ÑÅËÜÑÊÎÕÎÇßÉÑÒÂÅÍÍÀß ÁÈÎËÎÃÈß, 2014, ¹ 3, ñ. 77-82 Ñåëåêöèÿ ïøåíèöû: ãåíåòè÷åñêèå îñíîâû è in vitro òåõíîëîãèè ÓÄÊ 633.11:631.522/.524:576.3 ÖÈÒÎÃÅÍÅÒÈ×ÅÑÊÎÅ ÈÇÓ×ÅÍÈÅ ÊÎËËÅÊÖÈÈ ÑÈÍÒÅÒÈ×ÅÑÊÎÉ ÏØÅÍÈÖÛ ÈÇ ÍÀÖÈÎÍÀËÜÍÎÉ ÊÎËËÅÊÖÈÈ ÇËÀÊΠÑØÀ (NATIONAL SMALL GRAIN COLLECTION OF USDA-ARS)  ÓÑËÎÂÈßÕ ÍÅ×ÅÐÍÎÇÅÌÍÎÉ ÇÎÍÛ ÐÎÑÑÈÈ ∗ È.Ô. ËÀÏÎ×ÊÈÍÀ, È.Â. ÈÎÐÄÀÍÑÊÀß, Ã.Ë. ß×ÅÂÑÊÀß, ÀÄÕÀÌ ÀËÜ ËÀÁÁÀÍ Ñôîðìèðîâàííàÿ â ñîîòâåòñòâèè ñ ïðîãðàììîé CIMMYT (International Maize and Wheat Improvement Center — Ìåæäóíàðîäíûé öåíòð ïî óëó÷øåíèþ êóêóðóçû è ïøåíèöû) êîëëåêöèÿ ñèíòåòè÷åñêîé ïøåíèöû («ñèíòåòèêîâ») íàøëà øèðîêîå ïðèìåíåíèå â ñåëåêöèîííûõ ïðîãðàììàõ ìíîãèõ ñòðàí, òàê êàê îáðàçöû ëåãêî ñêðåùèâàþòñÿ ñ ñîðòàìè ìåñòíîé ñåëåêöèè, ÷òî çíà÷èòåëüíî ðàñøèðÿåò ãåíåòè÷åñêèé ïîòåíöèàë ãèáðèäîâ. Ýòà êîëëåêöèÿ «ñèíòåòèêîâ» ðàíåå íèêîãäà íå èñïûòûâàëàñü â ðåãèîíàõ Ðîññèè ïî óñòîé÷èâîñòè ê áîëåçíÿì è íå èñïîëüçîâàëàñü â ñåëåêöèîííûõ ïðîãðàììàõ. Ïðåäìåòîì íàøåãî èññëåäîâàíèÿ ñòàëà êîëëåêöèÿ èç 400 îáðàçöîâ ñèíòåòè÷åñêîé ïøåíèöû, ïîëó÷åííàÿ èç Íàöèîíàëüíîé êîëëåêöèè çëàêîâ ÑØÀ (øòàò Àéäàõî) â 2009 ãîäó äëÿ èñïûòàíèÿ åå â óñëîâèÿõ Íå÷åðíîçåìíîé çîíû íà èíôåêöèîííûõ ôîíàõ áóðîé ðæàâ÷èíû è ìó÷íèñòîé ðîñû.  ðåçóëüòàòå 2-ëåòíèõ èññëåäîâàíèé áûëè âûäåëåíû ïðîäóêòèâíûå îáðàçöû ñ óñòîé÷èâîñòüþ ê áóðîé ðæàâ÷èíå, ìó÷íèñòîé ðîñå è ñ ãðóïïîâîé óñòîé÷èâîñòüþ.  òî æå âðåìÿ ðåàêöèÿ íà ïðîíèêíîâåíèå âîçáóäèòåëÿ áóðîé ðæàâ÷èíû îêàçàëàñü ãåòåðîãåííîé, îòìå÷àëîñü òàêæå ðàñùåïëåíèå ïî ìîðôîëîãè÷åñêèì ïðèçíàêàì êîëîñà, ëèñòà, ñòåáëÿ. Äëÿ âûÿñíåíèÿ ïðè÷èí íåñòàáèëüíîñòè ó 71 îáðàçöà èç êîëëåêöèè «ñèíòåòèêîâ» èçó÷åí õàðàêòåð êîíúþãàöèè õðîìîñîì â ìåéîçå è ïîäñ÷èòàí ìåéîòè÷åñêèé èíäåêñ. Óñòàíîâëåíî, ÷òî òîëüêî 63 % îáðàçöîâ áûëè öèòîãåíåòè÷åñêè ñòàáèëüíûìè. Ó îñòàëüíûõ íàáëþäàëè âàðüèðîâàíèå ÷èñëà õðîìîñîì ó ðàñòåíèé îò 2n = 40 äî 2n = 43, îòìå÷àëè íàðóøåíèÿ â ìåòàôàçå I ìåéîçà (âûÿâëåíû óíèâàëåíòû è ìóëüòèâàëåíòû), è òàêèå ðàñòåíèÿ, êàê ïðàâèëî, èìåëè ïîíèæåííûé ìåéîòè÷åñêèé èíäåêñ. Ñäåëàíî çàêëþ÷åíèå, ÷òî â êîëëåêöèè ïðîäîëæàåòñÿ ôîðìîîáðàçîâàòåëüíûé ïðîöåññ, â îñíîâå êîòîðîãî ëåæèò ïîÿâëåíèå àíåóïëîèäíûõ ðàñòåíèé. Ýòèì îáúÿñíÿåòñÿ íàáëþäàåìîå ðàñùåïëåíèå âíóòðè îáðàçöîâ ïî ìîðôîëîãè÷åñêèì ïðèçíàêàì è óñòîé÷èâîñòè ê áîëåçíÿì. Äëÿ äàëüíåéøèõ ãåíåòè÷åñêèõ è ñåëåêöèîííî-ãåíåòè÷åñêèõ èññëåäîâàíèé âûäåëåíà ãðóïïà öèòîãåíåòè÷åñêè ñòàáèëüíûõ ôîðì, ñî÷åòàþùèõ óñòîé÷èâîñòü ê áîëåçíÿì ñ êðóïíîçåðíîñòüþ (ìàññà 1000 çåðåí äî 56 ã) è âûñîêîé ïðîäóêòèâíîñòüþ êîëîñà (äî 2,5 ã). Êëþ÷åâûå ñëîâà: ñèíòåòè÷åñêèå ãåêñàïëîèäíûå ïøåíèöû, êîëëåêöèÿ, ìåéîç. Âàæíûì èñòî÷íèêîì ðàñøèðåíèÿ ãåíåòè÷åñêîãî ðàçíîîáðàçèÿ ìÿãêîé ïøåíèöû ñëóæèò ïðèâëå÷åíèå åå äèêîðàñòóùèõ ñîðîäè÷åé, ïîëó÷åíèå ìåæâèäîâûõ è ìåæðîäîâûõ ãèáðèäîâ è âîâëå÷åíèå ýòèõ ãèáðèäîâ â äàëüíåéøåå ñêðåùèâàíèå. Ó èñòîêîâ ýòîãî íàïðàâëåíèÿ ñòîÿëè H. Kihara, McFadden è E.R. Sears (1-3), à â íàøåé ñòðàíå — Í.Â. Öèöèí è Ã.Ä. Ëàï÷åíêî (4, 5). Îäíàêî â ýòîì ñëó÷àå ðåøåíèå ïðîáëåìû óñëîæíÿåòñÿ òåì, ÷òî íàðÿäó ñ íóæíûìè ïðèçíàêàìè ãèáðèäíîìó ïîòîìñòâó ïåðåäàåòñÿ ìíîæåñòâî äðóãèõ íåæåëàòåëüíûõ ñâîéñòâ, õàðàêòåðíûõ äëÿ äèêèõ âèäîâ (6).  1990-å ãîäû â Êàíçàññêîì óíèâåðñèòåòå è â Ìåêñèêå â Ìåæäóíàðîäíîì öåíòðå ïî óëó÷øåíèþ êóêóðóçû è ïøåíèöû (CIMMYT — International Maize and Wheat Improvement Center) (7, 8) áûëè ðàçðàáîòàíû ïðîãðàììû ñîçäàíèÿ ñèíòåçèðîâàííûõ ãåêñàïëîèäíûõ (àìôèäèïëîèäíûõ) ôîðì ïøåíèöû («ñèíòåòèêîâ»). Ïî ïðîãðàììå, ïðåäëîæåííîé CIMMYT, áîëüøèíñòâî «ñèíòåòèêîâ» ïîëó÷åíî íà îñíîâå ñêðåùèâàíèÿ ñîâðåìåííîé ïøåíèöû durum (Triticum turgidum ssp. durum) c Aegilops tauschii (2n = 14) è ïîñëåäóþùåãî óäâîåíèÿ ÷èñëà õðîìîñîì, è ëèøü íåáîëüøîå ÷èñëî «ñèíòåòèêîâ» ñîçäàíî ñ ó÷àñòèåì T. dicoccoides è T. dicoccum. Ýòè «ñèíòåòèêè» íàøëè øèðîêîå ïðèìåíåíèå â ñåëåêöèîííûõ ïðî∗ Ðàáîòà âûïîëíåíà ïðè ïîääåðæêå ÌÍÒÖ, ïðîåêò 3352. 77 ãðàììàõ ìíîãèõ ñòðàí (9-12), òàê êàê ëåãêî ñêðåùèâàþòñÿ ñ ñîðòàìè ìåñòíîé ñåëåêöèè, ÷òî çíà÷èòåëüíî ðàñøèðÿåò ãåíåòè÷åñêèé ïîòåíöèàë ñîçäàâàåìûõ ãèáðèäîâ, â òîì ÷èñëå çà ñ÷åò ïåðåäà÷è íîâûõ ãåíîâ. Ñèíòåçèðîâàííûå ïøåíèöû ìîãóò áûòü èñïîëüçîâàíû â ñàìûõ ðàçíûõ íàïðàâëåíèÿõ ñåëåêöèè: â óâåëè÷åíèè ïîêàçàòåëåé ïî êîìïîíåíòàì óðîæàÿ, óñòîé÷èâîñòè ê áèîòè÷åñêèì è àáèîòè÷åñêèì ôàêòîðàì, â ïîâûøåíèè êà÷åñòâà çåðíà. Ïåðâûå êîììåð÷åñêèå ñîðòà Chunmai 38, Chunmai 42, Chunmai 43, Chunmai 47 ñ èñïîëüçîâàíèåì «ñèíòåòèêîâ» ïîëó÷åíû â Êèòàå (10). Êîëëåêöèÿ «ñèíòåòèêîâ» íèêîãäà íå èñïûòûâàëàñü â ðåãèîíàõ Ðîññèè ïî óñòîé÷èâîñòè ê áîëåçíÿì è íå èñïîëüçîâàëàñü â îòå÷åñòâåííûõ ñåëåêöèîííûõ ïðîãðàììàõ. Ïðåäìåòîì íàøåãî êîìïëåêñíîãî èññëåäîâàíèÿ ñòàëè 400 îáðàçöîâ ñèíòåòè÷åñêîé ïøåíèöû, ïîëó÷åííûõ èç Íàöèîíàëüíîé êîëëåêöèè çëàêîâ ÑØÀ (øòàò Àéäàõî) â 2009 ãîäó äëÿ èçó÷åíèÿ â óñëîâèÿõ Íå÷åðíîçåìíîé çîíû íà èíôåêöèîííûõ ôîíàõ áóðîé ðæàâ÷èíû è ìó÷íèñòîé ðîñû. Ïðîèñõîæäåíèå îáðàçöîâ êîëëåêöèè îïèñàíî ïîäðîáíî (http://www.ars-grin.gov/npgs/acc/acc_queries).  ðåçóëüòàòå 2-ëåòíèõ èññëåäîâàíèé áûëè âûäåëåíû ïðîäóêòèâíûå îáðàçöû ñ óñòîé÷èâîñòüþ ê áóðîé ðæàâ÷èíå, ìó÷íèñòîé ðîñå è ãðóïïîâîé óñòîé÷èâîñòüþ. Ó íåêîòîðûõ ìû èäåíòèôèöèðîâàëè ãåíû óñòîé÷èâîñòè ê áóðîé ðæàâ÷èíå ñ èñïîëüçîâàíèåì ìåòîäîâ ôèòîïàòîëîãè÷åñêîãî òåñòèðîâàíèÿ è ìîëåêóëÿðíûõ ìàðêåðîâ (13). Óñòàíîâëåíî òàêæå ïîâûøåííîå ñîäåðæàíèå áåëêà, êëåéêîâèíû è ìèêðîýëåìåíòîâ â çåðíå ïî ñðàâíåíèþ ñ àíàëîãè÷íûìè ïîêàçàòåëÿìè ó ñòàíäàðòíûõ ñîðòîâ ÿðîâîé ìÿãêîé ïøåíèöû (14). Èíûìè ñëîâàìè, îáðàçöû ñèíòåòè÷åñêîé ïøåíèöû äåéñòâèòåëüíî ìîãóò áûòü íîâûìè èñòî÷íèêàìè ïðèçíàêîâ äëÿ óëó÷øåíèÿ ìÿãêîé ïøåíèöû â Íå÷åðíîçåìíîé çîíå. Îäíàêî óæå â ïåðâûé ãîä èñïûòàíèÿ êîëëåêöèè áûëà îáíàðóæåíà íåîäíîðîäíîñòü ðàñòåíèé ó íåêîòîðûõ îáðàçöîâ. Îòìå÷àëè ðàñùåïëåíèå ïî ìîðôîëîãè÷åñêèì ïðèçíàêàì êîëîñà, ëèñòà, ñòåáëÿ. Íàáëþäàëè òàêæå ãåòåðîãåííóþ ðåàêöèþ íà ïðîíèêíîâåíèå áóðîé ðæàâ÷èíû (çíà÷åíèÿ îò 0-1 äî 3-4) ó ðàñòåíèé îäíîãî îáðàçöà. Äëÿ ãåíåòè÷åñêèõ èññëåäîâàíèé, êîòîðûå ïðåäïîëàãàþò óñòàíîâëåíèå ÷èñëà ãåíîâ, äåòåðìèíèðóþùèõ óñòîé÷èâîñòü ê áóðîé ðæàâ÷èíå, è èõ èäåíòèôèêàöèþ, íåîáõîäèìû ãåíåòè÷åñêè ñòàáèëüíûå ëèíèè. Ïîÿâèëèñü ñâåäåíèÿ, ÷òî íåêîòîðûå ñèíòåòè÷åñêèå ïøåíèöû èìåþò òàêîé æå âûñîêèé óðîâåíü íàðóøåíèé â ìåéîçå, êàê è òðèòèêàëå (15). Äëÿ âûÿñíåíèÿ ïðè÷èí íåñòàáèëüíîñòè îáðàçöîâ èçó÷àåìîé êîëëåêöèè áûëà ïîñòàâëåíà çàäà÷à ïî ïðîâåðêå ïîâåäåíèÿ õðîìîñîì â ìåéîçå è óñòàíîâëåíèþ ìåéîòè÷åñêîãî èíäåêñà êàê êîíå÷íîãî ïîêàçàòåëÿ ñòàáèëüíîñòè ãåíîòèïà. Ìåòîäèêà. Èçó÷èëè 71 îáðàçåö ñèíòåòè÷åñêîé ïøåíèöû èç 400, ïîëó÷åííûõ èç NSGC USDA-ARS Íàöèîíàëüíîé êîëëåêöèè çëàêîâ ÑØÀ (øòàò Àéäàõî). Àíàëèç ìåéîçà ïðîâîäèëè íà ïÿòè ðàñòåíèÿõ êàæäîãî èç îáðàçöîâ íà âðåìåííûõ äàâëåíûõ àöåòîêàðìèíîâûõ ïðåïàðàòàõ ïî îáùåïðèíÿòîé ìåòîäèêå (16). Ïðîñìàòðèâàëè ïðèìåðíî ïî 25 êëåòîê ñ êàæäîãî ðàñòåíèÿ, èçó÷àëè ïðîöåññ ñïàðèâàíèÿ õðîìîñîì â ìåòàôàçå I è îïðåäåëÿëè ìåéîòè÷åñêèé èíäåêñ (% íîðìàëüíî ñôîðìèðîâàâøèõñÿ òåòðàä). Ñòåïåíü ïîðàæåíèÿ ìó÷íèñòîé ðîñîé è áóðîé ðæàâ÷èíîé âûðàæàëè â ïðîöåíòàõ, òèï ðåàêöèè íà ïðîíèêíîâåíèå áóðîé ðæàâ÷èíû îïðåäåëÿëè ïî øêàëå Å.B. Mains è H.S. Jakson (17). Êîìïëåêñ õîçÿéñòâåííî öåííûõ ïðèçíàêîâ (âûñîòà ðàñòåíèÿ, ÷èñëî ñóòîê äî êîëîøåíèÿ, ìàññà çåðíà ñ ãëàâíîãî êîëîñà, ìàññà 1000 çåðåí) èçó÷àëè â òå÷åíèå 2009-2010 ãîäîâ ñ êîíòðàñòíûìè ïîãîäíûìè óñëîâèÿìè â ïîëåâûõ îïûòàõ (Ìîñêîâñêàÿ îáë.) 78 ×èñëî õðîìîñîì (2n), òèï êîíúþãàöèè â ìåòàôàçå I ìåéîçà è ìåéîòè÷åñêèé èíäåêñ (MI, %) ó èçó÷åííûõ îáðàçöîâ cèíòåòè÷åñêîé ïøåíèöû èç êîëëåêöèè NSGC-USDA-ARS (ÑØÀ) Îáðàçåö PI 613280 PI 613310 PI 648580 PI 648571 PI 648660 PI 648827 PI 648489 PI 648756 ×èñëî ðàñ÷èñëî òåíèé, øò. 2n ìåòàôàç I 5 6 2 1 2 1 4 1 5 1 4 5 5 74 126 13 6 28 8 62 5 77 8 33 45 74 42 42 41 43 42 40 42 41 42 41 42 42 42 Ìåòàôàçà òèï êîíúþãàöèè 21II¼20II + 2I¼19II + 4I 21II¼20II + 2I¼19II + 4I 20II + 1I¼19II + 3I 21II + 1I¼20II + 3I 21II¼20II + 2I 19II + 2I¼18II + 4I 20II + 2I¼18II + 6I 20II + 1I 21II 20II + 1I 21II¼20II + 2I 21II¼20II + 2I 21II¼19II + 4I Òåòðàäû äîëÿ íîðìàëüíûõ äîëÿ êëåòîê ñ ÷èñëî òåòðàä, øò. íàðóøåíèÿìè, % íîðìàëüíûõ ñ ìèêðîÿäðàìè òåòðàä (MI, %) 14,9 17,5 100 100 53,5 11,4 88,2 100 15,5 100 9,1 35,5 55.4 560 735 218 64 149 5 137 60 613 75 386 481 274 32 51 105 52 69 100 234 28 34 40 37 122 206 94,6 93,5 67,4 55,1 68,3 4,0 36,9 68,2 94,7 65,2 91,2 79,8 57,1 Õàðàêòåðèñòèêà îáðàçöà Ãåíîòèï ñòàáèëåí Ãåíîòèï ñòàáèëåí Îáðàçåö íåñòàáèëåí ïî 2n è õàðàêòåðó êîíúþãàöèè â ìåéîçå è èìååò íèçêèé MI Îáðàçåö íåñòàáèëåí ïî 2n è õàðàêòåðó êîíúþãàöèè â ìåéîçå è èìååò íèçêèé MI Îáðàçåö íåñòàáèëåí ïî 2n, íî ðàñòåíèÿ ñ 2n = 42 ñòàáèëüíû Îáðàçåö íåñòàáèëåí ïî 2n, íî ðàñòåíèÿ ñ 2n = 42 ñòàáèëüíû Îáðàçåö èìååò íèçêèé MI Îáðàçåö ñòàáèëåí ïî 2n, íî èìååò íàðóøåíèÿ â ìåéîçå è ïîíèæåííûé MI 79 ïðè ñðàâíåíèè ñî ñòàíäàðòîì (ñîðòà ìÿãêîé ïøåíèöû Ðîäèíà è Ëàäà). Ðàñùåïëåíèå ïî îïóøåíèþ êîëîñà, îñòèñòîñòè, íàëè÷èþ/îòñóòñòâèþ âîñêà íà êîëîñå ó÷èòûâàëè âèçóàëüíî â ïîëåâûõ óñëîâèÿõ íà âåãåòèðóþùèõ ðàñòåíèÿõ. Äîñòîâåðíîñòü ðàçëè÷èé ñî ñòàíäàðòíûì ñîðòîì îöåíèâàëè ïî ÍÑÐ05, êîòîðóþ ðàññ÷èòûâàëè ïî ìåòîäèêå, èçëîæåííîé Á.À. Äîñïåõîâûì (18), ñ èñïîëüçîâàíèåì ïðîãðàììû Microsoft Excel. Ðåçóëüòàòû. Ïîëó÷åííûå äàííûå ïîêàçàëè, ÷òî òîëüêî 45 îáðàçöîâ èç 71 áûëè öèòîëîãè÷åñêè ñòàáèëüíû. Îñòàëüíûå ëèíèè ðàñùåïëÿëèñü ïî ÷èñëó õðîìîñîì (2n), èìåëè íàðóøåíèÿ â ìåéîçå è õàðàêòåðèçîâàëèñü ðàçëè÷íîé ïëîòíîñòüþ êîíúþãàöèè õðîìîñîì. Ýòî äàëî âîçìîæíîñòü âñå èçó÷åííûå îáðàçöû ðàçäåëèòü íà ÷åòûðå óñëîâíûå ãðóïïû, íàèáîëåå òèïè÷íûå ïðåäñòàâèòåëè êîòîðûõ ïðèâåäåíû â òàáëèöå. Ó ðàñòåíèé I ãðóïïû áûëà óñòàíîâëåíà íåñòàáèëüíîñòü ÷èñëà õðîìîñîì (2n) è íàðóøåíèå ïðîöåññà ìåéîçà (ôîðìèðîâàíèå ïîâûøåííîãî ÷èñëà îòêðûòûõ áèâàëåíòîâ, óíèâàëåíòîâ è ìóëüòèâàëåíòîâ â ìåòàôàçå I è ôîðìèðîâàíèå òåòðàä ñ ìèêðîÿäðàìè). Ó 11 îáðàçöîâ îòìå÷àëè âàðüèðîâàíèå ÷èñëà õðîìîñîì îò 2n = 40 äî 2n = 43. Ýòè îáðàçöû èìåëè ïîíèæåííûé ìåéîòè÷åñêèé èíäåêñ (35-75 %) (PI 648571, PI 648599, PI 648767, PI 648842, PI 648760, PI 648757, PI 648580, PI 648529, PI 648687, PI 648843, PI 648491). Êî II ãðóïïå ðàñòåíèé îòíåñëè 10 ãåíîòèïîâ, êîòîðûå òàêæå ðàñùåïëÿëèñü ïî ÷èñëó õðîìîñîì, íî ïðè ýòîì 42-õðîìîñîìíûå ðàñòåíèÿ ïî òå÷åíèþ ìåéîçà áûëè ñòàáèëüíûìè è èìåëè âûñîêèé ìåéîòè÷åñêèé èíäåêñ (PI 648558, PI 648576, PI 648660, PI 648827, PI 648861, PI 648561, PI 648864, PI 648611, PI 648633, PI 648833).  III ãðóïïó îòîáðàëè 45 îáðàçöîâ ñ ïðàâèëüíûì òå÷åíèåì ìåéîçà è âûñîêèì (80-90 %) ìåéîòè÷åñêèì èíäåêñîì (öèòîãåíåòè÷åñêè ñòàáèëüíûå), êîòîðûå ïðèãîäíû äëÿ ïðîâåäåíèÿ èññëåäîâàíèé ïî ãåíåòèêå óñòîé÷èâîñòè ê áîëåçíÿì (PI 648528, PI 648557, PI 648559, PI 648573, PI 648575, PI 648581, PI 648586, PI 648598, PI 648603, PI 648648, PI 648650, PI 648716, PI 648724, PI 648726, PI 648572, PI 648588, PI 648649, PI 648758, PI 648752, PI 648664, PI 648681, PI 648826, PI 648508, PI 648606, PI 648865, PI 648645, PI 648629, PI 613280, PI 613306, PI 613310, PI 613320, PI 613321, PI 613322, PI 648556, PI 648608, PI 648651, PI 648655, PI 648656, PI 648661, PI 648691, PI 648693, PI 648787, PI 648851, PI 648859, PI 648860). Ïðè îöåíêå ýòèõ îáðàçöîâ â òå÷åíèå 2 ëåò â ïîëåâûõ óñëîâèÿõ ïî ñòåïåíè ïîðàæåíèÿ ìó÷íèñòîé ðîñîé è áóðîé ðæàâ÷èíîé, à òàêæå ïî êîìïëåêñó õîçÿéñòâåííî öåííûõ ïðèçíàêîâ îêàçàëîñü, ÷òî áîëüøèíñòâî èç ýòîé ãðóïïû (80 %) áûëè êðóïíîçåðíûìè è èìåëè ìàññó 1000 çåðåí áîëåå 45 ã è äîñòîâåðíî ïðåâîñõîäèëè ñîðòà ìÿãêîé ïøåíèöû Ðîäèíà è Ëàäà ïî ýòîìó ïðèçíàêó. Ìàêñèìàëüíîå ðàçâèòèå ïðèçíàêà ó ñîðòà Ðîäèíà ñîñòàâèëî 40,5 ã, à äëÿ îáðàçöîâ ñèíòåòè÷åñêîé ïøåíèöû (PI 648557, PI 648575, PI 613280, PI 613321, PI 648851, PI 648787) — 53,0-56,6 ã. Îñíîâíàÿ ÷àñòü îáðàçöîâ ñèíòåòè÷åñêîé ïøåíèöû ôîðìèðîâàëè äîñòàòî÷íî ïðîäóêòèâíûé êîëîñ (íà óðîâíå ñòàíäàðòíûõ ñîðòîâ). Òîëüêî äâà öèòîëîãè÷åñêè ñòàáèëüíûõ îáðàçöà óñòóïàëè ñòàíäàðòíûì ñîðòàì ïî ìàññå çåðíà ñ ãëàâíîãî êîëîñà (PI 648826 è PI 648606) — 1,1 ã. Ðÿä îáðàçöîâ (PI 648598, PI 648724, PI 648726, PI 648588, PI 648645, PI 613280, PI 613306, PI 648655, PI 648661, PI 648851) äîñòîâåðíî ïðåâîñõîäèëè ëó÷øèé ñîðò Ëàäà ïî ïðîäóêòèâíîñòè êîëîñà (2,2-2,5 ã ïðîòèâ 1,7 ã). Ó ýòèõ ãåíîòèïîâ ïðîäóêòèâíîñòü êîëîñà è ìàññà 1000 çåðåí íå ñíèæàëèñü è â óñëîâèÿõ íåáëàãîïðèÿòíîãî çàñóøëèâîãî 2010 ãîäà. 80 Êîíñòàíòíûå îáðàçöû ñèíòåòè÷åñêîé ïøåíèöû ïåðåäàíû äëÿ îöåíêè óñòîé÷èâîñòè ê æåëòîé è òåìíî-áóðîé ïÿòíèñòîñòÿì è èäåíòèôèêàöèè ãåíîâ óñòîé÷èâîñòè ê ðæàâ÷èííûì ãðèáàì âî Âñåðîññèéñêèé ÍÈÈ çàùèòû ðàñòåíèé (ã. Ñàíêò-Ïåòåðáóðã—Ïóøêèí).  IV ãðóïïó îòíåñëè 5 îáðàçöîâ, â êîòîðûõ âñå ïðîòåñòèðîâàííûå ðàñòåíèÿ èìåëè 2n = 42, íî â ìåòàôàçå I ìåéîçà îòìå÷àëèñü íàðóøåíèÿ: ïîâûøåííàÿ äîëÿ óíèâàëåíòîâ è ìóëüòèâàëåíòîâ, êîòîðûå ïðèâåëè ê ïîÿâëåíèþ òåòðàä ñ óâåëè÷åííûì ÷èñëîì ìèêðîÿäåð. Ýòè ðàñòåíèÿ èìåëè ïîíèæåííûé ìåéîòè÷åñêèé èíäåêñ (57,1-79,8 %). Ïûëüöà, êîòîðàÿ ñôîðìèðóåòñÿ èç òàêèõ ìèêðîñïîð, ìîæåò ñòàòü ïðè÷èíîé ïîÿâëåíèÿ àíåóïëîèäíûõ ðàñòåíèé â ñëåäóþùåì ïîêîëåíèè, òî åñòü ðàñòåíèÿ ýòèõ îáðàçöîâ åùå íåäîñòàòî÷íî ñòàáèëüíû (PI 648550, PI 648756, PI 648489, PI 648610, PI 648621). Ïîëó÷åííûå ðåçóëüòàòû ïîçâîëÿþò ïîíÿòü íàëè÷èå ðàñùåïëåíèÿ âíóòðè îáðàçöîâ ïî ìîðôîëîãè÷åñêèì ïðèçíàêàì, óñòîé÷èâîñòè ê áîëåçíÿì è ïðîäîëæèòü âûäåëåíèå è èçó÷åíèå ñòàáèëüíûõ ôîðì. Ìîæíî ñäåëàòü çàêëþ÷åíèå, ÷òî â êîëëåêöèè ñèíòåçèðîâàííûõ ïøåíèö ïðîäîëæàåò èäòè ôîðìîîáðàçîâàòåëüíûé ïðîöåññ, â îñíîâå êîòîðîãî ëåæèò ôîðìèðîâàíèå àíåóïëîèäíûõ ðàñòåíèé. Âåðîÿòíî, ÷òî íàáëþäàåìîå ðàñùåïëåíèå ïî îïóøåíèþ êîëîñà, îñòèñòîñòè, íàëè÷èþ/îòñóòñòâèþ âîñêà íà êîëîñå è ðàñòåíèè, ãåòåðîãåííàÿ ðåàêöèÿ íà ïðîíèêíîâåíèå áóðîé ðæàâ÷èíû (íàïðèìåð, îò 1/1 äî 1/3-4; îò 1/0 äî 25/3) â ïðåäåëàõ îäíîãî îáðàçöà îòðàæàåò ýòîò ôîðìîîáðàçîâàòåëüíûé ïðîöåññ. Òàêèì îáðàçîì, òîëüêî ó 63 % èç èçó÷åííûõ îáðàçöîâ ìåéîç ïðîèñõîäèë ïðàâèëüíî è ôîðìèðîâàëèñü íîðìàëüíûå òåòðàäû (ó îñòàëüíûõ íàáëþäàëè âàðüèðîâàíèå ÷èñëà õðîìîñîì ó ðàñòåíèé îò 2n = 40 äî 2n = 43, îòìå÷àëè íàðóøåíèÿ â ìåòàôàçå I ìåéîçà — áûëè âûÿâëåíû óíèâàëåíòû è ìóëüòèâàëåíòû, è òàêèå ðàñòåíèÿ, êàê ïðàâèëî, èìåëè ïîíèæåííûé ìåéîòè÷åñêèé èíäåêñ). Îáðàçöû ñ íîðìàëüíûì ìåéîçîì ìîæíî îòíåñòè ê öèòîãåíåòè÷åñêè ñòàáèëüíûì ôîðìàì, è îíè ïðèãîäíû äëÿ ãåíåòè÷åñêèõ èññëåäîâàíèé (îïðåäåëåíèÿ íàèáîëåå âåðîÿòíîãî ÷èñëà ãåíîâ, êîíòðîëèðóþùèõ óñòîé÷èâîñòü ê áóðîé ðæàâ÷èíå, ïîñòóëèðîâàíèÿ è èäåíòèôèêàöèè ãåíîâ óñòîé÷èâîñòè ìåòîäîì ôèòîïàòîëîãè÷åñêîãî òåñòèðîâàíèÿ è ÏÖÐ-àíàëèçà) è öåëåíàïðàâëåííîãî èñïîëüçîâàíèÿ âûäåëåííûõ äîíîðîâ óñòîé÷èâîñòè ê áóðîé ðæàâ÷èíå â ñåëåêöèîííûõ ïðîãðàììàõ. ËÈÒÅÐÀÒÓÐÀ 1. 2. 3. 4. 5. 6. 7. 8. 9. K i h a r a H. Discovery of the DD analyser, one of the ancestors of Triticum vulgare. Agricultural Horticulture, 1944, 19: 889-890. M c F a d d e n E.S., S e a r s E.R. The artificial synthesis of Triticum spelta. Rec. Genet. Soc. Am., 1944, 13: 26-27. M c F a d d e n E.S., S e a r s E.R. The origin of Triticum spelta and its free-threshing hexaploid relatives. J. Hered., 1946, 37: 81-89. Ö è ö è í Í.Â. Îòäàëåííàÿ ãèáðèäèçàöèÿ ðàñòåíèé. Ì., 1954. Ë à ï ÷ å í ê î Ã.Ä. Ïðèìåíåíèå ìåòîäà îòäàëåííîé ãèáðèäèçàöèè â ñåëåêöèè îçèìîé ïøåíèöû. Ñåëåêöèÿ è ñåìåíîâîäñòâî, 1967, 2: 33-38. C o x T.S., S e a r s R.G., B e q u e t t e R.K., M a r t i n T.J. Germplasm enhancement in winter wheat × Triticum tauschii backcross populations. Crop Sci., 1995, 35: 913-919. M u j e e b - K a z i A., R o s a s V., R o l d a n S. Conservation of the genetic variation of Triticum tauschii (Coss.) Schmalh. (Aegilops squarrosa auct. non L.) in synthetic hexaploid wheats (T. turgidum L. s. lat. × Triticum tauschii; 2n = 6x = 42, AABBDD) and its potential utilization for wheat improvement. Genetic Resources and Crop Evolution, 1996, 43: 129-134. E a s t w o o d R.F., L a g u d a h E.S., A p p e l s R., H a n n a h M., K o l l m o r g e n J.F. Triticum tauschii: a novel source of resistance to cereal cyst nematode (Heterodera avenae). Aust. J. Agr. Res., 1991, 42: 69-77. D r e c c e r F.M., B o r g o g n o n e G.M., O g b o n n a y a F.C., T r e t h o w a n R.M., 81 10. 11. 12. 13. 14. 15. 16. 17. 18. W i n t e r B. CYMMYT-selected derived synthetic bread wheats for rainfed environments: yield evaluation in Mexico and Australia. Field Crop. Res., 2007, 100: 218-228. Y a n g W., L i u D., L i J., Z h a n g L., W e i H., H u X., Z h e n g Y., H e Z., Z o u Y. Synthetic hexaploid wheat and its utilization for wheat genetic improvement in China. J. Genet. Genomics, 2009, 36(9): 539-546. W a r b u r t o n M.L., C r o s s a J., F r a n c o J., K a z i M., T r e t h o w a n R., R a j a r a m S., P f e i f f e r W., Z h a n g P., D r e i s i g a c k e r S., V a n G i n k e l M. Bringing wild relatives back into the family: recovering genetic diversity in CIMMYT improved wheat germplasm. Euphytica (on-line), 2006 (doi: 10.1007/s10681-005-9077-0). Z w a r t R.S., B a n s a l U.K., T h o m p s o n J.P., W i l l i a m s o n P.M., B a r i a n a H.S. QTL mapping of multiple disease resistance traits in a synthetic hexaploid bread wheat population. Proc. 11th Int. Wheat Genetic Symposium. Brisbane, 2008: 1-3. Ë à ï î ÷ ê è í à È.Ô., À ä õ à ì À ë ü Ë à á á à í , Ì à ê à ð î â à È.Þ., à à é í ó ë ë è í Í.Ð., Æ å ì ÷ ó æ è í à À.È. Îöåíêà è õàðàêòåðèñòèêà îáðàçöîâ êîëëåêöèè ñèíòåòè÷åñêîé ïøåíèöû (2n = 42) êàê íîâûõ èñòî÷íèêîâ óñòîé÷èâîñòè ê áóðîé ðæàâ÷èíå è ìó÷íèñòîé ðîñå â óñëîâèÿõ Íå÷åðíîçåìíîé çîíû ÐÔ. Èçâåñòèÿ ÒÑÕÀ, 2011, 6: 39-48. À ä õ à ì À ë ü Ë à á á à í, Ì è í å å â à Â.À., Ë à ï î ÷ ê è í à È.Ô. Ïèòàòåëüíàÿ öåííîñòü çåðíà ñèíòåòè÷åñêèõ ïøåíèö (T. aestivum) èç êîëëåêöèè Íàöèîíàëüíîãî Öåíòà ãåíåòè÷åñêèõ ðåñóðñîâ ÑØÀ. Òåîðåòè÷åñêèå è ïðèêëàäíûå ïðîáëåìû àãðîïðîìûøëåííîãî êîìïëåêñà, 2011, 4: 35-40. M a s o u m e h R e z a e i, A h m a d A r z a n i, B a d r a l d i n E b r a h i m S a y e d - T a b a t a b a e i. Meiotic behavior of tetraploid wheats (Triticum turgidum L.) and their synthetic hexaploid wheat derivates influenced by meiotic restitution and heat stress. J. Genet., 2010, 89(4): 401-407. ß ö ê î Â.Ï., ß ÷ å â ñ ê à ÿ Ã.Ë., Ï à í è í à Å.Á., Ô å ä î ð î â à Ò.Í. Ê ìåòîäèêå èçó÷åíèÿ ìåéîçà ó çëàêîâ. Öèòîëîãèÿ è ãåíåòèêà, 1972, 6(3): 253-258. M a i n s E.B., J a c k s o n H.S. Physiologic specialization on the leaf rust of wheat Puccinia triticina Erikss. Phytopathology, 1926, 16(1): 89-120. Ä î ñ ï å õ î â Á.À. Ìåòîäèêà ïîëåâîãî îïûòà. Ì., 1979. ÃÍÓ Ìîñêîâñêèé ÍÈÈ ñåëüñêîãî õîçÿéñòâà «Íåì÷èíîâêà» Ðîññåëüõîçàêàäåìèè, Ïîñòóïèëà â ðåäàêöèþ 21 ÿíâàðÿ 2013 ãîäà 143026 Ðîññèÿ, Ìîñêîâñêàÿ îáë., Îäèíöîâñêèé ð-í, ð.ï. Íîâîèâàíîâñêîå, óë. Êàëèíèíà, 1, e-mail: [email protected] CYTOGENETIC STUDY OF SYNTHETIC WHEATS FROM NATIONAL SMALL GRAIN COLLECTION OF USDA-ARS IN THE CONDITIONS OF NECHERNOZEMNAYA ZONE OF RUSSIA I.F. Lapochkina, I.V. Iordanskaya, G.L. Yachevskaya, Adkham Al Labban Moscow Research Institute of Agriculture «Nemchinovka», Russian Academy of Agricultural Sciences, 1, ul. Kalinina, r.p. Novoivanovskoe, Odintsovo Region, Moscow Province, 143026 Russia, e-mail [email protected] Received January 21, 2013 Supported by ISTC, project 3352 Abstract CIMMYT collection of synthetic wheats is widely used in breeding programs in many countries due to easy hybridization with domestic varieties, which enhances positive effects in the hybrids. In Russia this collection has never been checked for plant resistance to diseases or used for breeding. We investigated 400 samples of synthetic wheats, obtained in 2009 from National Small Grains Collection (NSGC USDA-ARS, Idaho, USA) to be tested in Nechernozemnaya zone of Russia under brown rust and mildew infection. During 2 years the high-yielding samples were found which were resistant to brown rust, to mildew, and to both pathogens. Nevertheless, the plant response to brown rust attack differed, and the segregation was observed on morphological parameters of ears, leaves and stems. To elucidate why the instability appears, the chromosome conjugation in meiosis was examined and a meiotic index was calculated in 71 synthetic wheats. It was shown that only 63 % of the samples were cytogenetically stable. In the remaining samples the chromosome number varied from 2n = 40 to 2n = 43, and different abnormalities (univalents and multivalents) were observed in meiotic metaphase I. Usually in these plants the meiotic index was lower. It is concluded that in the collection a genetic stabilization is still continuing by the appearance of aneuploids. For further genetic and breeding investigations, there are selected the forms with stable cytogenetic parameters, disease resistance and good yielding components (weight per 1000 kernels up to 56 g, and grain yield per ear up to 2.5 g). Keywords: synthetic hexaploid wheats, wheat collection, meiosis. 82