an sami bayanin hade-wuri mai yawa (zanan yatsan hannu), wanda gungun 25-29 suka wakilta (Forbes et al., 1998). 'Ya'yan Asexual (clonal) zasu sami bayanan martaba iri ɗaya. Ta hanyar tsara makada a kan electrophoretogram, mutum na iya yin hukunci da kamanceceniya da bambance-bambancen kwayoyin halittu idan aka kwatanta su.
Tsarin halittar DNA na mitochondrial
A mafi yawan kwayoyin eukaryotic, ana gabatar da mtDNA a cikin sifa ta madaidaiciyar madaidaiciyar kwayar halittar DNA, wanda, ba kamar chromosomes na nukiliya na kwayoyin eukaryotic ba, suna maimaita kwaro daya kuma ba a hade da kwayoyin sunadaran ba.
An tsara jigon halittar mitochondrial na P. infestans, kuma yawancin ayyukanda aka keɓe ga nazarin ƙuntataccen gutsuri (Carter et al, 1990, Goodwin, 1991, Gavino, Fry, 2002). Bayan Griffith da Shaw (1998) sun kirkiro hanya mai sauƙi da sauri don tantance mtDNA haplotypes, wannan alamar ta zama ɗayan shahararru a cikin karatun P. Infestans.Mahimmancin hanyar yana ƙunshe da haɓakawa bi-bi-bizu na wasu gutsuttukan DNA na mitochondrial guda biyu (daga asalin kwayar halitta) tare da fararen F2-R2 da F4-R4 (Table 3) da ƙuntatawarsu ta gaba tare da ƙuntatawa enzymes MspI (yanki 1) da EcoR1 (yanki na 2). Hanyar tana baka damar gano nau'ikan fasalin guda 4: Ia, IIa, Ib, IIb. Nau'in na II ya bambanta da nau'ikan I ta wurin kasancewar saka 1881 bp a cikin girma da kuma ta wani wurin daban na shafukan ƙuntatawa a yankuna P2 da P4 (Fig. 3).
Tun daga 1996, daga cikin matsalolin da aka tattara a kan yankin na Rasha, kawai alamun Ia da IIa ne aka lura (Elansky et al., 2001, 2015). Ana iya gano su bayan rabuwar samfuran ƙuntatawa tare da share fage na F2-R2 a cikin filin lantarki (Fig. 4, 5). Nau'ikan mtDNA ana amfani dasu a cikin kwatancen kwatanta damuwa da yawan jama'a. A cikin yawan karatun, an yi amfani da nau'ikan DNA na mitochondrial don keɓe layinn dunƙulen da keɓaɓɓu na ɓarnata P. Ta amfani da hanyar PCR-RFLP, an yanke shawarar cewa mtDNA yana da bambanci a cikin wannan nau'in P. infestans (Elansky da Milyutina, 2007). Yanayin haɓakawa: 2009x (2007 sec. 1 ° C), 500x (94 sec. 40 ° C, 30 sec. 90 ° C, 30 sec. 52 ° C); 90x (72 min. 1 ° C). Cakuda mai amsawa: (5 μl): 72 U Taq DNA polymerase, 20x 0,2 mM MgCl1-Taq buffer, 2,5 mM kowane dNTP, 2 pM share fage da 0,2 ng na DNA da aka binciko, ruwan da aka daskare - har zuwa 30 μl.
Outuntataccen samfurin PCR ana aiwatar dashi na awanni 4-6 a zazzabi na 37 ° C. Mixtureuntataccen cakuda (20 )l): 10x MspI (2 μl), maɓallin hana ƙuntatawa 10x (2 μl), ruwan da aka ƙaddara (6 μl), samfurin PCR (10 μl).
Tebur 3. An yi amfani da firamare don fadada yankunan mtDNA polymorphic
Cusungiyoyi | Farkon | Tsawon farko da sanyawa | PCR samfurin tsawon | Restuntata |
---|---|---|---|---|
P2 | F2: 5'- TTCCCTTTTGTCCTCTAGCGAT | 21; 13619-13639 | 1070 | MspI |
R2: 5'- TTACGGCGGTTTAGCACATACA | 22; 14688-14667 | |||
P4 | F4: 5'- TGGTCATCCAGAGGTTTATGTT | 22; 9329-9350 | 964 | EcoRI |
R4: 5 - CCGATACCGATACCAGCACAC | 22; 10292-10271 |
Omara ƙarfin share fage (RAPD)
Lokacin aiwatar da RAPD, ana amfani da share daya (wani lokaci ma firamare da yawa a lokaci ɗaya) tare da jerin nucleotide mara sabani, yawanci nucleotides 10 a tsayi, tare da babban abun ciki (daga 50%) na GC nucleotides da ƙarancin zafin jiki na annealing (kimanin 35 ° C). Irin waɗannan abubuwan share fage “kan ƙasa” akan yawancin shafukan yanar gizo masu haɗaka a cikin kwayar halitta. Bayan fadadawa, ana samun adadi mai yawa na amplicons. Lambar su ya dogara da abubuwan share fage (s) da aka yi amfani da su da kuma yanayin tasirin su (MgCl2 mai ɗorewa da zafin jiki na haɗuwa)
Gani na amplicons ana aiwatar dashi ta hanyar narkewa a cikin polyacrylamide ko gel na agarose. Yayin gudanar da bincike na RAPD, ya zama dole a kula da tsabtar kayan da aka bincika, saboda gurɓatawa tare da wasu abubuwa masu rai na iya haifar da ƙaruwa mai yawa a cikin kayan tarihi, waɗanda suke da yawa a cikin binciken abubuwa masu tsabta (Perez et al, 1998). Amfani da wannan hanyar a cikin binciken kwayar cutar ta infestans genome ya bayyana a cikin ayyuka da yawa (Judelson, Roberts, 1999, Ghimire et al., 2002, Carlisle et al., 2001). Zaɓin yanayin yanayin aiki da share fage (an yi nazarin 51 10-nucleotide primers) a cikin labarin Abu-El Samen et al., (2003).
Binciken Maimaita Microsatellite (SSR)
Maimaita tauraron dan adam (maimaitattun jerin maimaitawa, SSR) ana maimaita gajerun jerin 1-3 (wani lokacin har zuwa 6) nucleotides da ke cikin kwayoyin halittar nukiliya na dukkan eukaryotes. Adadin maimaitawa na baya-baya zai iya bambanta daga 10 zuwa 100. Microsatellite loci yana faruwa tare da madaidaiciyar mita kuma ana rarraba ko dai-dai a ko'ina cikin jigilar kwayoyin halitta (Lagercrantz et al., 1993). Polymorphism na jerin microsatellite jerin suna haɗuwa da bambance-bambance a cikin adadin maimaitawa na ainihin motif. Alamar tauraron dan adam suna da yawan gaske, wanda ke ba da damar amfani da su don nazarin tsarin yawan jama'a, ƙayyade dangi, hanyoyin ƙaura na jinsi, da dai sauransu. Daga cikin sauran fa'idodi na waɗannan alamomin, yakamata mutum ya lura da haɓakar polymorphism ɗinsa, mai iya haifuwa mai kyau, tsaka tsaki, da kuma ikon aiwatar da bincike na atomatik da kimantawa. Da farko, an gudanar da bincike ne tare da rabuwa da kayan aikin dauki akan gel polyacrylamide. Daga baya, ma'aikatan kamfanin Applied Biosystems sun ba da shawarar yin amfani da share fage masu haske tare da gano samfuran samfuran ta hanyar amfani da injin gano laser ta atomatik (Diehl et al., 1990), sannan kuma daidaitattun masu tsara DNA ta atomatik (Ziegle et al., 1992). Yin lakabi na share fage tare da launuka iri-iri masu haske suna ba da damar nazarin alamomi da yawa a lokaci ɗaya a kan layi ɗaya kuma, bisa ga haka, yana ƙaruwa ƙimar aikin hanyar da haɓaka ƙimar binciken.
Littattafan farko da aka keɓe don amfani da nazarin SSR don nazarin P. infestans sun bayyana a farkon 2000s. (Knapova, Gisi, 2002). Ba dukkan alamomin da marubutan suka gabatar ba sun nuna cikakken digiri na polymorphism, duk da haka, biyu daga cikinsu (4B da G11) an haɗa su a cikin saitin alamomi 12 SSR waɗanda Lees et al suka gabatar. (2006) kuma daga baya cibiyar sadarwa ta Eucablight ta karɓi su (www.eucablight) .org) a matsayin mizanin ma'auni na P. infestans. Bayan fewan shekaru daga baya, an wallafa wani bincike game da ƙirƙirar tsarin bincike na multiplex na P. infestans DNA dangane da alamomi takwas na SSR (Li et al., 2010). A ƙarshe, bayan kimanta duk alamomin da aka gabatar a baya kuma zaɓi waɗanda suka fi ba da bayani game da su, tare da inganta abubuwan share fage, alamun haske, da yanayin haɓakawa, rukuni ɗaya na marubutan sun gabatar da tsarin nazarin matakan sau ɗaya, gami da alamomi 12 (Table 4; Li et al. , 2013a). An zaɓi abubuwan share fage da aka yi amfani da su a cikin wannan tsarin kuma an lakafta su da ɗayan alamomi masu kyalli guda huɗu (FAM, VIC, NED, PET) don haka jeren jumloli masu girma na firamare masu alamu iri ɗaya ba su zo ɗaya ba.
Marubutan sun gudanar da binciken ne a kan na'urar kara karfin PTC200 (MJ Research, USA) ta amfani da kayan PCR na QIAGEN multiplex ko kuma QIAGEN Typeit Microsatellite PCR. Girman cakuda mai amsawa ya kasance 12.5 μL. Yanayin haɓakawa ya kasance kamar haka: don QIAGEN multiplex PCR: 95 ° C (15 min), 30x (95 ° C (20 sec), 58 ° C (90 sec), 72 ° C (60 sec), 72 ° C (20 min); don QIAGEN Type-it Microsatellite PCR: 95 ° C (5 min), 28x (95 ° C (30 sec), 58 ° C (90 sec), 72 ° C (20 sec), 60 ° C (30 min).
Rabuwa da gani na kayan PCR an gudanar dasu ta amfani da ABI3730 mai binciken kwayar halitta ta atomatik (Aiwatar da Biosystems).
Tebur 4. Halaye na alamun 12 na yau da kullun na SSR da aka yi amfani dasu don jigilar P. Infestans (Li et al., 2013a)
Title | Adadin alleles | Yanayin girma adalan (bp) | Firamare |
PiG11 | 13 | 130-180 | F: NED-TGCATATATTATTATCAGCGTGGG R: GTTTCAATCTGCAGAGCAGAGAGAGA |
Ft02 | 4 | 255-275 | F: NED-ACGGAGAACTACCGCCC R: GTTGACCACTTTTAGCAGCGGCTC |
Saukewa: FSSR11 | 4 | 325-360 | F: NED-TATAGGACCGGATATGAGCTG R: GTTAGAGAATTGTTTTAGTAGAGCAGCAGC |
D13 | 16 | 100-185 | F: FAM-TGCCCCCTGCTCACTC R: GCTCGAATATTTTGAGCAGAGTTG |
Saukewa: FSSR8 | 4 | 250-275 | F: FAM-AATGGATATATATCGCAACTGAGGG R: GTTTACAAGATACACACGAGGCAGCAGA |
Saukewa: FSSR4 | 7 | 280-305 | F: FAM-TATTGTTGAGTATGCAGAGA R |
Ft04 | 4 | 160-175 | F: VIC-AGCGGCTGCCGAGCGAGG R |
Ft70 | 3 | 185-205 | F: VIC-ATGAAAATACGTCAATGCTCG R: CGTTGGATATTATTATTATTATTATTATTAG |
Saukewa: FSSR6 | 3 | 230-250 | F: GTTTGGGGGGGCGGAGAGAGTAGT R |
Ft63 | 3 | 265-280 | F: VIC-ATGACGAGGATGAGAGAGAGAGG R |
Saukewa: FSSR2 | 3 | 165-180 | F: PET-CGACCATATATATATCAACCGGC R: GTTTGCTTGGAGCAGCAGCAGCAGCAGTTAGC |
Pi4B | 5 | 200-295 | F: PET-AAAATAAAGCCTTTGGTTATA R: GAGAAGCAGAGTTTAGAGAGTT |
Misali na ganin sakamakon bincike yana cikin Fig. 6. An yi nazarin sakamakon ta amfani da software ta GeneMapper 3.7 ta hanyar kwatanta bayanan da aka samu tare da bayanan sanannun saniyar ware. Don sauƙaƙe fassarar sakamakon binciken, ya zama dole a haɗa da keɓaɓɓun ra'ayoyin 1-2 tare da sanannen genotype a kowane binciken.
An gwada hanyar binciken da aka gabatar akan wasu adadi mai yawa na samfuran, bayan haka ne marubutan suka gudanar da daidaitattun ka'idoji tsakanin dakunan gwaje-gwaje na kungiyoyi biyu, The James Hutton Institute (UK) da Wageningen University & Research (Netherlands), wanda, tare da yiwuwar amfani da katunan FTA na yau da kullun don sauƙaƙa tattarawa da jigilar P. ya mamaye samfuran DNA, ya ba da damar magana game da yiwuwar cinikin wannan ci gaban. Bugu da kari, wata hanzari kuma madaidaiciyar hanya ta genotyping P. infestans kebewa ta amfani da bincike da yawa na SSR ya ba da damar gudanar da daidaitaccen nazarin yawan wannan kwayar cutar a kan sikelin duniya, da kuma kirkirar wani matattarar bayanai ta duniya a kan makararriyar cuta a cikin tsarin aikin na Eucablight (www.eucablight.org), gami da , ciki har da sakamakon nazarin microsatellite, ya ba da damar bin diddigin bayyanar da yaduwar sabbin kwayoyin halittu a duniya.
Mara girman ƙuntataccen guntun yanki polymorphism (AFLP). AFLP (haɓakar haɓakar tsinkayen polymorphism) fasaha ce don samar da alamomin kwayoyin bazuwar ta amfani da takamaiman share fage. A cikin AFLP, ana bi da DNA tare da haɗin enzymes na ƙuntatawa biyu. Adaayyadaddun masu haɗawa suna haɗe zuwa ƙarshen maƙogwaron gutsurin abubuwan hanawa.
Waɗannan gutsuttsura ana haɓaka su ta amfani da abubuwan share fage wanda ya dace da jerin adaftan da kuma shafin ƙuntatawa, kuma bugu da carryingari suna ɗauke da ɗaya ko fiye da bazuwar tushe a ƙarshen '3' ɗinsu. Saitin gutsutsuren da aka samo ya dogara da ƙayyadadden enzymes da zaɓaɓɓun nucleotides bazuwar a 3'-ƙarewar farkon (Vos et al., 1995). AFLP - ana amfani da genotyping don saurin nazarin bambancin kwayoyin halittu daban-daban.
An ba da cikakken bayanin hanyar a cikin ayyukan Mueller, Wolfenbarger, 1999, Savelkoul et al., 1999. Mafi yawan aikin kwatanta ƙudurin hanyoyin AFLP da SSR an gudanar da su ne ta hanyar masu binciken Sinawa. Anyi nazarin halaye masu kama da juna na 48 P. ƙananan dabbobi waɗanda aka tattara a yankuna biyar na Arewacin China. Wasannin AFLP sun nuna nau'ikan jinsin DNA guda takwas daban, sabanin halittar ta SSR, wacce ba a samu wani bambancin ba (Guo et al., 2008).
Plara haɓaka tare da abubuwan share fage masu kama da jerin abubuwan wayar hannu
Alamomin da aka samo daga jerin abubuwan da aka sake rubutawa sun dace sosai da zana taswirar kwayoyin halitta, nazarin bambancin kwayoyin halittu da tsarin juyin halitta (Schulman, 2006). Idan ana yin abubuwan share fage don taimakawa daidaitattun jerin wasu abubuwa na wayoyi, akwai yiwuwar fadada yankunan kwayar halittar dake tsakaninsu. A cikin nazarin wakili mai kawo ƙarshen tashin hankali, hanyar fadada yankuna masu amfani da kwayar halitta ta hanyar amfani da abin da ya dace da ainihin tsarin SINE (Short Interspersed Nuclear Elements) an yi amfani da retroazone cikin nasara (Lavrova da Elansky, 2003). Ta yin amfani da wannan hanyar, an bayyana bambance-bambance har ma a cikin zuriya ɗaya wanda ya keɓe ɗaya. Dangane da wannan, an yanke shawarar cewa hanyar-SINE - PCR tana da takamaiman bayani kuma yawan motsi na abubuwan SINE a cikin kwayar halittar Phytophthora yana da yawa.
A cikin kwayar halittar P. infestans, an gano iyalai 12 na gajerun retrotransposons (SINEs); an bincika rabe-raben jinsunan gajerun retrotransposons, an gano abubuwan (SINEs) wadanda ake samu a cikin kwayar halittar P. infestans (Lavrova, 2004).
Siffofin aikace-aikacen hanyoyin nazarin kwatancen wahala a cikin yawan karatu
Yayin tsara karatu, dole ne mutum ya fahimci manufofin da yake bi da kyau kuma yayi amfani da hanyoyin da suka dace. Don haka, wasu hanyoyin suna ba da damar samar da adadi mai yawa na alamun alamomi masu zaman kansu, amma a lokaci guda suna da ƙaramar haifuwa kuma suna dogara sosai da abubuwan da aka yi amfani da su, yanayin halayen, da kuma gurɓataccen kayan gwajin. Sabili da haka, a cikin kowane nazarin ƙungiyar damuwa, ya zama dole a yi amfani da daidaitattun daidaitattun abubuwa (tunani), amma har ma a wannan yanayin, sakamakon gwaje-gwajen da yawa yana da matukar wahalar haɗuwa.
Wannan rukunin hanyoyin sun hada da RAPD, AFLP, InterSSR, InterSINE PCR. Bayan fadadawa, ana samun adadi mai yawa na gutsuttsukan DNA na girma daban-daban. Yana da kyau a yi amfani da irin wadannan dabarun idan ya zama dole a samar da bambance-bambance tsakanin matsalolin da ke da nasaba da juna (mahaifa-zuriya, ire-iren halittun daji, da sauransu), ko kuma a lokuta da ake bukatar cikakken bincike kan karamin samfuri. Don haka, ana amfani da hanyar AFLP a cikin taswirar kwayoyin halittu na P. infestans (van der Lee et al., 1997) kuma a cikin nazarin yawan jama'a (Knapova, Gisi, 2002, Cooke et al, 2003, Flier et al, 2003). Irin waɗannan hanyoyin basu dace ayi amfani dasu ba yayin ƙirƙirar rumbunan adana bayanai na damuwa, saboda ba shi yiwuwa a iya haɗa lissafin sakamako yayin gudanar da bincike a cikin ɗakunan gwaje-gwaje daban-daban.
Duk da alamun sauki da saurin aiwatarwa (keɓewar DNA ba tare da tsarkakewa mai kyau ba, haɓakawa, kallon sakamakon), wannan rukunin hanyoyin yana buƙatar yin amfani da wata hanya ta musamman don tattara sakamakon: ɓarna a cikin polyacrylamide gel tare da tambarin (rediyoaktif ko luminescent) da haske mai zuwa na gaba ko hasken abu mai aiki da iska. Halin al'ada ethidium bromide agarose gel hoto bai dace da waɗannan hanyoyin ba saboda adadi mai yawa na sassan DNA na girma dabam na iya haɗuwa.
Sauran hanyoyin, akasin haka, suna ba ku damar ƙirƙirar ƙananan adadi na fasali tare da haɓaka mai girma sosai. Wannan rukunin ya hada da nazarin halittun halittar dan adam (haplotypes biyu na Ia da IIa ne kawai aka lura dasu a Rasha), nau'ikan saduwa (an raba yawancin kadarori zuwa nau'ikan 2: A1 da A2, ba a samun SF mai saurin haihuwa) da peptidase isozyme spectra (biyu loci Pep1 da Pep2 , wanda ya kunshi isozymes guda biyu kowannensu) da glucose-6-phosphate isomerase (a Rasha babu wani bambanci a cikin wannan halin, kodayake an lura da polymorphism mai mahimmanci a wasu ƙasashe na duniya). Yana da kyau a yi amfani da waɗannan sifofin yayin nazarin tarin abubuwa, tattara bayanan yanki da na duniya. Game da nazarin isozymes da haplotypes na mitochondrial DNA, yana yiwuwa a yi ba tare da daidaitattun matsaloli ba kwata-kwata, yayin da ake nazarin nau'ukan saduwa, ana buƙatar keɓaɓɓun gwaji guda biyu tare da sanannun nau'ikan saduwa.
Yanayin amsawa da reagents na iya shafar bambancin samfurin a kan electrophoretogram kawai; bayyanuwar kayan tarihi a cikin waɗannan nau'o'in karatun da wuya.
A halin yanzu, yawancin mutane a yankin Turai na Rasha suna wakiltar nau'ikan nau'ikan nau'ikan ma'aurata (Table 6), daga cikinsu akwai masu keɓewa da nau'ikan Ia da IIa na mitochondrial DNA (wasu nau'ikan mtDNA da aka samo a duniya ba a samo su a Rasha ba bayan 1993). Abubuwan da ke tattare da peptidase isozymes suna wakiltar nau'ikan nau'ikan halittu guda biyu a yankin Pep1 (100/100, 92/92 da heterozygote 92/100, kuma genotype na 92/92 ba kasafai ake samunsu ba (<0,3%)) kuma ta hanyar genotypes guda biyu a yankin Pep 2 (100/100) , 112/112 da heterozygote 100/112, tare da genotype 112/112 yana faruwa sau da yawa ƙasa da 100/100, amma kuma sau da yawa).
Babu wani bambanci a cikin nau'ikan isozymes na glucose-6-phosphate isomerase bayan 1993 (bacewar layin duniyan nan US-1); duk waɗanda aka keɓe masu keɓaɓɓu suna da nau'in nau'in 100/100 (Elansky da Smirnov, 2002).
Groupungiyar rukuni na uku na hanyoyi suna ba da izinin samun isassun rukuni na alamun alamomi masu zaman kansu tare da haɓakar haɓaka. A yau, wannan rukunin ya haɗa da bincike na RFLP-RG57, wanda ke samar da gutsuttsarin DNA 25-29 na girma dabam. Ana iya amfani da RFLP-RG57 duka yayin nazarin samfuran da tattara bayanai. Koyaya, wannan hanyar tayi tsada fiye da waɗanda suka gabata, yana cin lokaci, kuma yana buƙatar cikakken adadin DNA mai tsarkakakke. Sabili da haka, an tilasta mai bincike don iyakance girman abin da aka gwada.
Ci gaban RFLP-RG57 a farkon shekarun 90 na karnin da ya gabata ya ƙarfafa nazarin yawan jama'a game da mai haifar da cutar sankara. Ya zama tushen hanyar gwargwadon zaɓi da nazarin "Lines na Layi" (duba ƙasa). Tare da RFLP-RG57, nau'in mating, zanan yatsan DNA (hanyar RFLP-RG57), zane na peptidase da glucose-6-phosphate isomerase isoenzymes, da nau'in DNA na mitochondrial ana amfani dasu don gano layin dunƙulen. Godiya gareshi, an nuna al., 1994), maye gurbin tsoffin alƙaluma tare da sababbi (Drenth et al, 1993, Sujkowski et al, 1994, Goodwin et al, 1995a), sun bayyana layin duniyan da ke yaduwa a ƙasashe da yawa na duniya. Nazarin da aka yi game da ɓacin rai na Rasha ta amfani da wannan hanyar ya nuna babban kwayar halittar polymorphism na nau'ikan ɓangaren Turai da monomorphism na yawan ƙasashen Asiya da Gabas ta Tsakiya na Rasha (Elansky et al, 2001). Kuma yanzu wannan hanyar ta kasance babba a cikin karatun jama'a na P. infestans. Koyaya, yaɗuwarsa yana hanawa ta hanyar tsada mai yawa da ƙarfin aiki a aiwatarwa.
Wata dabarar da za a yi amfani da ita wacce ba kasafai a ke amfani da ita ba a karatun P. infestans shine sake nazarin microsatelite (SSR). A halin yanzu, ana amfani da wannan hanyar don rarrabe layukan layi. Don nazarin damuwa, irin waɗannan alamun alamun alamun kamar kasancewar ƙwayoyin cuta na ƙwayoyin cuta zuwa nau'in dankalin turawa (Avdey, 1995, Ivanyuk et al., 2002, Ulanova et al., 2003) da tumatir an yi amfani dasu ko'ina (kuma ana ci gaba da amfani da su). Zuwa yanzu, kwayoyin cuta na kwayar cuta zuwa nau'ikan dankalin turawa sun rasa kimarsu a matsayin alamun alama na karatun yawan jama'a saboda bayyanar matsakaicin (ko kusa da shi) adadin kwayar halittar cutar cikin mafi yawan kebantattun mutane. A lokaci guda, har yanzu ana amfani da kwayar cutar ta T1 don ƙwayoyin tumatir ɗauke da kwayar Ph1 daidai a matsayin alama ta alama (Lavrova et al., 2003; Ulanova et al., 2003).
A cikin karatun da yawa, ana amfani da juriya na kayan gwari azaman alama. Wannan halayen ba shi da kyau a yi amfani da shi a cikin nazarin yawan jama'a saboda sauƙin bayyanar sauyin rikitarwa a cikin layuka bayan an yi amfani da metalaxyl- (ko mefenoxam-) dauke da kayan gwari a cikin filin. Misali, an nuna manyan bambance-bambance a cikin matakin juriya a cikin layin clonal na Sib1 (Elansky et al., 2001).
Don haka, nau'in mating, peptidase isoenzyme bakan, nau'in DNA na mitochondrial, RFLP-RG57, SSR sune mafi kyawun alamomin alama don ƙirƙirar bankunan bayanai da lakabin damuwa a cikin tarin. Don kwatanta iyakance samfuran, idan ya zama dole ayi amfani da iyakar adadin alamun alama, zaka iya amfani da AFLP, RAPD, InterSSR, Inter-SINE PCR (Table 5). Koyaya, yakamata a tuna cewa waɗannan hanyoyin basuda ƙarfin sake haifuwa, kuma a kowane gwaji na mutum (fadada electrophoresis sake zagayowar) ya zama dole ayi amfani da keɓaɓɓun bayanan keɓancewa.
Tebur 5. Kwatanta hanyoyin daban-daban na binciken damuwa P. estan tsaran gida
rarrab | TS | Isofer 'yan sanda | MtDNA | RFLP-RG57 | RAPD | ISSR | SSR | Farashin AFLP | Rev |
---|---|---|---|---|---|---|---|---|---|
Adadin bayanai | Н | Н | Н | С | В | В | С | В | В |
Sake haifuwa | В | В | В | В | Н | Н | С | С | С |
Yiwuwar kayan tarihi | Н | Н | Н | Н | В | С | Н | С | В |
kudin | Н | С | Н | В | Н | Н | Н | С | Н |
Laborarfin aiki | Н | Н | Н | В | NS * | NS * | Н | С | NS * |
Gudanar da bincike ** | В | Н | Н | С | Н | Н | Н | Н | Н |
Lura: H - low, C - matsakaici, B - babba; НС * - laborarfin aiki ya yi ƙasa yayin amfani da gel na agarose ko atomatik
genotyper, matsakaici - ta hanyar narkewa a cikin polyacrylamide gel tare da alamun share fage,
** - Bata kirdadon lokacin da aka bata wajen girma mycelium don kebancewar DNA.
Tsarin yawan jama'a
Lines na dangi
Idan babu sake hadewa ko kuma wata muhimmiyar gudummawa da yake bayarwa ga yawan jama'a, yawan mutane ya kunshi wasu kwayoyi masu yawa, musanyar kwayoyin halitta tsakanin su wanda ba safai ake samunsu ba.
A cikin irin wannan yawan jama'a, ya fi ba da ilmi don yin nazari ba game da mahimmancin kwayoyin halittar mutum ba, amma maɓuɓɓuka na jinsin halitta waɗanda ke da asali guda ɗaya (layin clonal ko layin jikoki) kuma ya bambanta ne kawai ta hanyar maye gurbi. Nazarin yawan jama'a game da cututtukan cututtukan ƙarshen cuta da kuma nazarin layin dunƙulewa sun haɓaka sosai tun bayan zuwan hanyar RFLP-RG57 a farkon shekarun 90 na karnin da ya gabata. Tare da RFLP-RG57, nau'in mating, nau'ikan peptidase da glucose-6-phosphate isomerase isoenzymes, da nau'in DNA na mitochondrial ana amfani dasu don gano layin dunƙulen. An nuna halaye na layin dunƙulen yau da kullun a cikin Table 6.
Clone US-1 ya mamaye yawan jama'a a ko'ina har zuwa ƙarshen shekarun 80, bayan haka an fara maye gurbinsa da wasu kwayoyi masu ɓoyi kuma ya ɓace daga Turai da Arewacin Amurka. Yanzu ana samun sa a Gabas mai nisa (Philippines, Taiwan, China, Japan, Korea, Koh et al., 1994, Mosa et al, 1993), a cikin Afirka (Uganda, Kenya, Rwanda, Goodwin et al, 1994, Vega-Sanchez et al., 2000; Ochwo et al., 2002) da kuma a Kudancin Amurka (Ecuador, Brazil, Peru, Forbes et al., 1997, Goodwin et al., 1994). Babu wani nau'in mallakar layin US-1 da aka gano a Ostiraliya kawai. A bayyane yake, P. infestans ware sun zo Ostiraliya tare da wani motsi na ƙaura (Goodwin, 1997).
Clone US-6 sun yi ƙaura daga arewacin Mexico zuwa California a ƙarshen 70s kuma suka haifar da annoba a can cikin dankali da tumatir bayan shekaru 32 ba tare da cuta ba. Saboda tsananin tashin hankali, sai ya kori Amurka-1 ya fara mamayewa a gabar yammacin Amurka (Goodwin et al., 1995a).
An gano jinsin jinsin na US-7 da US-8 a Amurka a shekarar 1992, kuma tuni a shekara ta 1994 an yada shi sosai a Amurka da Kanada. A lokacin kakar wasa guda, clone US-8 na iya kusan kawar da Amurka-1 a cikin makircin dankalin turawa da farko ya kamu da duka kwafin a daidai daidaitawa (Miller da Johnson, 2000).
Clones BC-1 zuwa BC-4 an gano su a cikin British Columbia a cikin ƙananan adadin keɓewa daga Goodwin et al., 1995b). Clone US-11 ya bazu sosai a cikin Amurka kuma ya maye gurbin US-1 a Taiwan. Clones JP-1 da EC-1, tare da clone US-1, sun zama ruwan dare a Japan da Ecuador, bi da bi (Koh et al., 1994; Forbes et al., 1997).
SIB-1 haɗuwa ce wacce ta mamaye a Rasha akan yanki mai faɗi daga yankin Moscow zuwa Sakhalin. A cikin yankin Moscow, an gano shi a cikin 1993, kuma wasu yankuna na filin sun kunshi nau'ikan nau'ikan wannan layin na dunƙule-tsalle, masu tsananin ƙarfin metalaxyl. Bayan 1993, yaduwar wannan clone ya ragu sosai. A waje da Urals a cikin 1997-1998, an sami SIB-1 ko'ina, ban da Yankin Khabarovsk (clone SIB-2 ya bazu a wurin). Rarraba sararin samaniya na clones tare da nau'ikan ma'aurata daban-daban banda tsarin jima'i a Siberia da Gabas ta Tsakiya. A cikin yankin Moscow, ya bambanta da Siberia, yawancin mutane suna wakiltar kwayoyi masu yawa; kusan kowane keɓaɓɓe yana da nau'ikan nau'ikan nau'ikan nau'ikan launuka daban-daban (Elansky et al., 2001, 2015). Ba za a iya bayanin wannan bambancin ba kawai ta hanyar shigo da nau'ikan gwari daga sassa daban-daban na duniya tare da kayan iri da aka shigo da su. Tunda nau'ikan sadarwar biyu suna faruwa a cikin jama'a, akwai yuwuwar cewa yawanta kuma saboda sake haɗuwa yake. Don haka, a cikin British Columbia, fitowar jinsin halittu BC-2, BC-3, da BC-4 ana ɗauka ne saboda haɗuwa da kwayoyi masu juna biyu BC-1 da US-6 (Goodwin et al., 1995b). Zai yiwu ana samun nau'ikan haɗuwa a cikin jama'ar Moscow. Misali, damuwa MO-4, MO-8 da MO-11 heterozygous na yankin PEP na iya zama haɗuwa tsakanin ɓarna MO-12, MO-21, MO-22, suna da nau'ikan A2 na maɗauran aure da kuma homozygous don duk wata alama ta yankin PEP da damuwa MO-8, suna da nau'in A1 na mating da homozygous don ɗayan ɗayan wuraren. Kuma idan haka ne, kuma a cikin al'ummomin zamani na P. infestans akwai yiwuwar haɓaka aikin jima'i, to ƙimar bayani game da binciken kwayoyi masu yawa zai ragu (Elansky et al., 2001, 2015).
Bambanci a cikin layi
Har zuwa 90s na karni na 20, layin dunƙulen US-1 ya yadu a duniya. Yawancin yankuna da yankuna sun ƙunshi nau'ikan damuwa tare da nau'in jinsin Amurka-1. Koyaya, an lura da bambance-bambance tsakanin keɓaɓɓu, mai yuwuwa ta hanyar tsarin maye gurbi. Maye gurbi ya faru a duka nukiliya da mitochondrial DNA kuma abin ya shafa, a tsakanin sauran abubuwa, matakin juriya ga magungunan phenylamide da yawan kwayoyin halittar virulence. Lines da suka banbanta da asalin halittar asali ta hanyar maye gurbi ana nuna su ta ƙarin lambobi bayan digon da ke bin sunan asalin asalin (alal misali, layin mutant na Amurka-1.1 na layin dunƙulen Amurka-1). Layin DNA na yatsan hannu US-1.5 da US-1.6 dauke da layukan kayan aiki masu girma dabam (Goodwin et al., 1995a, 1995b); layin dunƙulen Amurka-6.3 kuma ya bambanta da US-6 a layin kayan haɗi ɗaya (Goodwin, 1997, Table 7).
A cikin binciken DNA na mitochondrial, an gano cewa a cikin layin duniyan nan US-1 ana samun nau'in 1b na DNA na mitochondrial kawai (Carter et al., 1990). Koyaya, a cikin nazarin nau'ikan wannan jinsi daga Peru da Philippines, an sami keɓaɓɓun waɗanda nau'in DNA na mitochondrial ya banbanta da 1b ta wurin kasancewar sakawa da sharewa (Goodwin, 1991, Koh et al., 1994).
Table 6. Multilocus genotypes na wasu P. infestans clonal Lines
Title | Nau'in ma'aurata | Isozymes | Alamar yatsun DNA | Nau'in 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 |
DA-1-EY | 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 |
Fadakarwa: * - babu data.
Table 7. Multilocus genotypes da layin mutant
Title | Nau'in ma'aurata | | DNA Yatsa (RG57) | Bayanan kula | |
GPI | Farashin PEP-1 | ||||
US-1 | A1 | 86/100 | 92/100 | 1011101011001101000110011 | Na asali na asali 1 |
US-1.1 | A1 | 86/100 | 100/100 | 1011101011001101000110011 | Mutuwa a cikin PEP |
US-1.2 | A1 | 86/100 | 92/100 | 1011101010001101000110011 | Sauyawa a cikin RG57 |
US-1.3 | A1 | 86/100 | 92/100 | 1011101001001101000110011 | Sauyawa a cikin RG57 |
US-1.4 | A1 | 86/100 | 100/100 | 1011101010001101000110011 | Sauyawa a cikin RG57 da PEP |
US-1.5 | A1 | 86/100 | 92/100 | 1011101011001101010110011 | Sauyawa a cikin RG57 |
US-6 | A1 | 100/100 | 92/100 | 1011111001001100010110011 | Na asali na asali 2 |
US-6.1 | A1 | 100/100 | 92 /92 | 1011111001001100010110011 | Mutuwa a cikin PEP |
US-6.2 | A1 | 100/100 | 92/100 | 1011101001001100010110011 | Sauyawa a cikin RG57 |
US-6.3 | A1 | 100/100 | 92/100 | 1011111001011100010110011 | Sauyawa a cikin RG57 |
US-6.4 | A1 | 100/100 | 100/100 | 1011011001001100010110011 | Sauyawa a cikin RG57 da PEP |
US-6.5 | A1 | 100/100 | 92/100 | 1011111001001100010010011 | Sauyawa a cikin RG57 |
BR-1 | A2 | 100/100 | 100/100 | 1011101000001100001111011 | Na asali na asali 3 |
BR-1.1 | A2 | 100/100 | 100/100 | 1010101000001100001110011 | Sauyawa a cikin RG57 |
Hakanan akwai canje-canje a cikin bakan isozymes. A matsayinka na mai mulki, ana haifar da su ne ta hanyar lalacewar kwayar halitta da farko heterozygous don wannan enzyme din zuwa masu kama da juna. A cikin 1993, akan 'ya'yan tumatir, mun gano damuwa tare da halaye iri iri na US-1: zanen yatsun RG57, nau'in DNA na mitochondrial, da jinsin 86/100 na glucose-6-phosphatizomerase, amma ya kasance homozygous (100/100) don farkon peptidase wuri maimakon wani nau'in heterozygote na 92/100 kwatankwacin wannan layi. Mun sanya nau'in nau'in wannan nau'in MO-17 (Table 6). Layin mutant US-1.1 da US-1.4 suma sun bambanta da US-1 ta maye gurbi a farkon peptidase locus (Table 7).
Maye gurbi da ke haifar da canje-canje a cikin yawan kwayar halittar kwayar cuta don dankalin turawa da tumatir iri-iri ne. An lura da su tsakanin keɓaɓɓun layin duniyan nan US-1 a cikin yawan jama'a daga Netherlands (Drenth et al., 1994), Peru (Goodwin et al., 1995a), Poland (Sujkowski et al., 1991), arewacin Arewacin Amurka (Goodwin et al., ,, 1995b). An kuma lura da bambance-bambance a cikin yawan kwayoyin cututtukan dankalin turawa tsakanin kebantattun layukan kasashen US-7 da US-8 a Kanada da Amurka (Goodwin et al., 1995a), a tsakanin keɓaɓɓun layin SIB-1 a yankin Asiya na Rasha (Elansky et al, 2001) ).
An keɓance tare da bambance-bambance masu ƙarfi a cikin matakan juriya ga magungunan phenylamide a cikin yawan mutanen yankin monoclonal, dukansu na layin dunƙule ne Sib-1 (Elansky et al, 2001, Table 1). Kusan dukkan nau'ikan layin duniyan Amurka-1 suna da matukar damuwa da metalaxyl; duk da haka, an keɓance maƙasudin wannan layin a cikin Philippines (Koh et al., 1994) da kuma Ireland (Goodwin et al., 1996).
Yawan mutanen P. infestans
Amurka ta Tsakiya (Meziko)
Yawan 'yan ƙabilar P. a cikin Meziko ya bambanta sosai da sauran al'ummomin duniya, wanda hakan ya samo asali ne saboda matsayinsa na tarihi. Yawancin karatu na wannan yawan da alaƙa da nau'in halittar P. na Phytophthora, da na gida na jinsi na Solanum, sun kai ga ƙarshe cewa juyin halittar kwayar cuta a tsakiyar Mexico ya faru tare da juyin halittar shuke-shuke masu karɓar bakuncin kuma yana da alaƙa da sake saduwa da jima'i (Grünwald, Flier , 2005). Dukkanin nau'ikan sadarwar suna da wakilci a cikin jama'a, kuma daidai gwargwado, da kasancewar wadatar zafin a cikin ƙasa, a kan tsire-tsire da tubers na dankali da jinsunan da ke da alaƙa da Solanum ya tabbatar da kasancewar tsarin jima'i a cikin jama'a (Fernández-Pavía et al., 2002). Karatun da aka yi kwanan nan game da kwarin Toluca da kewayenta (cibiyar da ake zaton asalin kwayar cutar ce) ta tabbatar da babban bambancin kwayar halittar mazauna yankin na P. infestans (134 multilocus genotypes a cikin samfurin samfuran 176) da kuma kasancewar an samu rarrabuwar mutane da yawa a yankin (Wang et al., 2017). Abubuwan da ke ba da gudummawa ga wannan bambance-bambancen shi ne rarraba sararin samaniya na yawan mutanen da ke tsakiyar tsaunukan tsakiyar Mexico, bambance-bambance a cikin yanayin namo da iri dankalin turawa da ake amfani da su a kwari da tsaunuka, da kuma kasancewar wasu nau'ikan Solanum da ke dauke da igiyar ruwa wadanda za su iya aiki a matsayin madadin runduna (Fry et al. , 2009).
Koyaya, ya kamata a sani cewa yawan mutanen P. wadanda ke cikin arewacin Mexico sun fi girma a yanayi kuma sun fi kama da jama'ar Arewacin Amurka, wanda na iya nuna cewa waɗannan sabbin halittu ne (Fry et al., 2009).
Arewacin Amurka
Yawan jama'ar Arewacin Amurka na P. infestans koyaushe suna da tsari mai sauƙi kuma halayensu na yau da kullun an kafa su tun kafin amfani da microsatellite analysis. Har zuwa 1987, layin dunƙulen Amurka-1 ya mamaye Amurka da Kanada (Goodwin et al., 1995). A tsakiyar shekarun 70, lokacin da kayan gwari masu tushen metalaxyl suka bayyana, wannan kwafin ya fara maye gurbinsa da wasu, nau'ikan nau'ikan jinsin da suka yi kaura daga Mexico (Goodwin et al., 1998). A ƙarshen 90s. jinsin jinsin Amurka-8 ya maye gurbin kwayar halittar ta US-1 a cikin Amurka kuma ya zama babban layi a kan dankali (Fry et al., 2009; Fry et al., 2015). Halin ya bambanta da tumatir, wanda koyaushe ke ƙunshe da layuka masu yawa, kuma halayen su ya canza daga shekara zuwa shekara (Fry et al., 2009).
A shekara ta 2009, wata mummunar annoba ta ƙarshen ɓarkewar cuta ta ɓarke a Amurka akan tumatir. Wani fasali na wannan annobar ita ce kusan lokaci ɗaya a wurare da yawa a arewa maso gabashin Amurka, kuma ya zama yana da alaƙa da tallace-tallace masu yawa na ƙwayoyin tumatir masu kamuwa da cuta a cikin manyan cibiyoyin lambu (Fry et al., 2013). Asarar amfanin gona tayi yawa. Binciken Microsatellite na samfuran da abin ya shafa ya nuna cewa cutar ta annoba ta kasance cikin layin da ke kusa da US-22 A2. A cikin 2009, rabon wannan nau'in kwayar halitta a cikin jama'ar Amurka na P. infestans sun kai 80% (Fry et al., 2013). A cikin shekaru masu zuwa, yawan yaduwar kwayoyin halitta US-23 (akasari akan tumatir) da US-24 (akan dankali) ya karu a hankali a cikin jama'a, amma, bayan 2011, adadin gano-US-24 ya ragu sosai, kuma har zuwa yau, kusan 90% na yawan masu cutar a Amurka ta wakilci samfurin-US genotype (Fry et al., 23).
A cikin Kanada, kamar a Amurka, a ƙarshen 90s. US-1 ta maye gurbin babban jinsin US-8, manyan mukamansu sun kasance basu canza ba har zuwa 2008. A Kanada, akwai mummunan annobar cutar ɓarna da ke da nasaba da sayar da ƙwayoyin tumatir da ke kamuwa da cutar, amma sun samo asali ne daga jinsin halittar US-2009 da US-2010 (Kalischuk et al., 23). Bambancin bambancin yanayin wannan jinsin ya kasance abin birgewa: US-8 ta mamaye lardunan yamma na Kanada (2012%), yayin da US-23 ta mamaye lardunan gabas (68%). A cikin shekaru masu zuwa, US-8 ya bazu zuwa yankuna na gabas; amma, gabaɗaya, rabon da yake da shi a cikin jama'a ya ɗan ragu saboda asalin bayyanar jinsin US-83 da US-23 a cikin ƙasar (Peters et al., 22). Zuwa yau, US-24 na riƙe da matsayi mafi girma a cikin Kanada; US-2014 suna cikin British Columbia, yayin da US-23 da US-8 suna cikin Ontario (Peters, 23).
Don haka, jama'ar Arewacin Amurka na P. infestans yawanci layuka ne na dunƙule-tsalle. A cikin shekaru 40 da suka gabata, yawan kwayoyin halittar da aka gano sun kai 24. Duk da cewa nau'ikan nau'ikan nau'ikan saduwa iri biyu suna nan a cikin jama'a, yiwuwar bayyanar wasu sabbin kwayoyin halittar sakamakon sake haduwar jima'i ya kasance kadan. Koyaya, a cikin shekaru 20 da suka gabata, an rubuta lamura da yawa game da bayyanar yawan mutanen da suka sake haduwa (Gavino et al., 2000; Danies et al., 2014; Peters et al., 2014), kuma a wata harka, sakamakon tsallakawa ya kasance jinsin jinsin Amurka-11 , wanda aka kafa shi a Arewacin Amurka shekaru da yawa (Gavino et al., 2000). Har zuwa shekara ta 2009, canje-canje a cikin tsarin yawan jama'a yana da alaƙa da fitowar sabon, mafi yawan nau'ikan halittar jini tare da ƙaurarsu da kuma sauyawa daga magabata da suka gabata. Abin da ya faru a shekarar 2009-2010 A cikin Amurka da Kanada, epiphytotics a karon farko ya nuna cewa a zamanin dunkulewar duniya, ɓarkewar cutar na iya haɗuwa da yaduwar sabbin ƙwayoyin halittar jini yayin sayar da kayan shuka da suka kamu.
Kudancin Amurka
Har zuwa kwanan nan, nazarin mutanen Kudancin Amurka na P. infestans ba na yau da kullun bane ko babba. Sananne ne cewa tsarin waɗannan alumman suna da sauƙi kuma sun haɗa da layin duniyan 1-5 a kowace ƙasa (Forbes et al., 1998). Don haka, daga 1998, jinsin halittar US-1 (Brazil, Chile) BR-1 (Brazil, Bolivia, Uruguay, Paraguay), EC-1 (Ecuador, Colombia, Peru da Venezuela), AR-1, AR an samo kan dankali -2, AR-3, AR-4 da AR-5 (Ajantina), PE-3 da PE-7 (kudancin Peru). Nau'in shigar da kayan A2 ya kasance a cikin Brazil, Bolivia da Argentina kuma ba a same shi ba bayan iyakar Bolivian-Peruvian a yankin Tafkin Titicaca, wanda bayan EC-1 A1 genotype ya mamaye Andes. A kan tumatir, US-1 ta kasance mafi rinjaye a cikin Kudancin Amurka.
Yanayin da yawa ko orasa ya ci gaba a cikin 2000s. Wani mahimmin mahimmanci shine gano sabon layin duniyan EC-2 na nau'in A2 akan dangin daji na dankali (S. brevifolium da S. tetrapetalum) a Arewacin Andes (Oliva et al., 2010). Karatun phylogenetic ya nuna cewa wannan layin bai zama daidai da P. infestans ba, kodayake yana da kusanci da shi, a cikin wannan haɗin an ba da shawarar yin la'akari da shi, haka kuma wani layin, EC-3, wanda aka keɓe daga bishiyar tumatir S. betaceum da ke girma a cikin Andes, wani sabon nau'in da ake kira P. andina; duk da haka, matsayin wannan jinsin (nau'in mai zaman kansa ko wani nau'in P. infestans tare da wasu layin da ba'a sani ba) har yanzu ba a sani ba (Delgado et al., 2013).
A halin yanzu, duk jama'ar Kudancin Amurka na P. infestans suna da ƙarfi. Duk da kasancewar nau'ikan nau'ikan ma'aurata biyu, ba a sake gano wasu mutanen da ke sake haduwa ba. A kan tumatir, sinadarin US-1 yana da ko'ina, wanda ga alama ya raba shi da dankali daga matsalolin gida, wanda har yanzu ba a san asalinsa ba. A cikin Brazil, Bolivia da Uruguay, samfurin BR-1 ya kasance; a cikin Peru, tare da US-1 da EC-1, akwai wasu jinsin halittu na cikin gida da yawa. A cikin Andes, babban matsayi ana riƙe shi ta layin dunƙule EC-1, dangantakar da ke tare da kwanan nan da aka gano P. andina ya kasance ba a bincika ba. Kadai "m" wuri inda na tsawon 2003-2013. akwai canje-canje masu mahimmanci a cikin jama'a, ya zama Chile (Acuña et al., 2012), inda a cikin 2004-2005. yawan kwayoyin cutar ya zama halin juriya da metalaxyl da sabon DNA haplotype na mitochondrial (Ia maimakon Ib da yake a yanzu). 2006 zuwa 2011 A cikin jama'a, nau'in 21 (a cewar SSR) ya mamaye, wanda kasonsa ya kai 90%, bayan haka dabino ya wuce zuwa genotype 20, yawan abin da ya faru a cikin shekaru biyu masu zuwa ya kasance kimanin 67% (Acuña, 2015).
Turai
A cikin tarihin Turai, akwai aƙalla raƙuman ruwa guda biyu na ƙaura na P. 'yan iska daga Arewacin Amurka: a cikin ƙarni na 1. (HERB-1) da farkon karni na 70 (US-1). Rarraba yaduwar abubuwa masu narkewa na metalaxyl a cikin shekaru XNUMX. ya haifar da ƙaura daga cikin jinsin jinsin Amurka-XNUMX da maye gurbinsa da sababbin genotypes. A sakamakon haka, a mafi yawan ƙasashen Yammacin Turai, yawancin ƙwayoyin cuta suna da wakilai da yawa ta hanyar layuka masu yawa.
Amfani da nazarin microsatellite don nazarin yawan ƙwayoyin cuta ya ba da damar gano canje-canje masu tsanani da suka faru a Yammacin Turai a cikin 2005-2008. A 2005, an gano sabon layin clonal a cikin Burtaniya, wanda ake kira 13_A2 (ko "Blue 13") kuma yana da nau'ikan A2 na mating , babban tashin hankali da juriya ga phenylamides (Shaw et al., 2007). An samo irin wannan nau'in a cikin samfuran da aka tattara a 2004 a cikin Netherlands da arewacin Faransa, yana nuna cewa ta yi ƙaura zuwa Burtaniya daga nahiyoyin Turai, mai yiwuwa da dankali iri (Cooke et al., 2007). Nazarin kwayar halittar wakilan wannan layin dunƙule-karfen ya nuna wani babban matakin polymorphism na jerin sa (zuwa shekara ta 2016, adadin ƙananan bambance-bambancensa sun kai 340) da mahimmin mataki na bambanci a cikin matakin nuna kwayar halitta, incl. kwayoyin tasiri yayin kamuwa da tsire-tsire (Cooke et al., 2012; Cooke, 2017). Waɗannan fasalulluka, tare da ƙarin tsawan lokaci na yanayin biotrophic, na iya haifar da ƙaruwar tashin hankali na 13_A2 da ikon sa cutar har ma da nau'o'in dankalin turawa mai jure cutar mara.
A cikin fewan shekaru masu zuwa, genotype ya bazu cikin sauri a ƙasashen Arewa maso Yammacin Turai (Burtaniya, Ireland, Faransa, Belgium, Netherlands, Jamus) tare da sauya ƙaura daga jinsin halittu da suka gabata 1_A1, 2_A1, 8_A1 (Montarry et al., 2010; Gisi et al. , 2011; Van den Bosch et al., 2011; Cooke, 2015; Cooke, 2017). A cewar shafin yanar gizon www.euroblight.net, rabon 13_A2 a cikin yawan waɗannan ƙasashe ya kai 60-80% da ƙari; kasancewar irin wannan jinsin an kuma rubuta shi a wasu ƙasashe na Gabas da Kudancin Turai. Koyaya, a cikin 2009-2012. 13_A2 ta rasa manyan mukaminta a Burtaniya da Faransa, ta hanyar layin 6_A1 (8_A1 a Ireland), kuma a cikin Netherlands da Belgium an maye gurbin ta da wasu nau'ikan jinsi 1_A1, 6_A1, da 33_A2 (Cooke et al., 2012; Cooke, 2017; Stellingwerf, 2017).
Zuwa yau, kusan kashi 70% na Yammacin Turai na P. infestans monoclonal ne. A cewar shafin yanar gizon www.euroblight.net, manyan jinsin halittu a cikin ƙasashen Arewacin Yammacin Turai (Birtaniya, Faransa,
Netherlands, Belgium) sun rage, kusan a daidai gwargwado, 13_A2 da 6_A1, kuma ƙarshen ba zai faru ba a wajen yankin da aka ayyana (ban da Ireland), amma tuni yana da aƙalla ƙananan ƙananan 58 (Cooke, 2017). Bambancin 13_A2 sun kasance a cikin sanannun lambobi a cikin Jamus, kuma ana kiyaye su lokaci-lokaci a cikin ƙasashen Tsakiya da Kudancin Turai. Genotype 1_A1 ya zama babban ɓangare na yawan jama'ar Belgium da kuma ɓangaren Netherlands da Faransa. Genotype 8_A1 ya daidaita a cikin jama'ar Turai a matakin 3-6%, ban da Ireland, inda take riƙe da matsayinta na jagora kuma ta kasu kashi biyu (Stellingwerf, 2017). A karshe, a shekarar 2016, an lura da karuwar yawan sabbin kwayoyin halittar 36_A2 da 37_A2, wadanda aka fara rubutawa a 2013-2014, an lura; har zuwa yau, ana samun waɗannan nau'ikan jinsin a cikin Netherlands da Belgium da kuma wani ɓangare a Faransa da Jamus, da kuma a kudancin Burtaniya (Cooke, 2017). Kusan 20-30% na yawan Yammacin Turai suna wakiltar nau'ikan nau'ikan jinsi na musamman kowace shekara.
Ba kamar Yammacin Turai ba, a lokacin da halittar 13_A2 ta bayyana, yawan mutanen Arewacin Turai (Sweden, Norway, Denmark, Finland) ba a wakilta su da layin dunƙuƙulen wuri ba, amma ta yawancin jinsin halittu na musamman (Brurberg et al.,
2011). A lokacin yaduwar aiki na 13_A2 a Yammacin Turai, ba a lura da kasancewar wannan nau'in a cikin Scandinavia ba sai a shekarar 2011, lokacin da aka fara gano shi a Arewacin Jutland (D Denmarknemark), inda yawancin potanyen dankalin turawa ke girma tare da amfani da sinadarin metalaxyl kayan gwari (Nielsen et al., 2014). A cewar www.euroblight.net, an gano jinsin halittar 13_A2 a cikin samfuran da yawa daga Norway da Denmark a cikin 2014 kuma a cikin samfuran Yaren mutanen Norway da yawa a cikin 2016; bugu da kari, a shekarar 2013, an gano kasancewar 6_A1 na genotype a cikin wani adadi kaɗan a cikin Finland. Babban dalilin gazawar 13_A2 da sauran layukan dunkulalliya a mamayar Scandinavia ana ganin shine bambance-bambancen canjin yanayi na wannan yankin daga kasashen Yammacin Turai.
Toari da gaskiyar lokacin bazara da lokacin sanyi suna inganta rayuwar ƙarancin kayan lambu mai yawa kamar oospores (Sjöholm et al., 2013), daskarewar ƙasa a lokacin hunturu (wanda yawanci ba ya faruwa a ƙasashe masu dumi na Yammacin Turai) yana ba da gudummawa ga aiki tare da tsire-tsire da tsire-tsire. dankali, wanda ke inganta matsayinsu a matsayin tushen kamuwa da cuta na farko (Brurberg et al., 2011). Har ila yau, ya kamata a sani cewa, a cikin yanayin arewa, ci gaban kamuwa da cuta daga oospores ya wuce ci gaban kamuwa da cuta mai raɗaɗɗen ƙwayar cuta, wanda a ƙarshe ya hana mamayar mawuyacin hali, amma daga baya ya ci gaba da yin layi (Yuen, 2012). Tsarin mafi yawan mutanen da aka fi sani da P. 'yan iska a cikin ƙasashen Gabashin Turai (Poland, jihohin Baltic) suna kama da na Scandinavia.
Dukansu nau'ikan sadarwar suna nan a nan, kuma yawancin jinsin halittar da binciken SSR ya tantance sune na musamman (Chmielarz et al., 2014; Runno-Paurson et al., 2016). Kamar yadda yake a Arewacin Turai, rarraba layin dunƙule (musamman na 13_A2 genotype) kusan bai shafi mazaunan cikin gida na kwayar cutar ba, wanda ke riƙe da babban matakin bambancin tare da rashin manyan layuka.
Ana lura da kasancewar 13_A2 lokaci-lokaci a cikin filaye tare da nau'in dankalin turawa na kasuwanci. A Rasha, yanayin yana ci gaba kamar haka. Binciken Microsatellite na P. infestans ya ware a cikin 2008-2011 a cikin yankuna 10 daban-daban na ɓangaren Turai na Rasha, ya nuna babban mataki na bambancin jinsin halitta da kuma cikakken rashin daidaituwa da layin duniyan Turai (Statsyuk et al., 2014). Shekaru da yawa bayan haka, nazarin samfuran ƙanana na P. waɗanda aka tattara a yankin Leningrad a cikin 2013-2014 ya nuna manyan bambance-bambance a tsakanin su da jinsin wannan yanki da aka gano a cikin binciken da ya gabata. A cikin karatun biyu, ba a sami jinsin Turai na Yammacin Turai ba (Beketova et al., 2014; Kuznetsova et al., 2016).
Babban bambancin kwayar halittar Yammacin Turai na P. infestans da kuma rashin manyan layuka a cikinsu na iya zama saboda dalilai da yawa. Na farko, kamar a Arewacin Turai, yanayin yanayi na ƙasashen da aka yi la’akari yana ba da gudummawa ga samuwar oospores a matsayin tushen asalin kamuwa da cuta (Ulanova et al., 2010; Chmielarz et al., 2014). Na biyu, yawan dankalin turawa da aka samar a cikin waɗannan ƙasashe ana girma ne a kan ƙananan gonaki masu zaman kansu, galibi kewaye da dazuzzuka ko wasu cikas ga ƙauracewar ƙwayoyin cuta (Chmielarz et al., 2014). A ka’ida, dankalin da ake girma a irin wannan yanayin kusan ba a kula da shi da sinadarai, kuma zaɓin nau’ikan ya dogara ne da ƙarshen juriya na ƙarshe, watau babu wani matsin lamba na zafin nama da juriya ga metalaxyl, wanda ke hana genotypes masu juriya, kamar su 13_A2, na fa'idodi akan sauran nau'ikan halittar (Chmielarz et al., 2014). Aƙarshe, saboda ƙananan filayen filaye, masu su galibi basa yin juyawar amfanin gona, suna girma dankali a wuri ɗaya tsawon shekaru, wanda ke taimakawa ga tara nau'ikan inoculum na jinsi daban-daban (Runno-Paurson et al., 2016; Elansky, 2015; Elansky et al. ,, 2015).
Asiya
Har zuwa kwanan nan, tsarin mutanen P. mazauna Asiya sun kasance ba su da fahimta sosai. An san cewa ana wakiltar shi ne ta hanyar layuka masu kima, kuma tasirin sake haduwar jima'i akan bayyanar sabbin kayan halittu yanada kadan. Don haka, misali, a cikin 1997-1998. A cikin yankin Asiya na Rasha (Siberia da Gabas ta Tsakiya), yawancin kwayoyin halittu uku ne kawai suka wakilci masu cutar tare da fifikon jinsin SIB-1 (Elansky et al., 2001). An nuna kasancewar layukan ƙwayoyin cuta a cikin ƙasashe kamar China, Japan, Korea, Philippines, da Taiwan (Koh et al., 1994; Chen et al., 2009). Layin ƙirar Amurka-1, wanda ya mamaye babban yankin Asiya, a ƙarshen 90s - farkon 2000s. kusan ko'ina an fara maye gurbinsu da wasu nau'ikan halittar jini, wadanda, daga baya, suka ba da sababbi. A mafi yawan lokuta, canje-canje a cikin tsari da yanayin yawan jama'a a cikin kasashen Asiya suna da alaƙa da ƙaurawar sabbin nau'in halittu daga waje. Don haka, a Japan, ban da jinsin JP-3, duk wasu nau'ikan jinsin Jafananci waɗanda suka bayyana bayan US-1 (JP-1, JP-2, JP-3) suna da ƙarancin asalin asalin waje (Akino et al., 2011) ... A kasar Sin, a halin yanzu akwai manyan cututtukan cuta guda uku tare da cikakken yanki; Babu wata kwayar halitta mai rauni ko rauni sosai tsakanin waɗannan jama'ar (Guo et al., 2010; Li et al., 2013b). Genotype 13_A2 ya bayyana a yankin ƙasar China a lardunan kudu (Yunnan da Sichuan) a cikin 2005-2007, da 2012-1014. an kuma gani a arewa maso gabashin kasar (Li et al., 2013b). A Indiya, 13_A2 ya bayyana mai yuwuwa a lokaci guda kamar a China, mai yiwuwa yana dauke da dankalin da ke dauke da kwayar cutar (Chowdappa et al., 2015), kuma a cikin 2009-2010. ya haifar da mummunan cututtukan epiphytotic na ƙarshen cutar tumatir a kudancin ƙasar, bayan haka ya bazu zuwa dankali kuma a cikin 2014 ya haifar da ɓarkewar cutar a West Bengal, wanda ya haifar da lalacewa da kashe manoma da yawa na gida (Fry, 2016).
Afrika
Har zuwa 2008-2010 ba a aiwatar da karatun tsari na P. infestans a cikin kasashen Afirka ba. A yanzu haka, ana iya raba al'ummomin Afirka na P. infestans zuwa rukuni biyu, kuma wannan rarrabuwa tana da alaƙa a sarari tare da gaskiyar shigo da dankali daga Turai.
A Arewacin Afirka, wanda ke shigo da dankalin turawa daga Turai, nau'ikan A2 na mating yana da wakilci a kusan dukkanin yankuna, wanda ke ba da damar yiwuwar bayyanar sabbin halittu a sakamakon sake hadewar jima'i (Corbière et al., 2010; Rekad et al., 2017). Bugu da kari, a Aljeriya, kasancewar alamun jinsi 13_A2, 2_A1, da 23_A1 an lura dasu da nuna karfi na farkonsu, haka nan kuma a hankali ana samun raguwar nau'ikan jinsin halittu na musamman don kammala bacewa (Rekad et al., 2017). Ya bambanta da sauran yankin, a Tunisia (ban da arewa maso gabashin ƙasar), yawancin ƙwayoyin cuta suna wakiltar nau'in A1 na mating (Harbaoui et al., 2014).
Layin girma na NA-01 ya mamaye nan. Gabaɗaya, yawan adadin layukan duniyan a cikin jama'a shine kawai 43%. A Gabas da Kudancin Afirka, inda yawan shigo da iri ya yi kadan (Fry et al., 2009), P. 'yan iska ne kawai ke da wakilcin layin A1 guda biyu, US-1 da KE-1, kuma na biyun yana rarar tsohuwar kan dankali ( Pule et al., 2012; Njoroge et al., 2016). Zuwa yau, duk waɗannan nau'ikan jinsin suna da sanannen adadin bambancin subclonal.
Australia
Rahoton farko na jinkirin kamuwa da dankali a cikin Ostiraliya ya samo asali ne daga shekarar 1907, kuma epiphytotia na farko, mai yuwuwa ne sanadiyyar ruwan sama mai karfi a cikin watannin bazara, ya faru ne a cikin 1909-1911. (Drenth et al., 2002). Gabaɗaya, kodayake, jinkirin blight bashi da mahimmancin tattalin arziƙi ga ƙasar. Barkewar cutar lokaci zuwa lokaci, wanda yanayin yanayi ke bayarwa wanda ke samar da danshi mai zafi, ba ya faruwa fiye da sau daya duk bayan shekaru 5-7 kuma an fi saninsu a arewacin Tasmania da tsakiyar Victoria. Dangane da abin da ke sama, wallafe-wallafen da aka keɓe don nazarin tsarin jama'ar Australiya na P. ƙarancin dabbobi kusan ba su nan. Sabbin bayanan da aka samo daga 1998-2000. (Drenth et al., 2002). A cewar marubutan, yawan jama'ar jihar Victoria sun kasance tsatson dangin Amurka-1.3, wanda a kaikaice ya tabbatar da hijirar wannan nau'in halittar daga Amurka. Samfurori na Tasmania an rarraba su a matsayin AU-3, ya bambanta da jinsin halittar da ake da su a wancan lokacin a wasu sassan duniya.
Fasali na ci gaban ƙarshen buguwa a Rasha
A cikin Turai, kamuwa da cuta da aka gabatar tare da tubers iri, wanda ya mamaye ƙasa, da kuma zoosporangia da iska ta kawo daga shuke-shuke da aka tsiro daga filayen da aka dasa a filayen bara (shuke-shuke '' masu sa kai ''), ko kuma a kan tsibiyoyin waɗanda aka ɗora, ana ɗaukarsu a matsayin babban maganin rashin lafiya a kan dankali. alamar don ajiyar tubers. Daga cikin wadannan, ana daukar shuke-shuke da suka tsiro a tsibiyoyin tubers da aka jefar a matsayin mafi hatsarin tushen kamuwa da cuta. a can, yawan adadin tubers da suka toho galibi yana da mahimmanci, kuma ana iya ɗaukar zoosporangia daga gare su a kan nesa mai nisa. Sauran hanyoyin (oospores, "sa kai" shuke-shuke) basu da hatsari sosai, saboda ba al'ada ba ce shuka shuke-shuke a fannoni iri ɗaya fiye da sau ɗaya kowace shekara 3-4. Kamuwa da cuta daga tubers iri mai cuta shima kadan ne saboda kyakkyawan tsarin kula da ingancin iri.
Gabaɗaya, adadin inoculum a cikin jama'ar Turai yana da iyakancewa, sabili da haka ƙaruwar annobar ba ta da sauƙi kuma ana iya sarrafa shi cikin nasara ta amfani da shirye-shiryen fungicidal. Babban aiki a cikin yanayin Turai shine yaƙi da kamuwa da cuta a cikin lokacin lokacinda yaduwar zoosporangia daga tsire-tsire masu tasiri ya fara.
A Rasha, lamarin ya sha bamban. Mafi yawan noman dankalin turawa da tumatir ana shuka su ne a cikin kananan lambuna masu zaman kansu; ba a aiwatar da matakan kariya a kan su kwata-kwata, ko kuma ana yin magungunan fungicidal a cikin ƙarancin adadi kuma ana farawa bayan bayyanar marigayi ɓarna a saman. A sakamakon haka, lambunan kayan lambu masu zaman kansu sun zama tushen asalin kamuwa da cuta, wanda daga ita iska ke dauke da zoosporangia zuwa dasa kasuwancin. Wannan ya tabbata ne ta hanyar lura da mu kai tsaye a cikin Moscow, Bryansk, Kostroma, Ryazan yankuna: ana lalata lalacewar shuke-shuke a cikin lambuna masu zaman kansu tun kafin a fara maganin fungicide na dasa kasuwancin. Bayan haka, annoba a cikin manyan fannoni an taƙaita ta amfani da shirye-shiryen fungicidal, yayin da a cikin lambuna masu zaman kansu akwai saurin ci gaban marigayi.
Dangane da maganin rashin dacewa ko na "kasafin kuɗi" na dasa shuki na kasuwanci, har ila yau ana samun ɓarna a cikin filayen; daga baya suna haɓaka haɓaka, suna rufe manyan yankuna (Elansky, 2015). Kamuwa da cuta a cikin lambuna masu zaman kansu suna da tasirin gaske akan annoba a cikin fagen kasuwanci. A duk yankuna masu noman dankalin Rasha, yankin da dankali ya mamaye shi a lambuna masu zaman kansa ya nunka yawan filayen manyan masu kerawa sau da yawa. A cikin irin wannan yanayin, ana iya kallon lambunan kayan lambu masu zaman kansu azaman kayan aikin koyarwa na duniya don filayen kasuwanci. Bari muyi ƙoƙari don gano waɗancan kaddarorin waɗanda ke halayyar jinsin halittu a cikin lambuna masu zaman kansu.
Dasa shuki da rashin kera dankalin turawa, iri na tumatir da aka samu daga masu kera kasashen waje, noman dankali da tumatir na dogon lokaci a yankuna daya, maganin rashin kayan gwari ko rashin rashi gaba daya suna haifar da cututtukan epiphytoties a kamfanoni masu zaman kansu, sakamakon su kyauta ne hayewa, hadewa da samuwar wadatar iska a lambuna masu zaman kansu. A sakamakon haka, ana lura da bambancin jinsin halittu masu yawa, yayin da kusan kowace iri kebantacciya a cikin jinsinta (Elansky et al., 2001, 2015). Shuka dankali iri daban-daban daga asalin halittar da ke haifar da wuya cewa layukan dunkulallum wadanda suka kware wajen kai hari kan wani nau'ikan iri daban daban zasu fito. Thewayoyin da aka zaɓa a cikin irin wannan yanayin ana rarrabe su ta hanyar amfani da su dangane da nau'ikan da abin ya shafa, mafi yawansu suna da kusanci ga matsakaicin adadin ƙwayoyin cuta. Wannan ya sha bamban da tsarin "layin dunƙuƙulen dunƙule" na al'ada ga manyan filayen masana'antun noma tare da ingantaccen tsarin kariya daga kamuwa da cutar marigayi. "Lines na layi" (lokacin da duk wani nau'in kwayar cutar mai saurin lalacewa a filin yake da wakilci ɗaya ko fiye da haka) suna ko'ina cikin ƙasashe inda ake gudanar da noman dankalin turawa musamman ta manyan gonaki: Amurka, Netherlands, Denmark, da dai sauransu. A Ingila, Ireland, Poland, inda filayen gida kuma a al'adance suke yaɗuwa girma dankalin turawa, akwai kuma mafi girma genotypic bambancin a cikin gidãjen Aljanna masu zaman kansu. A ƙarshen karni na 20, "layukan dunƙuƙu" sun yadu a sassan Asiya da Gabas ta Tsakiya na Rasha (Elansky et al., 2001), wanda a bayyane yake saboda amfani da iri iri iri na musamman dan shuka. Kwanan nan, halin da ake ciki a waɗannan yankuna kuma ya fara canzawa zuwa ƙaruwa game da yawan jinsin mutane.
Rashin ingantattun magunguna tare da shirye-shiryen fungicidal na da wani, sakamako na kai tsaye - babu tarin ƙwayoyi masu juriya a cikin gidajen Aljanna. Tabbas, sakamakonmu yana nuna cewa ana samun ƙananan nau'in juriya na metalaxyl sau da yawa a cikin lambuna masu zaman kansu fiye da shuka na kasuwanci.
Kusancin dankalin turawa da tumatir, wanda aka saba da shi a lambun masu zaman kansu, yana sa hijirar damuwa tsakanin wadannan albarkatun gona, sakamakon hakan, a cikin shekaru goman da suka gabata, a tsakanin matsalolin da aka ware daga dankali, gwargwadon nauyin da ke dauke da kwayar halitta don juriya da nau'ikan tumatir na tumatir (T1), a da ana yinsa ne kawai don " tumatir "damuwa. Inswarewa tare da kwayar T1 a mafi yawan lokuta suna da saurin fushi ga duka dankali da tumatir.
A cikin 'yan shekarun nan, ƙarshen cutar tumatir ya fara bayyana a lokuta da dama fiye da na dankali. Ciyawar tumatir za a iya mamaye ta da ciyawar da ke cikin ƙasa, ko kuma wadatar da ke cikin ƙwayoyin tumatir ko manne da su (Rubin et al., 2001). A cikin shekaru 15 da suka gabata, yawancin wadatattun tsaba da aka shigo dasu, akasari wadanda aka shigo dasu, sun bayyana a shagunan, kuma yawancin kananan masu kera kayan sun canza zuwa amfani dasu. Theaƙƙarfan na iya ƙunsar ƙwayoyi iri iri waɗanda ke da alaƙar yankuna yankunan noman su. A nan gaba, wadannan nau'ikan jinsin an hade su a cikin tsarin jima'i a cikin lambuna masu zaman kansu, wanda ke haifar da bayyanar sabon jinsi gaba daya.
Don haka, ana iya bayyana cewa lambuna masu zaman kansu “tukunyar narkewa” ce ta duniya wanda, sakamakon musayar kayan ƙirar, ana aiwatar da nau'ikan jinsin da ke yanzu kuma sababbin sababbi sun bayyana. Bugu da ƙari, zaɓinsu yana faruwa a ƙarƙashin yanayin da ya sha bamban da waɗanda aka kirkira don dankali a manyan gonaki: rashin fungicidal press, daidaito iri-iri na shuka, fifikon shuke-shuke da ke fama da nau'ikan ƙwayoyin cuta da ƙwayoyin cuta, kusancin tumatir da daji na dare, ƙetare aiki da ƙoshin lafiya, yiwuwar don oospores suyi aiki azaman tushen kamuwa da cuta shekara mai zuwa.
Duk wannan yana haifar da babban bambancin jinsin al'umman bayan gida. A karkashin yanayin epiphytotic, ƙarshen bala'i yana yaduwa cikin hanzari a cikin lambunan kayan lambu kuma ana fitar da adadi mai yawa, wanda ke tashi zuwa dasa kasuwancin nan kusa. Koyaya, kasancewar sun shiga fagen kasuwanci tare da ingantaccen tsarin fasahar noma da kariya ta sinadarai, abubuwan da suka shigo ba su da wata dama ta fara epiphytotics a cikin filin, wanda hakan ya faru ne saboda rashin layukan dunƙule masu tsayayya da kayan gwari da ƙwarewa iri-iri da aka noma.
Wata hanyar ta asali na farko na iya kasancewa tubers marasa lafiya waɗanda ke cikin tsiron kasuwanci. Waɗannan tubers ɗin sun girma, a matsayin mai mulkin, a cikin filayen da ke da kyakkyawar fasahar noma da kariya ta sinadarai mai ƙarfi. Abubuwan jinsin wadanda aka ware wadanda suka shafi tubers an daidaita su da ci gaban nasu. Waɗannan nau'ikan suna da haɗari sosai don dasa kasuwanci fiye da tsarin ilimin asali daga lambuna masu zaman kansu. Sakamakon karatun mu kuma yana tallafawa wannan tunanin. Jama'a da aka keɓe daga manyan filaye tare da kiyaye sinadarai yadda ya kamata da kuma fasahar aikin gona mai kyau ba sa bambanta a cikin babban bambancin halittar jini. Yawancin lokaci waɗannan layuka ne masu yawa waɗanda ke da tsananin tashin hankali.
Rarara daga kayan shuka na kasuwanci na iya shigar da jama'a a cikin lambunan kayan lambu kuma su shiga cikin ayyukan da ke gudana a cikin su. Koyaya, a cikin lambun kayan lambu, fafatawarsu zata kasance ƙasa da ta filin kasuwanci, kuma ba da daɗewa ba zasu daina wanzuwa ta hanyar layin dunƙule, amma ana iya amfani da kwayoyin halittar su a cikin yawan "lambun".
Cutar da ta ɓullo akan tsire-tsire na '' sa kai '' da tarin tarin tubers yayin girbi bai dace da Rasha ba, saboda A cikin manyan yankuna masu girma na dankalin turawa na Rasha, ana lura da daskarewar ƙasa mai zurfin hunturu, kuma tsire-tsire daga tubers waɗanda suka yi damuna a cikin ƙasa ba safai suke haɓaka ba. Bugu da ƙari, kamar yadda gwaje-gwajenmu suka nuna, ƙarshen cututtukan ƙwayar cuta ba ya rayuwa a yanayin ƙarancin yanayi ko da na tubers waɗanda suka ci gaba da aiki. A yankin bushewa, inda ake yin noman dankali na farko, ƙarshen bushewa yana da wuya saboda yanayin rani da zafi.
Don haka, a halin yanzu muna lura da rarrabuwar kawunan mutanen P. zuwa cikin "filin" da kuma "yawan lambu". Koyaya, a cikin recentan shekarun nan, an lura da matakai da ke haifar da haɗuwa da haɗuwa da jinsin halittu daga waɗannan al'ummomin.
Daga cikin su, mutum na iya lura da cikakken ƙaruwa a cikin karatu na ƙananan masu kera, fitowar ƙananan fakiti masu araha na ɗanke dankali, yaɗuwar shirye-shiryen kayan gwari a cikin ƙananan fakiti, da kuma asarar tsoron “ilmin sunadarai” ta yawan jama’a.
Yanayi na faruwa ne lokacin da, saboda ƙwazon aikin mai siyarwa ɗaya, an shuka dukkanin ƙauyuka da tubers iri iri iri sannan aka samar musu da ƙananan fakiti iri ɗaya na maganin ƙwari. Ana iya ɗauka cewa za a sami dankali iri iri iri a kan dasa kasuwancin nan kusa.
A gefe guda kuma, wasu kamfanonin kasuwanci na magungunan kashe qwari suna inganta dabarun maganin sinadarai na "kasafin kudi". A wannan yanayin, ba a raina adadin magungunan da aka ba da shawara kuma ana bayar da kayan arha mafi arha, kuma ba a mai da hankali kan hana ci gaban ɓarkewar tashin hankali ba har zuwa yanke saman, amma a kan wani jinkiri a cikin epiphytoty don ƙara yawan amfanin ƙasa. Irin waɗannan tsare-tsaren suna da hujjar tattalin arziki lokacin da ake shuka dankali daga kayan ƙasa mai ƙarancin ƙarfi, alhali kuwa bisa ƙa'ida babu batun samun ɗimbin riba. Koyaya, a wannan yanayin, ya bambanta da yawan mutanen lambun, yanayin asalin ɗankalin da dankalin turawa ya bayar ya ba da gudummawar zaɓin takamaiman jinsi na ilimin kimiyyar lissafi, waɗanda ke da haɗari sosai ga wannan nau'ikan.
Gabaɗaya, abubuwan da ake nufi da haɗuwa da hanyoyin "lambu" da "filin" na samar da dankalin turawa suna neman zama mana haɗari. Don hana mummunan sakamakonsu, a cikin gida da kuma sassan kasuwanci, zai zama wajibi ne a sarrafa duka nau'ikan iri na dankali da kewayon kayan gwari da aka ba masu keɓaɓɓu a cikin ƙananan marufi, da bin diddigin tsare-tsaren kariya da dankalin turawa da kuma amfani da shirye-shiryen kayan gwari a ɓangaren kasuwanci.
A cikin yankunan kamfanoni masu zaman kansu, akwai ci gaba mai ƙarfi ba kawai ɓacin lokaci ba, har ma da Alternaria. Yawancin masu mallakar filaye masu zaman kansu ba sa ɗaukar matakai na musamman don kariya daga Alternaria, suna ɓatar da cigaban Alternaria don ɓarnatarwar saman ko ci gaban latti. Sabili da haka, tare da babban ci gaban Alternaria akan ire-iren abubuwa masu saukin kamuwa, filayen gida na iya zama tushen tushen inoculum don kasuwancin shuka.
Hanyoyin canzawa
Tsarin maye gurbi
Tunda faruwar lamarin maye gurbi tsari ne wanda yake gudana ba tare da wata matsala ba, faruwar lamarin maye gurbi a kowane wuri ya dogara da yawan canjin wannan wurin da kuma yawan mutane. Lokacin nazarin yawan maye gurbi na P. infestans damuwa, yawan yankuna da suka girma akan zaɓaɓɓen kafofin watsa labarai na abinci mai gina jiki bayan jiyya tare da mutagens na jiki ko na jiki yawanci ana ƙaddara su. Kamar yadda ake iya gani daga bayanan da aka gabatar a cikin Table 8, saurin maye gurbi na iri ɗaya a wuri daban-daban na iya bambanta da umarni da yawa na girma. Babban yawan maye gurbi a cikin juriya da metalaxyl na iya zama ɗayan dalilan tarin ƙwayoyin dake jurewa da ita a yanayi.
Mitar lokaci-lokaci ko maye gurbi, wanda aka lasafta shi bisa ga gwaje-gwajen dakunan gwaje-gwaje, ba koyaushe yake dacewa da matakan da ke faruwa a cikin al'umman ƙasa ba, saboda dalilai masu zuwa:
1. Tare da rikice-rikicen makaman nukiliya, ba shi yiwuwa a kimanta yawan maye gurbi a kowane ƙarni ɗaya na nukiliya. Sabili da haka, yawancin gwaje-gwajen suna ba da bayani ne kai tsaye game da yawan maye gurbi, ba tare da rarrabewa tsakanin abubuwa biyu na maye gurbi da wani abu da ya biyo bayan mitosis ba.
2. Sauyawa sau da yawa yakan rage daidaituwar kwayar halitta, saboda haka, tare da sayan sabuwar kadara, gaba dayan kwayar halittar tana raguwa. Yawancin maye gurbi na gwaji yana da rage tashin hankali kuma ba'a rubuta shi a cikin jama'ar ƙasa ba. Don haka, daidaito tsakanin daidaituwar juriya na P. infestans mutants zuwa sinadarin phenylamide da haɓakar haɓaka akan mahalli na wucin gadi ya kasance matsakaita (-0,62), da juriya ga kayan gwari da tashin hankali akan ganyen dankalin turawa (-0,65) (Derevyagina et al. , 1993), wanda ke nuna ƙarancin ƙarancin yanayin mutants. Har ila yau, maye gurbin juriya ga dimethomorph ya kasance tare da raguwar saurin aiki (Bagirova et al., 2001).
3. Mafi yawa daga rikitarwa da rikirkita rikirkitattun abubuwa ba su da tasiri kuma ba sa bayyana kansu a zahiri a gwaje-gwajen, amma sun zama wata buyayyar wuri ta bambancin yanayin halittun. Mutananan mutant da aka keɓe a cikin gwaje-gwajen gwaje-gwaje suna ɗaukar rinjaye ko maye gurbinsu gaba ɗaya (Kulish da Dyakov, 1979). A bayyane yake, difiloma na nukiliya ya bayyana ƙoƙarin da ba a yi nasara ba don samo maye gurbi a ƙarƙashin tasirin tasirin iska ta UV wanda ke da lahani a kan nau'ikan da ke da juriya a baya (McKee, 1969). Dangane da lissafin marubucin, irin wannan maye gurbi na iya faruwa tare da saurin da bai gaza 1: 500000 ba. Canjin yanayi na maye gurbi zuwa maye gurbi, yanayin da aka bayyana a bayyane zai iya faruwa saboda jima'i ko sake haifar da jima'i (duba ƙasa). Koyaya, koda a wannan yanayin, ana iya rufe maye gurbi ta hanyar manyan alamomin nau'ikan halittar daji a cikin cenotic (multinucleated) mycelium kuma an daidaita su ne kawai a lokacin samuwar zoospores mononuclear.
Tebur 8. Mitar P. na maye gurbi zuwa abubuwa masu hana girma a ƙarƙashin aikin nitrosomethylurea (Dolgova, Dyakov, 1986; Bagirova et al., 2001)
Haɗin kai | Tsarin maye gurbi |
Oxytetracycline | 6,9 10 x-8 |
Labarin 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 |
Har ila yau, yawan mutane yana taka muhimmiyar rawa wajen aukuwar sauyin yanayi. A cikin adadi mai yawa, wanda yawan ƙwayoyin N> 1 / a, inda a shine ƙimar maye gurbi, maye gurbi ya daina zama baƙon abu (Kvitko, 1974).
Lissafi na nuna cewa tare da matsakaicin matsakaicin gonar dankalin turawa (tabo 35 a kowace shuka), ana samar da fuloti 8x1012 a kowace rana a hekta daya (Dyakov da Suprun, 1984). A bayyane yake, irin waɗannan al'ummomin suna ƙunshe da duk maye gurbi da izinin musayar yake a kowane wuri. Ko da sauyin yanayi, wanda ke faruwa tare da mita 10-9, mutane dubu ne zasu saya daga miliyoyin da ke rayuwa a hekta daya a filin dankalin turawa. Don maye gurbi da ke faruwa tare da maɗaukakiyar mita (misali, 10-6), a cikin irin wannan yawan, sauye-sauye masu haɗuwa da juna na iya faruwa kowace rana (lokaci guda a loci biyu), watau tsarin canjin yanayi zai maye gurbin sake hadewa.
Hijira
Ga 'yan infestans, an san manyan nau'ikan ƙaura biyu: don rufe nesa (a cikin filin dankalin turawa ko filayen maƙwabta) ta hanyar yada zoosporangia ta igiyar iska ko feshin ruwan sama, da kuma zuwa nesa mai nisa - tare da dasa shuki na tubers ko tumatir' ya'yan itacen da aka ɗora. Hanya ta farko ta tanadar da faɗaɗa abin da cutar ta fi mayar da hankali, na biyu - ƙirƙirar sabbin masarufi a wuraren da ke nesa da na farko.
Yaɗuwar kamuwa da ƙwayoyin tumatir da fruitsa fruitsan itace ba kawai yana ba da gudummawar ɓullar cutar ba a sabbin wurare, amma kuma shine babban tushen bambancin ƙwayoyin halittu a cikin jama'a. A cikin yankin Moscow, an shuka dankali, an kawo shi daga yankuna daban-daban na Rasha da Yammacin Turai. Ana kawo 'ya'yan tumatir daga yankunan kudancin Rasha (yankin Astrakhan, yankin Krasnodar, Arewacin Caucasus). Itatuwan tumatir, wanda kuma zai iya zama tushen kamuwa da cuta (Rubin et al., 2001), ana kuma shigo da su daga yankunan kudancin Rasha, China, ƙasashen Turai da sauran ƙasashe.
Dangane da lissafin da E. Mayr (1974) yayi, canjin dabi'un halittu da akayi a wani gari wanda yake haifar da maye gurbi da wuya ya wuce 10-5 a kowane guri, yayin da yake a bayyane, musayar saboda yawan kwararar kwayoyin halitta akalla 10-3 - 10-4.
Hijira a cikin tubers da ke dauke da kwayar cutar ita ce sanadiyyar shigar da 'yan iska na P. zuwa Turai, yana yaduwa zuwa dukkan yankuna na duniya inda ake noman dankali; sun haifar da canje-canje mafi yawan mutane. Larshen bala'i akan dankali ya bayyana a yankin daular Rasha kusan lokaci guda tare da bayyanarsa a Yammacin Turai.
Tunda aka fara lura da cutar a cikin 1846-1847 a cikin Yankin Baltic kuma kawai a cikin shekaru masu zuwa suka bazu a Belarus da yankunan arewa maso yamma na Rasha, asalin Yammacin Turai bayyane yake. Tushen farko na ƙarshen ɓarna a cikin Tsohuwar Duniya ba a bayyane yake ba. Maganar da Fry et al. (Fry et al., 1992; Fry, Goodwin, 1995, Goodwin et al., 1994) ya ba da shawarar cewa cutar ta fara ne daga Mexico zuwa Arewacin Amurka, inda ta bazu a kan amfanin gona, sannan aka kai ta Yammacin Turai. (Fig. 7).
Sakamakon guguwar da aka maimaita (sakamako biyu na "kwalbar kwalbar"), kwayoyi guda sun isa Turai, zuriyarsu sun haifar da annoba a ko'ina cikin ƙasar Tsohuwar Duniya inda ake shuka dankali. A matsayin shaidar wannan tsinkayen, marubutan sun bayyana, da farko, yaduwar nau'ikan nau'ikan ma'aurata guda daya (A1) kuma, abu na biyu, daidaituwar jinsin jinsin da aka yi nazari daga yankuna daban daban (dukkansu suna kan alamomin kwayoyin ne, gami da 2 isozyme loci, alamun yatsun DNA, da tsarin mitochondrial DNA iri daya ne, kuma yayi daidai da clone US-1 da aka bayyana a Amurka). Koyaya, wasu bayanan suna haifar da shakku game da aƙalla wasu daga cikin tanadin abin da aka ambata. Tattaunawa game da P. infestans mitochondrial DNA da aka keɓe daga samfuran dankalin turawa da ke ɗauke da cutar a lokacin farkon epiphytotic na 40s ya nuna cewa sun bambanta a cikin tsarin mitochondrial DNA daga clone US-1, wanda, saboda haka, ya kasance aƙalla ba kawai tushen kamuwa da cuta a Turai ba (Ristaino et al, 2001).
Matsayin marigayi na ƙarshe ya sake tsanantawa a cikin 80s na karni na XX. Wadannan canje-canje sun faru:
1) Matsakaicin tashin hankali na jama'a ya karu, wanda hakan ya haifar da shi, musamman, zuwa yaduwar mummunar hanyar cutarwa ta marigayi - lalacewar petioles da mai tushe.
2) Ya kasance akwai motsi a lokacin ƙarshen buguwa a kan dankali - daga ƙarshen Yuli zuwa farkon Yuli har ma zuwa ƙarshen Yuni.
3) Nau'in A2, wanda a baya baya cikin Tsohuwar Duniya, ya zama ko'ina.
Sauye-sauye sun gabaci abubuwa biyu: yawan amfani da sabon kayan gwari metalaxyl (Schwinn da Staub, 1980) da kuma bayyanar Mexico a matsayin mai fitar da dankali a duniya (Niederhauser, 1993). Dangane da wannan, an gabatar da dalilai guda biyu don canjin yawan mutane: jujjuyawar nau'in ma'adanai a ƙarƙashin tasirin metalaxyl (Ko, 1994) da kuma gabatarwar sabbin ƙwayoyi tare da tubers masu cutar daga Mexico (Fry da Goodwin, 1995). Kodayake ba Ko kawai ya samo rikice-rikice na nau'ikan saduwa a ƙarƙashin tasirin metalaxyl ba, har ma a cikin ayyukan da aka gudanar a dakin gwaje-gwaje na Jami'ar Jihar Moscow (Savenkova, Chherepennicova-Anikina, 2002), zato na biyu ya fi kyau. Tare da bayyanar nau'ikan nau'ikan na biyu, canje-canje masu mahimmanci sun faru a cikin jinsin jinsin jinsin P. P. na Rasha, gami da cikin ƙwayoyin halitta masu tsaka (isozyme da RFLP loci), da kuma cikin tsarin mitochondrial DNA. Ba za a iya bayanin hadaddun waɗannan canje-canje ta hanyar aikin metalaxyl ba; a'a, akwai wadatattun shigo da sababbin matsaloli daga Mexico, wanda, kasancewa mafi tsananin tashin hankali (Kato et al., 1997), ya raba tsofaffin damuwa (US-1), ya zama rinjaye a cikin jama'a. Canji a cikin ƙididdigar yawan jama'ar Turai ya faru a cikin ɗan gajeren lokaci - daga 1980 zuwa 1985 (Fry et al., 1992). A kan yankin tsohuwar USSR, an sami “sababbin damuwa” a cikin tarin daga Estonia a cikin 1985, wato, a baya fiye da Poland da Jamus (Goodwin et al., 1994). Lokaci na karshe da "tsohuwar damuwa US-1" a Rasha ta keɓe daga tumatir da ke kamuwa da cutar a yankin Moscow a cikin 1993 (Dolgova et al., 1997). Har ila yau a Faransa, an sami nau'in "tsofaffi" a cikin shukar tumatir har zuwa farkon shekarun 90, ma'ana, bayan sun daɗe da ɓacewa a kan dankali (Leberton da Andrivon, 1998). Canje-canje a cikin matsalolin 'yan ƙabilar P. ya shafi halaye da yawa, gami da waɗanda ke da mahimmancin amfani, kuma ya ƙaru da cutarwar marigayi
Haɗuwa da jima'i
Domin sake saduwa da jima'i don ba da gudummawa ga bambance-bambancen, ya zama dole, na farko, kasancewar nau'ikan nau'ikan saduwa biyu a cikin jama'a a cikin rabo kusa da 1: 1, kuma, na biyu, kasancewar canjin yawan jama'a na farko.
Rabon nau'ikan ma'aurata ya bambanta sosai a cikin alumomi daban-daban har ma a cikin shekaru daban-daban a cikin jama'a ɗaya (Table 9,10, 90). Ba a san dalilan da ke haifar da irin wannan canjin canjin a cikin mitar nau'ikan nau'ikan ma'aurata a cikin jama'a ba (kamar, alal misali, a Rasha ko a Isra'ila a farkon shekarun 2002 na karnin da ya gabata), amma an yi imanin cewa hakan ya faru ne saboda gabatarwar wasu kwayoyi masu gasa (Cohen, XNUMX).
Wasu bayanan kai tsaye suna nuna yadda tsarin jima'i yake a wasu shekaru kuma a wasu yankuna:
1) Nazarin yawan jama'a daga yankin Moscow ya nuna cewa a cikin yawan mutane 13 wanda rabon nau'in nau'in A2 bai kai 10% ba, jimillar jinsin halittu da aka lissafa don isozyme loci uku ya kasance 0,08, kuma a cikin mutane 14 wanda rabon A2 ya wuce 30%, bambancin kwayoyin ya ninka biyu (0,15) (Elansky et al., 1999). Don haka, mafi girman yuwuwar yin jima'i, ya fi girma game da bambancin halittar jama'a.
2) An lura da alaƙar da ke tsakanin yanayin nau'ikan sadarwar aure a cikin yawan jama'a da ƙarfin samuwar oospore a cikin Isra'ila (Cohen et al., 1997) da kuma Holland
(Flier et al., 2004). Karatunmu ya nuna cewa a cikin yawan mutanen da ke kebance da nau'in A2 na ma'aurata sun kai 62, 17, 9, da 6%, an samu oospores a cikin 78, 50, 30, da 15% na ganyen dankalin turawa (da ke da maki 2 ko fiye).
Samfurori tare da 2 ko sama da tabo sun fi dacewa da ƙunshin abubuwa fiye da samfuran tare da tabo 1 (32 da 14% na samfuran, bi da bi) (Apryshko et al., 2004).
Oospores sun fi yawa a cikin ganyen tsakiya da ƙananan layin dasa dankalin turawa (Mytsa et al., 2015; Elansky et al., 2016).
3) A wasu yankuna, an gano nau'ikan jinsin halittu na musamman, wanda ake alakanta shi da sake saduwa da jima'i. Don haka, a cikin Poland a cikin 1989 da Faransa a 1990, nau'ikan homozygous na glucose-6-
isomerase na phosphate (GPI 90/90). Tunda a baya kawai an sadu da heterozygotes 10/90 kawai tsawon shekaru 100, ana danganta homozygosity da sake haɗuwa da jima'i (Sujkowski et al., 1994). A Colombia (Amurka), keɓance haɗa A2 da GPI 100/110 da A1 tare da GPI 100/100 gama gari ne, amma a ƙarshen lokacin 1994 (Agusta 16 da 9 ga Satumba), damuwa da nau'ikan halittar jini (A1 GPI 100/110) da A2 GPI 100/100) (Miller et al., 1997).
4) A cikin wasu al'ummomi daga Poland (Sujkowski et al., 1994) da Arewacin Caucasus (Amatkhanova et al., 2004), rarraba yatsan DNA na yatsa da lozy protein loci ya dace da rarraba Hardy-Weinberg, wanda ke nunawa
game da babban rabo na gudummawar saduwa da jima'i zuwa bambancin yawan jama'a. A wasu yankuna na Rasha, ba a sami wata wasiƙa ba game da rarraba Hardy-Weinberg a cikin yawan jama'a, amma an nuna kasancewar haɗin ƙwayar cuta, yana nuna fifikon haifuwa a cikin ƙira (Elansky et al., 1999).
5) Bambancin kwayar halitta (GST) tsakanin damuwa tare da nau'ikan nau'ikan ma'aurata (A1 da A2) sun kasance ƙasa da tsakanin mutane daban-daban (Sujkowski et al., 1994), wanda a kaikaice ke nuna gicciyen jima'i.
A lokaci guda, gudummawar sake haduwar jima'i zuwa bambancin jama'a ba zai iya zama mai yawa ba. An kirga wannan gudummawar don yawan jama'ar yankin Moscow (Elansky et al., 1999). Dangane da lissafin Lewontin (1979) “sake hadewa, wanda zai iya samar da sabbin bambance-bambancen daga loci biyu tare da mitar da bata wuce samfurin heterozygosities dinsu ba, zai zama yana da tasiri ne kawai idan dabi’un heterozygosity na duka allles sun riga sun zama manya”.
Tare da nauyin nau'ikan nau'i biyu, wanda ya saba da yankin Moscow, daidai yake da 4: 1, yawan sake haɗawa zai zama 0,25. Yiwuwar yiwuwar ƙetare damuwa zai kasance mai ban sha'awa ga biyu daga cikin ukun da aka yi nazarin isozygous loci a cikin yawan mutanen da aka yi nazari shine 0,01 (damuwa 2 daga 177). Sakamakon haka, yiwuwar faruwar heterozygotes sau biyu sakamakon sake hadewa kada ya wuce samfurin su wanda aka ninka shi da yiwuwar wucewa (0,25x0,02x0,02) = 10-4, watau masu haɗuwa da jima'i yawanci basa fada cikin samfurin nazarin damuwa. Waɗannan lissafin an yi su ne don yawan jama'a daga yankin Moscow, waɗanda ke da ɗan bambanci sosai. A cikin yawan mutane kamar na Siberia, tsarin jima'i, koda kuwa ya faru ne a cikin daidaikun jama'a, ba zai iya yin tasiri ga bambancin jinsinsu ba.
Bugu da ƙari, P. infestans yana da halin saurin chromosome a cikin meiosis, wanda ke haifar da aneuploidy (Carter et al., 1999). Irin wannan take hakki ya rage yawan haihuwa na matasan.
Sake haɗuwa da 'yar madigo, juzu'in juzu'in halitta
A cikin gwaje-gwajen da aka yi akan yaduwar cututtukan P. masu haɗari tare da maye gurbi don juriya da masu hana ci gaban daban, an sami fitowar misolates masu tsayayya ga masu hana su biyu (Shattock da Shaw, 1975; Dyakov, Kuzovnikova, 1974; Kulish, Dyakov,
1979). Rashin damuwa ga masu hana ci gaban guda biyu ya tashi ne sakamakon jujjuyawar maganin mycelium, kuma a wannan yanayin sun manne yayin haifuwa ta zoospores mononuclear (Judelson, Ge Yang, 1998), ko kuma ba su rabe a cikin zuriya ba, saboda suna da tetraploid (tun lokacin da aka keɓe su na farko diploid) nuclei (K , 1979). Heterozygous diflomasiyya da aka keɓe a ƙananan mitar saboda haploidization, chromosome nondisjunction, da kuma mitotic mararraba a kan (Poedinok et al., 1982). Za'a iya ƙara yawan waɗannan matakan tare da taimakon wasu tasirin tasirin difloma na heterozygous (alal misali, saka ƙwallafawar UV na ƙwayoyin cuta).
Kodayake samuwar ganyayyaki tare da juriya biyu yana faruwa ba kawai a cikin vitro ba, har ma a cikin tubers na dankalin turawa da ke haɗuwa da cakuda masu rikida (Kulish et al., 1978), yana da wuya a tantance rawar da ake samu na sake saduwa da mace da namiji a cikin ƙarni na sabbin jinsin halittu a cikin jama'a. Mitar segregants da aka samu sakamakon fashewar abubuwa, rashin hada-hada na chromosomes da tsallakawa ta hanyar mitotic ba tare da wani tasiri na musamman ba zai zama abin kulawa (kasa da 10-3).
Abinda ke faruwa na segregants na homozygous na cututtukan heterozygous na iya dogara ne akan ƙetarewar mitotic da canjin canjin mitotic, wanda a cikin P. sojae ke faruwa da saurin 3 x 10-2 zuwa 5 x 10-5 a kowane wuri, ya dogara da damuwa (Chamnanpunt et al. , 2001).
Kodayake yawan abin da ya faru na heterokaryons da difloma na heterozygous ya zama ya zama babba ba zato ba tsammani (ya kai kashi goma cikin ɗari), wannan aikin yana faruwa ne kawai lokacin da aka jujjuya al'adun mutant da aka samo daga iri ɗaya. Lokacin amfani da nau'ikan iri daban daban daga yanayi, heterokaryotization baya faruwa (ko yana faruwa tare da ƙananan ƙananan mita) saboda kasancewar rashin jituwa ta ciyayi (Poedinok da Dyakov, 1981; Anikina et al., 1997b; Cherepennikova-Anikina et al., 2002). Sakamakon haka, za a iya rage rawar recombination na ɗan luwaɗi kawai zuwa sake haɗawar intraclonal a cikin heterozygous nuclei da kuma sauyawar halittar mutum zuwa yanayin homozygous ba tare da tsarin jima'i ba. Wannan tsari na iya zama muhimmiyar mahimmanci na annoba a cikin damuwa tare da rikicewar rikicewa ko maye gurbin maye gurbin kayan gwari. Canjin sa zuwa wani yanayi na homozygous saboda aiwatarwar da ake yi tsakanin maza da mata zai kara karfin juriya mai rikida (Dolgova, Dyakov, 1986).
Gabatarwar kwayoyin halitta
Hetrothallic jinsin Phytophthora suna da ikon yin haɗuwa tare da samuwar ƙananan oospores (duba Vorob'eva da Gridnev, 1983; Sansome et al., 1991; Veld et al., 1998). Naturalarancin halittar jinsin Phytophthora guda biyu ya kasance mai tsananin tashin hankali wanda ya kashe dubunnan alder a Burtaniya (Brasier et al., 1999). P. infestans na iya faruwa tare da wasu nau'ikan jinsi (P. erythroseptica, P. nicotianae, P. Cactorum, da sauransu) akan tsire-tsire masu masaukin baki da kuma cikin ƙasa, amma akwai ɗan bayanai kaɗan a cikin wallafe-wallafen game da yiwuwar cakudaddu. A karkashin yanayin dakin gwaje-gwaje, an sami matasan tsakanin P. infestans da P. Mirabilis (Goodwin da Fry, 1994).
Tebur 9. Matsayin P. infestans damuwa tare da A2 mating type a kasashe daban-daban na duniya a tsakanin 1990 zuwa 2000 (gwargwadon bayanan maɓuɓɓun wallafe-wallafe da shafuka www.euroblight.net, www.eucablight.org)
kasar | 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) | ||||||||||
Ingila | 4 (26) | 3 (630) | 9 (336) | ||||||||
Finland | 0 (15) | 19 (117) | 12 (16) | 21 (447) | 6 (509) | 9 (432) | 43 (550) | ||||
Faransa | 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) | ||||||||||
Arewa. Ireland | 10 (41) | 9 (58) | 1 (106) | 0 (185) | 0 (18) | 0 (56) | 0 (35) | 0 (26) | |||
Netherlands | 7 (41) | 5 (276) | 24 (377) | 44 (353) | 23 (185) | ||||||
Norway | 25 (446) | 28 (156) | 8 (39) | 18 (257) | 38 (197) | ||||||
Peru | 0 (34, 1984 -86) | 0 (287, 1997-98) | 0 (112) | 0 (66) | |||||||
Poland | 19 (180) | 21 (142) | 33 (256) | 26 (149) | 35 (70) | ||||||
Scotland | 25 (147) | 11 (163) | 22 (189) | 5 (22) | |||||||
Sweden | 25 (263) | 62 (258) | 49 (163) | ||||||||
Wales | 0 (16) | 7 (97) | 0 (48) | 0 (25) | |||||||
Koriya | 36 (42) | 10 (130) | 15 (98) | ||||||||
China | 20 (142, 1995-98) | 0 (6) | 0 (8) | 0 (35) | |||||||
Colombia | 0 (40, 1994-2000) | ||||||||||
Uruguay | 100 (25, 1998-99) | ||||||||||
Morocco | 60 (108, 1997-2000) | 52 (25) | 42 (40) | ||||||||
Serbia | 76 (37) | ||||||||||
Mexico (Toluca) | 28 (292, 1988-89) | 50 (389, 1997-98) |
Tebur 10. Matsayin P. infestans damuwa tare da A2 nau'in mating a cikin ƙasashe daban-daban na duniya a tsakanin shekarun 2000 zuwa 2011
kasar | 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) | |||||||
Jamus | 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) | |||||
Ingila | 4 (47) | 10 (96) | 31 (55) | 55 (790) | 68 (862) | 70 (552) | 68 (299) | ||||
Finland | 47 (162) | 12 (218) | 42 | ||||||||
Faransa | 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 | |||||||
Arewa. Ireland | 0 (38) | 0 (58) | 0 (40) | 0 (24) | 5 (54) | 0 (18) | 27 (578) | 45 (239) | 36 (213) | 82 (60) | 10 (80) |
Netherlands | 66 (24) | 93 (15) | 91 (11) | ||||||||
Norway | 39 (328) | 3 (115) | 12 (19) | ||||||||
Peru | 0 (36) | ||||||||||
Poland | 25 (46) | 10 (30) | 85 (20) | 38 (44) | 75 (66) | 55 (56) | 65 (35) | 72 (81) | 85 (21) | ||
Scotland | 3 (213) | 2 (474) | 24 (135) | 86 (337) | 88 (386) | 74 (172) | |||||
Sweden | 60 (277) | 39 (87) | |||||||||
Slovakia | 0 (36) | 14 (26) | 62 (26) | 0 (26) | |||||||
Wales | 25 (12) | 68 (106) | 80 (88) | 92 (143) | 75 (45) | ||||||
Koriya | 46 (26) | ||||||||||
Brazil | 0 (49) | 0 (30) | |||||||||
China | 10 (30) | 0 (6) | 0 (6) | ||||||||
Vietnam | 0 (294, 2003-04) | ||||||||||
Yuganda | 0 (8) |
Dynamics of genotypic abun da ke ciki na yawan jama'a
Canje-canje a cikin tsarin halittar jini na P. yawan mutanen da ke addabar mutane na iya faruwa a ƙarƙashin tasirin ƙaurawar sabbin kwayoyi daga wasu yankuna, ayyukan noma (canjin iri, aikace-aikacen kayan gwari), da yanayin yanayi. Tasirin waje yana shafar kwayoyi daban-daban a matakai daban-daban na rayuwar rayuwa; sabili da haka, yawan jama'a kowace shekara suna fuskantar canje-canje na cyclical a cikin ƙananan ƙwayoyin halittar da aka zaɓa, saboda canji a cikin mahimmin rawar kwayar kwazo da zaɓi.
Tasirin iri-iri
Sabbin al'adu tare da ingantattun kwayoyin halitta don juriya ta tsaye (R-genes) babban zaɓi ne mai zaɓin zaɓi wanda ke zaɓar kwafin kwayoyi tare da ƙarin ƙwayoyin cuta na jijiyoyin jiki a cikin P. yawan jama'a. Idan babu juriya mara ma'ana a cikin nau'ikan dankalin turawa wanda ke hana ci gaban yawan kwayoyin cuta, aikin maye gurbin manyan kwayoyi masu yawa a cikin jama'a yana faruwa da sauri. Don haka, bayan yaduwa a cikin yankin Moscow na nau'ikan Domodedovsky, wanda ke da kwayar R3, yawan kwayoyi masu cutar da wannan iri sun karu daga 0,2 zuwa 0,82 a cikin shekara guda (Dyakov, Derevjagina, 2000).
Koyaya, canjin canjin yanayin kwayar halittar kwayar cuta (pathotypes) a cikin yawan jama'a yana faruwa ba kawai ƙarƙashin rinjayar nau'o'in dankalin turawa. Misali, a cikin Belarus har zuwa 1977, kwayoyi masu dauke da kwayar halitta ta 1 da 4 sun mamaye, wanda ya samo asali ne daga noman dankalin turawa tare da kwayar halittar R1 da R4 (Dorozhkin, Belskaya, 1979). Koyaya, a ƙarshen shekarun 70 na karni na 2002, kwayoyi sun bayyana tare da ƙwayoyin cuta daban-daban da haɗuwarsu, kuma ba a taɓa amfani da ƙwayoyin juriya masu haɗuwa a cikin kiwo da dankalin turawa (ƙarin ƙwayoyin cuta na ƙwayoyin cuta) (Ivanyuk et al., XNUMX). Dalilin bayyanar irin wadannan kwayoyi masu kunnen doki, ga alama, saboda ƙaura zuwa Turai na kayan kamuwa daga Mexico tare da tubers dankalin turawa. A cikin gida, waɗannan kwafin halittun sun haɓaka ba wai kawai a kan dankalin da aka noma ba, har ma a kan nau'ikan daji waɗanda ke ɗauke da ƙwayoyin halittar juriya da yawa; sabili da haka, haɗuwa da yawancin ƙwayoyin cuta a cikin kwayar halitta ya zama dole don rayuwa a waɗancan yanayi.
Game da nau'ikan da ke da juriya mara ma'ana, su, ta hanyar rage saurin yaduwar kwayar cuta, suna jinkirta canjin yawan jama'arta, wanda, kamar yadda aka ambata, aiki ne na lamba. Tunda tashin hankali polygenic ne, kwayoyi masu kunshe da yawancin kwayoyin halitta don "zafin rai" suna tarawa da sauri girman mutane. Sabili da haka, jinsi masu saurin tashin hankali ba samfuri ne na daidaitawa ga nau'ikan da aka noma tare da juriya mara ma'ana ba, amma, akasin haka, ana iya gano su a cikin dasa nau'ikan ƙwayoyi masu saukin kamuwa waɗanda sune masu tarawar ƙwayoyin cuta.
Don haka, a cikin Rasha an sami yawan mutanen da ke cikin tashin hankali na P. Infestans a cikin yankuna na shekara-shekara epiphytoties (yawan jama'a daga Sakhalin, Leningrad, da Bryansk yankuna). Tsananin tashin hankalin wadannan al'umman ya zama ya fi na mutanen Mexico (Filippov et al., 2004).
Bugu da kari, an samu karancin warkoki a cikin ganyen nau'ikan da ke jurewa fiye da wadanda ke iya kamuwa da cutar (Hanson da Shattock, 1998), ma'ana, rashin jituwa ta musamman da ke tattare da ire-iren wadannan kuma yana rage karfin sake kamuwa da kwayoyin cutar da yiwuwar wasu hanyoyin na hunturu.
Tasirin kayan gwari
Fungicides ba kawai rage yawan fungi na phytopathogenic ba, watau shafi halaye na adadi na yawan mutanen su, amma kuma suna iya canza mitar yanayin halittar mutum, watau tasiri tasirin ƙididdigar yawan jama'a. Daga cikin mahimman alamu na yawan mutanen da ke canzawa a ƙarƙashin tasirin kayan gwari sune masu zuwa: canje-canje a cikin juriya da magungunan gwari, sauye-sauye cikin zafin rai da lalata jiki, da canje-canje a tsarin haihuwa.
Tasirin kayan gwari akan juriya da ta'adi na yawan jama'a
Matsayin wannan tasirin ya ƙaddara, da farko, ta hanyar nau'in kayan gwari da ake amfani da shi, wanda za'a iya rarraba shi cikin yanayi zuwa polysite, oligosite da monosite.
Na farkon ya hada da yawancin masu amfani da kayan gwari. Juriya a gare su (idan yana yiwuwa a kowane) ana sarrafa shi ta hanyar adadi mai yawa na kwayoyin halitta masu rauni sosai. Waɗannan kaddarorin suna ƙayyade rashin canje-canje na bayyane a cikin juriya na jama'a bayan jiyya tare da kayan gwari (kodayake a cikin wasu gwaje-gwajen an sami ƙarin ƙaruwa a juriya). Yawan naman gwari da aka adana bayan an fesa shi tare da magungunan kashe gwari ya kunshi kungiyoyi biyu na damuwa:
1) Hanyoyin da aka adana a wuraren tsire-tsire waɗanda ba a magance su da magani. Tunda babu wata hulɗa da kayan gwari, ta'adi da juriya na waɗannan matsalolin ba ya canzawa.
2) insarfafawa a cikin hulɗa tare da kayan gwari, ƙwarewar da yake a wuraren tuntuɓar ta ƙasa da ta mutuwa. Kamar yadda aka ambata a sama, juriyar wannan ɓangaren ma ba ta canzawa ba, duk da haka, saboda lahanin da ake samu na kayan gwari ko da kuwa a cikin kwayar halitta ne kan kwayar halittar fungal, ƙoshin lafiyar gaba ɗaya da ɓangarenta na parasitic, tashin hankali, raguwa (Derevyagina da Dyakov, 1990).
Don haka, koda wani ɓangare na yawan mutanen da basu mutu ba, wanda aka fallasa su tuntuɓar mai amfani da kayan gwari, yana da rauni mai rauni kuma ba zai iya zama tushen epiphytotics ba. Sabili da haka, kula da hankali wanda ke rage yawan adadin yawan mutanen da basa hulɗa da kayan gwari yanayi ne na nasarar matakan kariya. Rashin jituwa da oligosite kayan gwari yana sarrafawa ta yawancin kwayoyin halitta.
Maye gurbi na kowace kwayar halitta yana haifar da daɗaɗa juriya, kuma gabaɗaya matakin juriya saboda ƙari da irin wannan maye gurbi ne. Sabili da haka, ƙaruwa cikin juriya yana faruwa ne kai tsaye. Misali na ƙaruwa a cikin tsayayyar gaba ita ce maye gurbi a juriya da dimethomorph na fungicide, wanda ake amfani da shi sosai don kare dankali daga ƙarshen cutar. Dimethomorph juriya ne polygenic da ƙari. Sauyawa mataki-mataki sau ɗaya kaɗan yana ƙaruwa da ƙarfi.
Kowace maye gurbi na gaba yana rage girman makirci kuma, sakamakon haka, yawan maye gurbi na gaba (Bagirova et al., 2001). Inara yawan matsakaicin juriya na jama'a bayan maimaita jiyya tare da fungicide na oligosite yana faruwa a hankali kuma a hankali. Saurin wannan tsari an ƙaddara shi aƙalla aƙalla abubuwa guda uku: yawan maye gurbi na ƙwayoyin juriya, gwargwadon ƙarfin juriya (rabon kashi na mutuwa mai saurin haɗari dangane da mai damuwa) da kuma sakamakon maye gurbi a cikin kwayoyin juriya akan dacewa.
Yawan faruwar kowane maye gurbi yana zuwa ƙasa da na baya; sabili da haka, aikin yana da ɗabi'a mai lalacewa (Bagirova et al., 2001). Koyaya, idan hanyoyin sake haɗawa (jima'i ko ɗan luwaɗi) suka faru a cikin yawan jama'a, to yana yiwuwa a haɗu da maye gurbi daban-daban na iyaye a cikin wani nau'in haɗari da kuma hanzarta aikin. Saboda haka, al'ummomin panmix suna samun juriya da sauri fiye da na agamic, kuma a karshen, yawan mutanen da basu da shingen rashin daidaituwa na ciyayi da sauri fiye da yawan mutanen da aka raba ta irin waɗannan shingen. Dangane da wannan, kasancewar damuwa a cikin al'ummomin da suka banbanta a cikin nau'ikan saduwa da juna yana hanzarta aiwatar da samin juriya ga kayan gwari oligosite.
Abubuwa na biyu da na uku ba sa ba da gudummawa ga saurin tarin ƙwayoyin dimethomorph masu juriya a cikin jama'a. Kowace maye gurbi na zuwa kusan ninki biyu na juriya, wanda ba shi da mahimmanci, kuma a lokaci guda yana rage yawan ci gaban da ke cikin yanayin wucin gadi da tashin hankali (Bagirova et al., 2001; Stem, Kirk, 2004). Wataƙila wannan shine dalilin da ya sa kusan babu wata damuwa mai tsayayya tsakanin ƙwayoyin halittu na P., har ma waɗanda aka tattara daga dasa dankalin turawa da aka kula da su tare da dimethomorph.
Jama'ar da aka yiwa magani tare da fungicide na oligosite zasu hada da rukuni biyu na damuwa: wadanda basu hadu da maganin ba, sabili da haka basu canza halaye na farko ba (idan ana samun damuwa masu karfi a tsakanin wannan rukunin, ba zasu tara ba saboda tsananin tashin hankali da kuma gasa masu wahala) da kuma damuwa a cikin hulɗa tare da abubuwan da ake amfani da su na kayan gwari. Yana daga cikin ƙarshen cewa tarin ƙwayoyin cuta masu juriya abu ne mai yiwuwa, saboda a nan suna da fa'idodi akan waɗanda ke da laushi.
Sabili da haka, lokacin amfani da kayan gwari na oligosite, ba cikakken magani bane wanda ke da mahimmanci azaman mahimmin maganin ƙwayoyi, sau da yawa ya fi na kisa, saboda tare da matakan mutagenesis, farkon juriya na yanayin maye gurbi yana da ƙasa.
Aƙarshe, maye gurbi a juriya da magungunan kashe kayan masarufi yana da ma'ana sosai, ma'ana, maye gurbi ɗaya na iya bayar da rahoton babban matakin juriya, har zuwa ƙarshen hasarar hankali. Sabili da haka, ƙaruwa cikin juriya na yawan jama'a yana faruwa da sauri sosai.
Misalin irin wadannan kayan gwari sune phenylamides, gami da mafi yawan kayan gwari, metalaxyl. Maye gurbi na juriya da shi ya tashi tare da babban mitar, kuma matakin juriya a cikin maye gurbi yana da girma ƙwarai - ya wuce saurin damuwa da kusan dubu ko fiye (Derevyagina et al., 1993). Kodayake yawan ci gaba da tashin hankali na rikirkita rikirkicewa yana raguwa dangane da asalin mutuwar nau'in mai saukin kamuwa daga kayan gwari, amma yawan mutanen da ke juriya suna girma cikin sauri kuma, a layi daya, ta'addancin nasa yana karuwa. Saboda haka, bayan shekaru da yawa na amfani da kayan gwari, tsananin tashin hankali na damuwa ba zai iya daidaita zaluncin masu damuwa ba, har ma ya zarce shi (Derevyagina da Dyakov, 1992).
Tasiri kan sake haɗuwa da jima'i
Tunda yawan abin da ke faruwa a A2 a cikin P. yawan jama'a ya yi daidai da tsananin amfani da metalaxyl akan ƙarshen ɓarayi, an ba da shawarar cewa metalaxyl yana haifar da sauya nau'in ma'aurata. A cikin P. parasitica, irin wannan jujjuyawar a ƙarƙashin aikin chloroneb da metalaxyl an gwada ta da gwaji (Ko, 1994). Hanya guda daya akan matsakaici tare da karancin sinadarin metalaxyl ya haifar da bayyanar rabewar homothallic daga wani nau'in P. wadanda suka kamu da cutar metalaxyl tare da nau'in A1 (Savenkova da Cherepnikova-Anikina, 2002). Yayin da wasu hanyoyin da zasu biyo baya a kafafen yada labarai tare da karfin metalaxyl, ba a gano wani kebantaccen nau'in A2 wanda yake hade da juna ba, duk da haka, mafi yawan masu kebewa, lokacin da aka haye su tare da A2, maimakon oospores, sun kirkiro tarin kwayoyin mycelium kuma basu da amfani. Hanyoyin da ke tattare da nau'in nau'in nau'in nau'in A2 akan kafofin watsa labaru tare da babban ƙwayar metalaxyl ya ba mu damar gano nau'ikan nau'ikan canje-canje iri uku: 1) cikakkiyar haihuwa lokacin da aka haye tare da A1 da A2 keɓewa; 2) homotallism (samuwar abubuwan fida a cikin wata al'ada); 3) jujjuyawar nau'ikan A2 na ma'aurata zuwa A1. Don haka, metalaxyl na iya haifar da canje-canje a cikin nau'ikan saduwa a cikin mazaunan P. kuma saboda haka, faruwar sake haɗuwa da jima'i a cikinsu.
Tasiri kan sake haɗuwa da ciyayi
Wasu kwayoyin halittar juriya sun kara yawan karfin karfin jini da difloma na nukiliya (Poedinok da Dyakov, 1981). Kamar yadda muka gani a baya, heterokaryotization na hyphae yayin haɗuwa da nau'ikan iri daban-daban na P. infestans yana faruwa sosai da wuya saboda abin da ya faru na rashin daidaituwa da ciyayi a cikin wannan naman gwari. Koyaya, kwayoyin don jure wasu maganin rigakafi na iya haifar da illa, wanda aka bayyana kan shawo kan rashin dacewar ciyayi. An mallaki wannan kayan ta hanyar maye gurbin 1S-1 mutant streptomycin gene gene. Kasancewar irin wadannan mutant a cikin yawan mutanen da ke cikin halittar jiki na iya kara yaduwar kwayoyin halittu tsakanin kwayoyin halitta da kuma hanzarta saurin karbuwa ga dukkan mutane zuwa sabbin iri ko kayan gwari.
Wasu masu amfani da kayan gwari da magungunan kashe kwayoyin cuta na iya yin tasiri kan yawan sake hadewar jiki, wanda kuma zai iya sauya mitocin jinsin mutane. Benomyl mai yaduwa da ake amfani dashi yana ɗaure ne zuwa beta-tubulin, furotin wanda ake gina microtubules na cytoskeleton, kuma hakan yana dagula ayyukan rabuwa na chromosome a cikin anaphase na mitosis, yana ƙaruwa da saurin sake haɗawar mitotic (Hastie, 1970).
Magungunan fungicide para-fluorophenylalanine, da ake amfani da shi don magance cutar Dutch a cikin tsautsayi, yana da dukiya iri ɗaya. Para-fluorophenylalanine ya haɓaka yawan sake haɗawa a cikin heterozygous diploids P. infestans (Poedinok et al., 1982).
Canje-canje mai canzawa a cikin jigon halittar mutane na rayuwar rayuwar P. infestans
Tsarin rayuwar yau da kullun na P. infestans a cikin yankin mai sanyin yanayi ya ƙunshi fasali 4.
1) Matsakaicin haɓaka yawan jama'a (polycyclic phase) tare da gajerun ƙarni. Wannan lokaci yakan fara ne a watan Yuli kuma yana ɗaukar watanni 1,5-2.
2) Lokaci na dakatar da karuwar jama'a saboda raguwar kaɗan na adadin kayan da ba a taɓa su ba ko farkon yanayin rashin kyau. Wannan matakin a gonakin da ke aiwatar da cire ganyen farko kafin girbi ya sauka daga zagayowar shekara-shekara.
3) Lokaci na hunturu a tubers, tare da raguwa mai yawa a cikin yawan mutane saboda haɗarin kamuwa da bazara, jinkirin ɓullar kamuwa da cuta a cikinsu, rashin sake kamuwa da cutar tubers, ruɓewa da cushe tubers da abin ya shafa a ƙarƙashin yanayin ajiya na yau da kullun.
4) Lokaci na jinkirin cigaba a cikin ƙasa da kan seedlings (monocyclic phase), wanda tsawon lokacin ƙarni zai iya kaiwa wata ɗaya ko fiye (ƙarshen Mayu - farkon Yuli). Yawancin lokaci a wannan lokacin, ganyayyaki marasa lafiya suna da wuyar ganewa, koda kuwa da lura na musamman.
Lokaci na haɓaka yawan mutane (polycyclic phase)
Abubuwan lura da yawa (Pshedetskaya, Kozubova, 1969; Borisenok, 1969; Osh, 1969; Dyakov, Suprun, 1984; Rybakova, Dyakov, 1990) sun nuna cewa a farkon epiphytoty, ƙananan ƙwayoyin cuta masu ƙarancin ƙarfi da ƙarancin ƙarfi, waɗanda aka maye gurbinsu da waɗanda suka fi ƙarfin da ƙarfi. yawan karuwar tashin hankali na jama'a shine mafi girma, mafi ƙarancin juriya da nau'ikan tsire-tsire masu masaukin baki.
Yayinda yawan jama'a ke ƙaruwa, haɓakar halittu masu mahimmanci masu mahimmanci waɗanda aka gabatar dasu cikin nau'ikan kasuwanci (R1-R4) da tsaka tsaki zaɓaɓɓe (R5-R11) yana ƙaruwa. Don haka, a cikin yawan mutanen kusa da Moscow a cikin 1993, matsakaicin tashin hankali daga ƙarshen Yuli zuwa tsakiyar Agusta ya karu daga 8,2 zuwa 9,4, kuma mafi girman haɓaka an lura da kwayar cutar kwayar cuta ta R5 (daga 31 zuwa 86% na kwayoyi masu illa) (Smirnov, 1996) ).
Rage yawan ci gaban yawan mutane yana tare da raguwar ayyukan parasitic na yawan jama'a. Sabili da haka, a cikin shekarun damuwa, duka yawan jinsi da kuma yawan jinsi masu rauni sun fi na epiphytotic (Borisenok, 1969). Idan a tsayin yanayi na yanayin yanayin epiphytotic ya canza zuwa mara kyau don jinkirin bugun ƙarewa da ƙarancin dankalin turawa, ƙarancin kwayoyi masu saurin tashin hankali da raguwa suma suna raguwa (Rybakova et al., 1987).
Inara yawan ƙwayoyin halittar da ke shafar larura da tashin hankali na yawan jama'a na iya zama saboda zaɓin ƙarin kwayoyi masu banƙyama da haɗari a cikin cakudaddiyar jama'a. Don nuna zaɓin, an kirkiro wata hanya don nazarin maye gurbin tsaka-tsakin yanayi, wanda aka yi amfani da shi cikin nasara a yawancin mutanen yisti (Adams et al., 1985) da Fusarium graminearum (Wiebe et al., 1995).
Yawan canzawar yanayin mutants wanda ya yi hannun riga da blasticidin S a cikin yawan mutanen P. infestans ya ragu daidai yake da ci gaba a cikin zafin rai na yawan jama'a, wanda ke nuna canji a cikin manyan kwayoyi yayin haɓakar jama'a (Rybakova et al., 1987).
Lokacin cin nasara a cikin tubers
A lokacin hunturu a cikin tubers dankalin turawa, tsananin karfi da zafin rai na P. infestans damuwa yana raguwa, kuma raguwar cutar na faruwa a hankali fiye da tashin hankali (Rybakova da Dyakov, 1990). A bayyane yake, a ƙarƙashin yanayin da ya dace da saurin ƙaruwar jama'a (zaɓin r), "ƙarin" ƙwayoyin cuta da tsananin tashin hankali suna da amfani, sabili da haka ci gaban epiphytotics yana tare da zaɓin mafi mawuyacin kwayoyi masu zafi da ƙarfi. A cikin yanayin jikewar muhalli, lokacin da ba yawan haihuwa ba, amma juriya da wanzuwa a cikin yanayi mara kyau (K-zaɓi) yana taka muhimmiyar rawa, "ƙarin" kwayoyin halittar ƙwayoyin cuta da tashin hankali suna rage ƙoshin lafiya, kuma kwayoyi masu ɗauke da waɗannan ƙwayoyin halitta sune farkon waɗanda zasu mutu, don haka matsakaiciyar tashin hankali da yawan kwayar cutar na raguwa.
Lokaci na ciyayi a cikin ƙasa
Wannan lokaci shine mafi ban mamaki a cikin rayuwar rayuwa (Andrivon, 1995). Kasancewarsa an tsara shi ne kawai - saboda karancin bayani game da abin da ke faruwa ga mai cutar tsawon lokaci (wani lokaci fiye da wata daya) - daga fitowar dankalin turawa zuwa bayyanar alamun farko na cutar a kansu. Dangane da abubuwan lura da gwaje-gwajen, an sake gina halayen fungus a wannan zamanin na rayuwa (Hirst da Stedman, 1960; Boguslavskaya, Filippov, 1976).
Sporulation na naman gwari na iya samuwa kan tubers mai dauke da cutar a cikin ƙasa. Sakamakon spores yayi girma tare da hyphae, wanda zai iya ciyawa na dogon lokaci a cikin ƙasa. Firamare (wanda aka kafa akan tubers) da sakandare (akan mycelium a cikin ƙasa) spores ya tashi zuwa farfajiyar ƙasa ta hanyar igiyar ruwa, amma ya sami ikon kamuwa da dankali kawai bayan leavesan ganyensa ya sauko kuma ya sadu da farfajiyar ƙasa. Irin waɗannan ganyayyaki (wato, alamun farko na cutar ana samun su) basu cika nan da nan ba, amma bayan ci gaba mai ɗaci da ci gaban saman dankalin turawa.
Don haka, tsarin ciyawar saprotrophic shima yana iya kasancewa a cikin rayuwar rayuwar P. infestans. Idan a cikin yanayin yanayin rayuwar rayuwa tashin hankali shine mafi mahimmancin yanayin motsa jiki, to a cikin zaɓin saprotrophic ana nufin rage abubuwan da ke cikin parasitic, kamar yadda aka nuna a gwaji don wasu fungi na phytopathogenic (duba Carson, 1993). Sabili da haka, a wannan lokacin na sake zagayowar, yakamata a rasa dukiyoyi masu ƙarfi sosai. Amma har yanzu ba a gudanar da gwaje-gwajen kai tsaye ba don tabbatar da zaton da ke sama ba.
Sauye-sauye na yanayi ba wai kawai tasirin ƙwayoyin cuta na P. infestans ba, har ma da juriya ga kayan gwari, wanda ke tsiro a cikin kwayar polycyclic (yayin epiphytoties), kuma yana raguwa yayin ajiyar hunturu (Derevyagina et al., 1991; Kadish da Cohen, 1992). An lura da raguwar tsayayyar metalaxyl tsakanin dasa shukokin da abin ya shafa da bayyanar alamun farko na cutar a filin.
Ingantaccen fasaha da sauyi
P. infestans yana haifar da annoba a cikin amfanin gona biyu masu muhimmanci, dankali da tumatir. Epiphytoties akan dankali ya fara jim kaɗan bayan naman gwari ya shiga sabbin yankuna. Hakanan an lura da kayar da tumatir jim kaɗan bayan bayyanar kamuwa da cuta a kan dankali, amma ba a lura da epiphytoties a kan tumatir sai bayan ɗari bayan haka - a tsakiyar ƙarni na XNUMX. Ga abin da Hallegli da Niederhauser suka rubuta game da shan tumatir a cikin Amurka
(1962): “Kimanin shekaru 100 bayan mummunan cutar cututtukan fata na 1845, kalilan ne ko kusan ba a yi ƙoƙarin samo irin tumatir mai jurewa ba. Kodayake an fara narkar da cutar ta tumatir a tumatir tun daga shekarar 1848, amma hakan bai zama abin kulawa ga masu kiwo a wannan shuka ba har sai da cutar ta yi karfi sosai a shekarar 1946. A cikin ƙasar Rasha an yiwa tumatir ƙarshen bugun tumatir a ƙarni na 60. “Tsawon lokaci, masu bincike ba su mai da hankali kan wannan cutar ba, tunda ba ta haifar da babbar illa ga tattalin arziki ba. Amma a cikin 70s da 1979s. Hakanan ana lura da epiphytoties na ƙarshen ƙarni na kamu da tumatir a Soviet Union, galibi a yankin Lower Volga, a cikin Ukraine, North Caucasus, a Moldova ... ”(Balashova, XNUMX).
Tun daga wannan lokacin, cutar tumatir ta ƙarshen ɓarayi ta zama ta shekara-shekara, ta bazu a duk faɗin ƙasar masana'antu da noman gida kuma yana haifar da babbar illa ga tattalin arzikin wannan amfanin gona. Me ya faru? Me yasa bayyanar cutar ta farko akan dankali da cututtukan epiphytotic na wannan amfanin gona ya faru kusan lokaci guda, kuma me yasa ya dauki karni kafin epiphytotic ya bayyana akan tumatir? Wadannan bambance-bambance suna tallafawa dan Mexico maimakon tushen kamuwa da Kudancin Amurka. Idan jinsunan Phytophthora infestans suka zama kwayoyin parasicic na mexico masu dauke da tuber na jinsi na Solanum, to zamu iya fahimtar dalilin da yasa dankalin da aka kebanta shi yake daya daga cikin jinsin halittar kamar yadda jinsin Mexico yake da karfi sosai, amma saboda rashin cigaban juyin halitta tare da kwayar cutar, wacce ba ta samar da wasu dabaru na musamman ba.
Tumatir na wani bangare ne daban na jinsi, nau'in musayarsa yana da manyan bambance-bambance daga nau'ikan nau'ikan bututu, saboda haka, duk da cewa tumatir baya wajen ƙwarewar abinci na P. infestans, ƙarfin lalacewar sa bai isa ba ga asarar tattalin arziki mai tsanani.
Bayyanar cututtukan epiphytoties akan tumatir saboda tsananin canjin kwayoyin halitta a cikin kwayar cutar, wanda ya kara karfin gwiwa (pathogenicity) yayin cutar ta parasitism. Mun yi imanin cewa sabon salon da aka keɓe don inganta tumatir shine tseren T1 wanda M. Gallegly ya bayyana, wanda ke shafar nau'ikan tumatir na tumatir (Red Cherry, Ottawa), mai jure tseren T0 da ya bazu a kan dankali (Gallegly, 1952). A bayyane yake, maye gurbi (ko jerin maye gurbi) wanda ya juya tseren T0 zuwa tseren T1 kuma ya haifar da bayyanar kwayayen kwayoyi masu dacewa sosai don kayar da tumatir. Kamar yadda yake faruwa sau da yawa, ƙaruwa a cikin cuta ga mai gida ɗaya yana tare da raguwa a ciki zuwa wani, ma'ana, na farko, ba tukuna cikakke ƙwarewar ƙwarewa ya tashi - zuwa dankali (tseren T0) da tumatir (tseren T1).
Menene hujja ga wannan zato?
- Faruwa akan dankali da tumatir. Akan ganyen tumatir, tseren T1 ya mamaye, yayin da akan ganyen dankalin turawa yake da wuya. A cewar S.F.Bagirova da T.A. Oreshonkova (ba a buga shi ba) a cikin yankin Moscow a cikin 1991-1992, faruwar tseren T1 a cikin shukar dankalin turawa ya kai kashi 0%, kuma a cikin tumatir - 100%; a 1993-1995 - 33% da 90%, bi da bi; a 2001 - 0% da 67%. An samo irin wannan bayanan a cikin Isra'ila (Cohen, 2002). Gwaje-gwaje game da kamuwa da cutar tumatir dankalin turawa tare da keɓaɓɓiyar jinsin T1 da cakuda masu keɓewa T0 da T1 sun nuna cewa keɓancewar T1 tsere ba a kiyaye shi sosai a cikin tubers kuma an maye gurbinsu da keɓewar T0 (Dyakov et al., 1975; Rybakova, 1988).
2) Dynamics of the T1 race a cikin dasa tumatir. Cutar farko ta ganyen tumatir ana aiwatar da ita ne ta hanyar keɓancewar jinsin T0, wanda ya mamaye yin nazarin kamuwa da cuta a wuraren farko da aka kafa akan ganyen. Wannan yana tabbatar da makircin da aka yarda dashi na ƙaura mai cutar: Babban kamuwa da cuta daga dankali ana yin sa ne ta tseren T0, amma, ƙananan ƙwayoyi na T1 da aka adana a cikin dankali, sau ɗaya akan tumatir, sun sauya tseren T0 kuma sun taru zuwa ƙarshen lokacin epiphytotic. Zai yiwu kuma akwai wani madadin tushen kamuwa da ganyen tumatir tare da jinsin T1, wanda ba shi da ƙarfi kamar tubers dankali da ganye, amma na yau da kullun. Saboda haka, wannan tushe yana da rauni mai tasiri akan tsarin kwayar halittar jama'ar da ke kamuwa da tumatir, amma daga baya yana tantance tarin jinsin T1 (Rybakova, 1988; Dyakov et al., 1994).
3) Tsanani ga dankali da tumatir. Kamuwa da cutar tumatir da ganyen dankalin turawa tare da keɓaɓɓun jinsunan T0 da T1 ya nuna cewa na farko sun fi nuna damuwa ga dankali fiye da na tumatir, kuma na biyun sun fi tumatir ƙarfi fiye da na dankalin turawa. Wadannan bambance-bambance sun bayyana a cikin kawar da keɓewar jinsin da ba “mallakarsa” ba daga yawan mutanen da aka gauraya a yayin da ganyen ganye a cikin gidan haya (D'yakov et al., 1975) da kuma cikin filayen filaye (Leberton et al., 1999); bambance-bambance a cikin mafi ƙarancin ƙwayar cuta, lokacin latency, girman wuraren cuta da samar da ƙwayoyin cuta (Rybakova, 1988; Dyakov et al., 1994; Legard et al., 1995; Forbes et al., 1997; Oyarzun et al., 1998; Leberton et al. al., 1999; Vega-Sanchez et al., 2000; Knapova, Gisi, 2002; Sussuna et al., 2004).
Tsananin tashin hankali na tseren T1 zuwa nau'ikan tumatir da basu da kwayar halittar juriya yana da yawa har waɗannan rabe-raben suna zube a kan ganyayyaki kamar kan matsakaitan abinci mai gina jiki ba tare da lalata ƙwayar cuta ba (Dyakov et al., 1975; Vega-Sanchez et al., 2000).
4) Nutsuwa ga dankali da tumatir. Gasar T1 tana shafar nau'ikan tumatir na ceri tare da kwayar juriya ta Ph1, yayin da tseren T0 ba zai iya kamuwa da waɗannan nau'in ba, watau yana da ƙananan ƙwayar cuta. Dangane da masu rarrabewa
R-genes din dankalin yana da alaka da juna, watau nau'ikan da ke ware daga ganyen tumatir ba su da ƙarfi kamar ƙwayoyin "dankalin turawa" (Table 11).
5) Alamun tsakani. Nazarin alamomin tsaka tsaki a cikin yawan mutanen P. infestans parasitizing akan dankali da tumatir shima ya tabbatar da zaɓin intraspecific multidirectional. A cikin jama'ar Brazil na P. infestans, ganyen tumatir ya zama na layin ne US-1, kuma waɗanda ke ganyen dankalin turawa na layin BR-1 ne (Suassuna et al., 2004). A cikin Florida (Amurka), tun daga 1994, clone US-90 ya fara mamaye kan dankali (tare da faruwar sama da 8%), kuma kwafin US-11 da US-17 akan tumatir, kuma keɓewar na ƙarshen ya fi zalunci ga tumatir fiye da dankalin turawa (Weingartner , Tombolato, 2004). Mahimmancin bambance-bambance a cikin nau'ikan halittar jini (zanan yatsun DNA) a cikin dankalin turawa da tumatir an keɓe shi don 1200 P. ɓarna da aka tattara a Amurka daga 1989 zuwa 1995 (Deahl et al., 1995).
Yin amfani da hanyar AFLP ya ba da damar raba iri 74 da aka tattara daga dankalin turawa da ganyen tumatir a cikin 1996-1997. a Faransa da Switzerland, a rukuni 7. Nauyin dankalin da na tumatir bai banbanta sosai ba, amma nau'ikan "dankalin" ya sha bamban da na "tumatir". An samo tsohuwar a cikin dukkanin gungu bakwai, kuma na biyun, kawai a cikin huɗu, wanda ke nuna ƙirar ƙirar ƙwararriyar ta ƙarshen (Knapova da Gisi, 2002).
6) Hanyoyin keɓewa. Idan yawan kwayoyin halittar dake jikin wasu jinsin shuke-shuke biyu suka canza zuwa ga takaita kwarewar zuwa ga mai gidan nasu, to wasu dabaru na gaba da na baya-baya sun tashi wadanda zasu hana musayar kwayoyin halittar juna (Dyakov da Lekomtseva, 1984).
Yawancin karatu sun binciko tasirin asalin damuwa na iyaye game da ingancin haɓakar haihuwa. Lokacin da ake tsallake wasu nau'ikan da ke ware daga jinsuna daban-daban na Solanum a Ecuador (Oliva et al., 2002), an gano cewa damuwa da nau'ikan A2 daga Solanaceae na daji (layin layi EC-2) sune mafi munin ketare da damuwa daga tumatir (layin EC -3), kuma mafi inganci ya ketare tare da dankalin turawa (EC-1).
Dukkanin matasan da aka samu basu da wata cuta. Marubutan sun yi imanin cewa ƙananan kashi na haɓakawa da raguwar ƙwayoyin cuta a cikin haɗuwa sun kasance ne saboda hanyoyin aikin gaba na keɓance haihuwar jama'a.
A cikin gwaje-gwajen Bagirova et al. (1998), an haye yawancin dankalin turawa da tumatir tare da kaddarorin jinsunan T0 da T1. Wadanda ke da matukar inganci sune giciye na nau'ikan T1xT1 wadanda aka kebe daga tumatir (36 oospore a cikin microscope fagen gani, 44% na germination oospore), mafi karancin tasiri shine tsallakar jinsi na T0xT1 wanda aka kebe daga wasu runduna daban-daban (wani adadi mai yawa na ci gaba da tsire-tsire masu tsire-tsire) ... Ingancin gicciye tsakanin keɓancewar tseren T0 wanda aka ware daga dankali ya kasance tsaka-tsaki. Tunda babban nau'in damuwa na tseren T0 yana shafar dankali, yana da tushen abin dogaro na damuna - tubers dankalin turawa, sakamakon haka mahimmancin sifa kamar ƙanƙantar da keɓaɓɓu ga ƙananan mutane daga dankali ya yi ƙasa. Daidaitaccen “nau'in tumatir” yana iya yin hunturu a kan tumatir a matsayin oospores (duba ƙasa) sabili da haka yana riƙe da haɓaka mafi girma na tsarin jima'i. Saboda yawan haihuwa, T1 ta sami damar cin gashin kansa na kamuwa da cutar tumatir. Sakamakon da Knapova et al. (Knapova et al., 2002) ana iya fassara su ta hanya ɗaya. Maganganun matsalolin da aka ware daga dankali tare da damuwa daga tumatir sun ba da mafi yawan adadin fure - 13,8 a kowace sq.mm. matsakaici (tare da yaduwar 5-19) da kuma matsakaiciyar kashi na yaduwar oospores (6,3 tare da yaduwar 0-24). Rage igiyar da aka keɓe daga tumatir ya ba da kashi mafi ƙasƙanci na magogin ruwa (7,6 tare da yaɗuwar 4-12) tare da mafi yawan ƙwayoyinsu (10,8). Giciye tsakanin matsalolin da aka keɓe daga dankali ya ba da matsakaiciyar adadin furofesoshi (8,6 tare da babban watsa bayanai - 0-30) da mafi ƙarancin kashi na ƙwayoyin cuta na oospores (2,7). Don haka, damuwa daga dankali ba shi da wadatuwa fiye da na tumatir, amma gicciyen mutane bai ba da sakamakon da ya fi na mutanen ciki ba. Zai yiwu cewa bambance-bambance tare da bayanan da ke sama na Bagirova et al. an bayyana ta gaskiyar cewa masu binciken na Rasha sun yi aiki tare da ɓacin rai da aka keɓe a farkon shekarun 90 na ƙarni na ashirin, kuma masu binciken Switzerland - tare da ɓacin rai da aka ware a ƙarshen 90s.
Dalili don ƙarancin haihuwa na iya zama heteroploidy na damuwa. Idan a cikin jama'ar Meziko, inda tsarin jima'i da kamuwa da cuta ta farko tare da zuriya mai cike da ɗabi'a na yau da kullun ne, yawancin yawancin binciken na P. Infestans na difloma ne, to a cikin ƙasashe na Tsohon Worldwayar intrapopulation polymorphism na ploidy ana lura da (nau'in di-, tri- da tetraploid, da kuma cututtukan heterokaryotic tare da mahaifa heteroploid) , da kuma nau'ikan nau'ikan nau'ikan hanyoyin saduwa, watau masu amfani da juna, sun banbanta a shirin nukiliya (Therrien et al., 1989, 1990; Whittaker et al., 1992; Ritch, Daggett, 1995). Bambancin nuclei a cikin antheridia da oogonia na iya zama dalilin ƙarancin haihuwa.
Game da musayar makaman nukiliya tsakanin hyphae a lokacin anastomoses, wannan ya hana ta rashin daidaituwa tsakanin ciyayi, wanda ke raba yawan mutane masu zuwa a cikin kwayoyi masu yawa (Poedinok da Dyakov, 1987; Gorbunova et al., 1989; Anikina et al., 1997b).
7) Yawan jama'a. Bayanai na sama sun nuna cewa hadewa tsakanin "dankalin turawa" da "tumatir" P. 'yan infestans iri ne mai yiwuwa. Sake kamuwa da sake kamuwa da cuta na runduna daban-daban shima yana yiwuwa, kodayake yana rage yawan tashin hankali.
Nazarin alamomin yawan jama'a a keɓe daga gonakin dankalin turawa da tumatir a shekara ta 1993 ya nuna cewa kimanin kashi ɗaya cikin huɗu na keɓewar daga ganyen tumatir an ɗauke su daga filin dankalin da ke makwabtaka da su (Dolgova et al., 1997). A ka'ida, ana iya ɗauka cewa bambancin yawan jama'a akan runduna biyu zai haɓaka kuma zai haifar da fitowar wasu nau'ikan siffofin ƙwararru (f.sp. dankalin turawa da tumatir f.sp.), musamman tun da oospores na iya ci gaba cikin tarkacen shuka (Drenth et al., 1995 ; Bagirova, Dyakov, 1998) da kuma tumatir (Rubin et al., 2001). Sakamakon haka, tumatir a halin yanzu suna da tushen sabunta bazara mai zaman kansa daga tubers dankalin turawa.
Koyaya, komai ya faru daban. Winarfafawa tare da oospores ya ba da damar parasite ɗin don guje wa mataccen matse a cikin rayuwarsa - matakin monocyclic na ciyayi a cikin ƙasa, a lokacin da kayan masarufin ke raguwa, waɗanda a hankali ake mayar da su cikin yanayin polycyclic a lokacin bazara.
Tebur 11. Mitar yanayin kwayar halitta zuwa nau'ikan bambance-bambancen dankalin Turawa a cikin ƙwayoyin cuta na P.
kasar | Shekara | Matsakaicin adadin ƙwayoyin cuta na jijiyoyi a cikin damuwa | marubucin | |
daga dankali | daga tumatir | |||
Faransa | 1995 | 4.4 | 3.3 | Leberton et al., 1999 |
1996 | 4.8 | 3.6 | Leberton, Andrivon, 1998 | |
Faransa, Switzerland | 1996-97 | 6.8 | 2.9 | Knapova, Gisi, 2002 |
United States | 1989-94 | 5 | 4.8 | Goodwin et al., 1995 |
Amurka, Zap. Washington | 1996 | 4.6 | 5 | Dorrance et al., 1999 |
1997 | 6.3 | 3.5 | " | |
Ecuador | 1993-95 | 7.1 | 1.3 | Oyarzun et al., 1998 |
Isra'ila | 1998 | 7 | 4.8 | Cohen, 2002 |
1999 | 6 | 5.7 | " | |
2000 | 6.7 | 6.1 | " | |
Rasha, Mosk. yanki | 1993 | 8.9 | 6.7 | Smirnov, 1996 |
Rasha, yankuna daban-daban | 1995 | 9.4 | 8 | Kozlovskaya da sauransu. |
1997 | 9.2 | 9.2 | " | |
2000 | 8.7 | 4.8 | " |
Firayim gidan zoosporangia da zoospores, wanda ke fitar da ospores, suna da babban aikin kwayar cutar, musamman idan aka samar da kuliyoyin da zaran an sami sauyin yanayi a karkashin tasirin kwayoyin halittar wani irin yanayi tare da kishiyar nau'in saduwa. Sabili da haka, kayan cututtukan akan tumatir da aka tsiro daga ƙwayoyin da suka kamu da oospores suna da matukar damuwa ga tumatir da dankalin turawa.
Waɗannan canje-canje sun haifar da sake fasalin yawan jama'a, wanda aka bayyana a cikin mahimman canje-canje masu zuwa daga ra'ayi na annoba:
- Maganin tumatir da ya kamu ya zama muhimmiyar hanyar kamuwa da dankali na farko (Filippov, Ivanyuk, saƙonnin sirri).
- Epiphytoties akan dankali ya fara kiyayewa tun a watan Yuni, kimanin wata daya da ya wuce yadda aka saba.
- A cikin dasa dankalin turawa, yawan tseren T1 ya karu, wanda a baya aka ci karo da shi a can cikin ƙarancin adadin (Ulanova et al., 2003).
- Yanayin da aka ware daga ganyen tumatir ya daina bambanta da nau'ikan dankalin turawa a cikin ɓarkewa a kan bambancin dankalin ƙwallon ƙafa kuma ya fara fin ƙarfin "dankalin turawa" a cikin zafin rai ba kawai a kan tumatir ba, har ma da dankali (Lavrova et al., 2003; Ulanova et al. , 2003).
Don haka, maimakon rarrabuwar kawuna, sai aka sami daidaituwa tsakanin jama'a, bayyanar yawan mutane a kan shuke-shuke masu daukar bakuna guda biyu masu tsananin kazantawa da zafin rai ga duka jinsunan.
ƙarshe
Don haka, duk da fiye da shekaru fiye da 150 na zurfin nazarin P. infestans, a cikin ilmin halitta, gami da ilimin halittar jama'a na wannan wakili mai haifar da mahimman cututtukan cututtukan tsire-tsire masu haɓaka, ba a san da yawa ba. Ba a bayyana yadda yadda kowane matakin rayuwar mutum yake shafar tsarin yawan jama'a ba, menene hanyoyin kwayar halittar da ke iya haifar da bambance-bambance da tashin hankali, yaya rabon tsarin haihuwa da haihuwa a cikin yawan jama'a, yadda rashin cin ganyayyaki da ciyawa suka gada, menene matsayin dankali da tumatir a cikin cutar farko ta wadannan amfanin gona da kuma a menene tasirin su akan tsarin yawan mutane masu cutar. Har zuwa yanzu, ba a warware irin waɗannan batutuwa masu mahimmanci kamar su hanyoyin halittar ɗan adam don canza tashin hankali na larura ko zaizawar ƙwarin dankalin turawa da ba shi da takamaimai. Tare da zurfafawa da fadada bincike kan dankalin turawa a ƙarshen cutar, cutar ta haifar da sabon ƙalubale ga masu bincike. Koyaya, haɓaka ƙwarewar gwaji, fitowar sabbin hanyoyin hanyoyin magudi da ƙwayoyin halitta da sunadarai ya bamu damar fatan samun nasarar warware tambayoyin da aka gabatar.
An buga labarin a cikin mujallar "Kariyar Dankali" (A'a. 3, 2017)