Селекция пшеницы: генетические основы и in vitro технологии

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 ÇËÀÊÎÂ ÑØÀ
(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). Â 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.
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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
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Ö è ö è í Í.Â. Îòäàëåííàÿ ãèáðèäèçàöèÿ ðàñòåíèé. Ì., 1954.
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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
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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., Ã 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.
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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.
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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: inna-lapochkina@yandex.ru
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 inna-lapochkina@yandex.ru
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.
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