Динамика популяций и изменчивость генома у лошадей

ÑÅËÜÑÊÎÕÎÇßÉÑÒÂÅÍÍÀß ÁÈÎËÎÃÈß, 2014, ¹ 4, ñ. 35-41
Äèíàìèêà ïîïóëÿöèé è èçìåí÷èâîñòü ãåíîìà ó ëîøàäåé
ÓÄÊ 636.12/.13:636.082:575.222.5:577.2:575.174
doi: 10.15389/agrobiology.2014.4.35rus
ÈÍÁÐÈÄÈÍÃ È ÑÒÅÏÅÍÜ
ÃÎÌÎÇÈÃÎÒÍÎÑÒÈ ÌÈÊÐÎÑÀÒÅËËÈÒÍÛÕ ËÎÊÓÑÎÂ Ó ËÎØÀÄÅÉ
(Equus caballus) ÎÐËÎÂÑÊÎÉ ÐÛÑÈÑÒÎÉ ÏÎÐÎÄÛ
Ë.À. ÕÐÀÁÐÎÂÀ, Í.Â. ÁËÎÕÈÍÀ, À.Â. ÓÑÒÜßÍÖÅÂÀ
Îöåíêà æèâîòíûõ ïî ðîäîñëîâíûì íå ïîçâîëÿåò îòñëåæèâàòü ôàêòè÷åñêîå ñõîäñòâî
ïðîáàíäà ñ ïðåäêàìè, â òîì ÷èñëå ïðè èñïîëüçîâàíèè èíáðèäèíãà. Îáùåå ÷èñëî îïðåäåëÿåìûõ ó ëîøàäåé ìàðêåðíûõ ãåíîâ óæå äîñòèãëî íåñêîëüêèõ ñîòåí, ÷òî äàåò âîçìîæíîñòü íàäåæíî êîíòðîëèðîâàòü çíà÷èòåëüíóþ ÷àñòü ãåíîìà æèâîòíûõ è èñïîëüçîâàòü ãåíåòè÷åñêóþ
èíôîðìàöèþ â ïðàêòè÷åñêîé ñåëåêöèè. Ðàíåå ìû ïîêàçàëè, ÷òî ó ÷èñòîêðîâíîé âåðõîâîé ïîðîäû ñòåïåíü ãîìîçèãîòíîñòè ïî ìèêðîñàòåëëèòíûì ëîêóñàì íå èìååò ïðÿìîé çàâèñèìîñòè
îò ñòåïåíè èíáðèäèíãà â ðîäîñëîâíîé, è ñàìûé áîëüøîé ðàçìàõ âàðèàáåëüíîñòè ýòîãî ïîêàçàòåëÿ íàáëþäàåòñÿ â ãðóïïå èíáðåäíûõ ëîøàäåé.  íàñòîÿùåé ðàáîòå íàìè áûëî èçó÷åíî
âëèÿíèå èíáðèäèíãà íà èíäèâèäóàëüíîì óðîâíå ó ëîøàäåé îðëîâñêîé ðûñèñòîé ïîðîäû. Ïðîâåäåí àíàëèç âçàèìîñâÿçè ìåæäó ñòåïåíüþ ãîìîçèãîòíîñòè ïî 16 ìèêðîñàòåëëèòíûì ëîêóñàì
(AHT4, AHT5, ASB2, ASB17, ASB23, CA425, HMS1, HMS2, HMS3, HMS6, HMS7, HTG4,
HTG6, HTG7, HTG10, VHL20) è ïîêàçàòåëåì èíáðèäèíãà ó 1194 îñîáåé èç 14 õîçÿéñòâ (âåäóùèå êîííûå çàâîäû è ïëåìåííûå ôåðìû) áîëüøèíñòâà ðåãèîíîâ Ðîññèè (æåðåáöû-ïðîèçâîäèòåëè — 190 ãîë., ìàòêè — 688 ãîë., ìîëîäíÿê — 316 ãîë.). Äàííûå ïî ëîêóñó LEX 3, ëîêàëèçîâàííîìó â Õ-õðîìîñîìå, íå ó÷èòûâàëè, ÷òîáû ïîëó÷èòü ñîïîñòàâèìûå ðåçóëüòàòû ïî ñòåïåíè
ãîìî-/ãåòåðîçèãîòíîñòè ó ëîøàäåé ðàçíîãî ïîëà. Áûëî îïðåäåëåíî, ÷òî ñòåïåíü ãîìîçèãîòíîñòè
îðëîâñêèõ ðûñàêîâ âàðüèðóåò â î÷åíü øèðîêîì èíòåðâàëå (îò 0 äî 75,0 %) è èìååò íîðìàëüíîå
ðàñïðåäåëåíèå ïðè ïîëîæèòåëüíîì êîýôôèöèåíòå àñèììåòðèè As = 0,186 è íåáîëüøîì îòðèöàòåëüíîì ýêñöåññå Ex = −0, 039. Íàèáîëüøèé ðàçáðîñ ïîêàçàòåëåé ñòåïåíè ãîìîçèãîòíîñòè çàðåãèñòðèðîâàëè â ãðóïïå ëîøàäåé ñ óìåðåííûì èíáðèäèíãîì (Fx = 0,1-1,0 %). Ñòåïåíü ãîìîçèãîòíîñòè àóòáðåäíûõ ëîøàäåé òàêæå âàðüèðîâàëà â øèðîêîì äèàïàçîíå — îò 0 (100 % ãåòåðîçèãîòû) äî 56,3 %, òî åñòü äîñòèãàëà çíà÷åíèé ãîðàçäî âûøå è íèæå ñðåäíåãî ïîêàçàòåëÿ ïî ïîïóëÿöèè.  ïðåäûäóùèõ èññëåäîâàíèÿõ òàêàÿ æå çàêîíîìåðíîñòü áûëà îòìå÷åíà ó ëîøàäåé ÷èñòîêðîâíîé âåðõîâîé ïîðîäû.  ñðåäíåì êîýôôèöèåíò èíáðèäèíãà ó ïðîòåñòèðîâàííîãî ïîãîëîâüÿ
ñîñòàâèë 1,61 % ïî Ðàéòó è ïðàêòè÷åñêè íå èçìåíèëñÿ çà ÷åòûðå ïîñëåäíèõ äåñÿòèëåòèÿ. Ïðè
ýòîì ñðåäíèé êîýôôèöèåíò èíáðèäèíãà ó ïðîäóöèðóþùèõ æåðåáöîâ è ìàòîê äîñòèã ñîîòâåòñòâåííî 1,65 è 1,67 %, ó ðûñèñòîãî ìîëîäíÿêà — ñîñòàâèë 1,45 %. Ïî ìåðå óâåëè÷åíèÿ êîýôôèöèåíòà èíáðèäèíãà ñòåïåíü ãîìîçèãîòíîñòè ìåäëåííî âîçðàñòàëà âî âñåõ ïîëîâîçðàñòíûõ ãðóïïàõ ëîøàäåé. Çàìåòíîå ïîâûøåíèå ãîìîçèãîòíîñòè áûëî âûÿâëåíî òîëüêî ïðè ñòåïåíè èíáðèäèíãà 6,10 % è áîëåå, òî åñòü íà ãðàíèöå èíáðåäíîé äåïðåññèè ïî õîçÿéñòâåííî ïîëåçíûì ïðèçíàêàì. Êîýôôèöèåíò êîððåëÿöèè ìåæäó ñòåïåíüþ ãîìîçèãîòíîñòè ìèêðîñàòåëëèòíûõ ëîêóñîâ
è ñòåïåíüþ èíáðèäèíãà ñîñòàâèë 0,079, êîýôôèöèåíò ðåãðåññèè — 0,546. Âî âñåõ èññëåäîâàííûõ ãðóïïàõ îðëîâñêèõ ðûñàêîâ îòìå÷àëîñü ñíèæåíèå ñòåïåíè ãîìîçèãîòíîñòè ïðè èíáðèäèíãå
5,1-6,0 %, ÷òî îáóñëîâëåíî ïðèìåíåíèåì êîìïëåêñíîãî èíáðèäèíãà (íà äâóõ ïðåäêîâ è áîëåå).
Êëþ÷åâûå ñëîâà: ãîìîçèãîòíîñòü, èíáðèäèíã, ìèêðîñàòåëëèòû, ÄÍÊ, îðëîâñêàÿ ðûñèñòàÿ ïîðîäà ëîøàäåé.
Ñîâåðøåíñòâîâàíèå îöåíêè ïëåìåííûõ êà÷åñòâ ñ èñïîëüçîâàíèåì
ïîêàçàòåëåé, àññîöèèðîâàííûõ ñ ôóíêöèîíàëüíî âàæíûìè ãåíàìè èëè èõ
êîìïëåêñàìè, ñîñòàâëÿåò îñíîâíîå íàïðàâëåíèå ãåíåòè÷åñêèõ èññëåäîâàíèé â ñîâðåìåííîì æèâîòíîâîäñòâå.  íàñòîÿùåå âðåìÿ â ñåëåêöèè ñåëüñêîõîçÿéñòâåííûõ ïîðîä âñå øèðå èñïîëüçóþòñÿ ìîëåêóëÿðíî-ãåíåòè÷åñêèå ìåòîäû, ðàçðàáàòûâàþòñÿ è âíåäðÿþòñÿ òåõíîëîãèè ïîëíîãî ãåíîìíîãî àíàëèçà (1-4). Áëàãîäàðÿ óñïåøíîìó çàâåðøåíèþ ïðîåêòà ïî èçó÷åíèþ
ãåíîìà ëîøàäè îáùåå ÷èñëî ìàðêåðíûõ ãåíîâ, îïðåäåëÿåìûõ ó ýòîé ãåíåòè÷åñêîé ãðóïïû, óæå äîñòèãëî íåñêîëüêèõ ñîòåí, ÷òî ïîçâîëÿåò íàäåæíî
êîíòðîëèðîâàòü çíà÷èòåëüíóþ ÷àñòü ãåíîìà è èñïîëüçîâàòü ïîëó÷àåìóþ
èíôîðìàöèþ â ïðàêòè÷åñêîé ñåëåêöèè (5-7).
Îäèí èç ïåðñïåêòèâíûõ ìåòîäîâ ÄÍÊ-òåñòèðîâàíèÿ æèâîòíûõ —
èçó÷åíèå ìèêðîñàòåëëèòíûõ ëîêóñîâ, õàðàêòåðèçóþùèõñÿ âûñîêèì ïîëè35
ìîðôèçìîì. Ïåðåõîä áîëüøèíñòâà ãåíåòè÷åñêèõ ëàáîðàòîðèé íà òåñòèðîâàíèå è êîíòðîëü ïðîèñõîæäåíèÿ ëîøàäåé ïî ïîëèìîðôíûì ëîêóñàì ìèêðîñàòåëëèòîâ ÄÍÊ ñóùåñòâåííî ðàñøèðèë êðóã ãåíåòè÷åñêèõ ìàðêåðîâ,
èìåþùèõñÿ â ðàñïîðÿæåíèè ñåëåêöèîíåðîâ. Äëÿ ïðàêòè÷åñêîé ñåëåêöèè
âñåõ âèäîâ æèâîòíûõ îñîáåííî âàæíî, ÷òî øèðîêî ðàñïðîñòðàíåííûå â
ãåíîìå ëîêóñû ñàòåëëèòíîé ÄÍÊ ìîãóò áûòü èñïîëüçîâàíû â êà÷åñòâå ãåíåòè÷åñêèõ ìàðêåðîâ ïðè îïðåäåëåíèè ñòåïåíè ãåòåðîçèãîòíîñòè æèâîòíûõ êàê âàæíîé ñîñòàâëÿþùåé èíäèâèäóàëüíîãî ãåíîìíîãî àíàëèçà.
Äëÿ ïðàêòè÷åñêîé ñåëåêöèè íåñîìíåííûé èíòåðåñ ïðåäñòàâëÿåò
îöåíêà ýôôåêòèâíîñòè ñèñòåì ðàçâåäåíèÿ æèâîòíûõ íà ãåíîìíîì óðîâíå.
Ñ ãåíåòè÷åñêîé òî÷êè çðåíèÿ ãåíîòèïè÷åñêàÿ îöåíêà æèâîòíûõ ïî ðîäîñëîâíûì íåäîñòàòî÷íî èíôîðìàòèâíà, òàê êàê íå ïîçâîëÿåò êîíòðîëèðîâàòü ôàêòè÷åñêîå ñõîäñòâî ïðîáàíäà ñ ïðåäêàìè, â òîì ÷èñëå ïðè èñïîëüçîâàíèè èíáðèäèíãà (8, 9). Íàøè èññëåäîâàíèÿ íà ëîøàäÿõ ÷èñòîêðîâíîé
âåðõîâîé ïîðîäû ïîêàçàëè, ÷òî ñòåïåíü èõ ãîìîçèãîòíîñòè ïî ìèêðîñàòåëëèòíûì ëîêóñàì íå èìååò ïðÿìîé çàâèñèìîñòè îò ñòåïåíè èíáðèäèíãà â
ðîäîñëîâíîé è ñàìûé áîëüøîé ðàçìàõ âàðèàáåëüíîñòè ýòîãî ïîêàçàòåëÿ
íàáëþäàåòñÿ â ãðóïïå èíáðåäíûõ ëîøàäåé (10). Èíòåðåñíûå ðåçóëüòàòû ïî
âçàèìîñâÿçè ìåæäó ñòåïåíüþ ãåòåðîçèãîòíîñòè ïî ìèêðîñàòåëëèòíûì ëîêóñàì, èíáðèäèíãîì è ìîðôîëîãè÷åñêèìè ïðèçíàêàìè ïîëó÷åíû ïðè èçó÷åíèè ëîøàäåé ëèïèöàíñêîé ïîðîäû (11). Óñòàíîâëåíî, ÷òî íà ñòåïåíü ãåòåðîçèãîòíîñòè èíáðåäíûõ ëîøàäåé çàìåòíî âëèÿþò ãåíåòè÷åñêèå ïàðàìåòðû êàæäîé ïîïóëÿöèè, ïîýòîìó ïðèìåíåíèå ðîäñòâåííîãî ðàçâåäåíèÿ (äàæå
â ïðåäåëàõ îäíîé ïîðîäû) â ðàçíûõ õîçÿéñòâàõ ïðèâîäèò ê íåîäèíàêîâûì
ðåçóëüòàòàì. Òàêæå áûëî ïîêàçàíî, ÷òî ïî ìåðå óâåëè÷åíèÿ êîýôôèöèåíòà
èíáðèäèíãà ó ëîøàäåé íàáëþäàåòñÿ óìåíüøåíèå äëèíû øåè è ïÿñòè (11).
Ïðîâåäåííîå â íàøåé ëàáîðàòîðèè èçó÷åíèå ãåíåòè÷åñêîé ñòðóêòóðû çàâîäñêèõ è ìåñòíûõ ïîðîä ëîøàäåé ïî ëîêóñàì ìèêðîñàòåëëèòîâ
ÄÍÊ ïîêàçàëî, ÷òî îðëîâñêèé ðûñàê — îäíà èç ñàìûõ ãåíåòè÷åñêè ñàìîáûòíûõ ïîðîä: äëÿ íåãî õàðàêòåðíû âûñîêèå ïîêàçàòåëè ãåíåòè÷åñêîãî
ðàçíîîáðàçèÿ ïðè ñòàáèëüíîñòè àëëåëîôîíäà (12).
 íàøèõ èññëåäîâàíèÿõ áûëà ïîñòàâëåíû çàäà÷à èçó÷èòü âîçìîæíîñòü èñïîëüçîâàíèÿ ìèêðîñàòåëëèòíûõ ëîêóñîâ, ïðèìåíÿåìûõ ïðè êîíòðîëå ïðîèñõîæäåíèÿ, äëÿ îöåíêè ñòåïåíè ãåòåðîçèãîòíîñòè ëîøàäåé îðëîâñêîé ðûñèñòîé ïîðîäû, à òàêæå âëèÿíèå èíáðèäèíãà íà ñòåïåíü ãîìîçèãîòíîñòè ðûñàêîâ íà èíäèâèäóàëüíîì óðîâíå.
Ìåòîäèêà. Îáúåêòîì ãåíåòèêî-ïîïóëÿöèîííîãî àíàëèçà, ïðîâåäåííîãî â 2010-2013 ãîäàõ, áûëè ëîøàäè îðëîâñêîé ðûñèñòîé ïîðîäû: 190 æåðåáöîâ-ïðîèçâîäèòåëåé, 688 ìàòîê è 316 ãîë. ìîëîäíÿêà (âñåãî 1194 ãîë.),
ïðèíàäëåæàùèõ 14 âåäóùèì õîçÿéñòâàì è ÷àñòíûì âëàäåëüöàì èç ðàçíûõ
ðåãèîíîâ Ðîññèè.
ÄÍÊ èç âîëîñÿíûõ ëóêîâèö è îáðàçöîâ êðîâè ëîøàäåé âûäåëÿëè ñ
èñïîëüçîâàíèåì íàáîðîâ ExtraGeneÒÌ DNA Prep 200 è DiatomÒÌ DNA
Prep 200 (ÎÎÎ «Ëàáîðàòîðèÿ Èçîãåí», Ðîññèÿ). Àìïëèôèêàöèþ ÄÍÊ ïðîâîäèëè â ÏÖÐ ñ ìóëüòèïëåêñíûì íàáîðîì ïðàéìåðîâ StockMarks® for
Horses Equine Genotyping Kit («Applied Biosystems», ÑØÀ). Äëÿ ýëåêòðîôîðåòè÷åñêîãî ðàçäåëåíèÿ ïðîäóêòîâ àìïëèôèêàöèè èñïîëüçîâàëè 4-êàíàëüíûé àâòîìàòè÷åñêèé àíàëèçàòîð ABI 3130 («Applied Biosystems», ÑØÀ).
Òåñòèðîâàíèå âûïîëíÿëè ïî 16 ëîêóñàì ìèêðîñàòåëëèòíîé ÄÍÊ: AHT4,
AHT5, ASB2, ASB17, ASB23, CA425, HMS1, HMS2, HMS3, HMS6, HMS7,
HTG4, HTG6, HTG7, HTG10, VHL20.
Ñòåïåíü ãîìîçèãîòíîñòè ëîøàäåé (Ñà) îïðåäåëÿëè êàê ñðåäíèé
ñóììàðíûé ïîêàçàòåëü ïî ïðîòåñòèðîâàííûì ìèêðîñàòåëëèòíûì ëîêóñàì.
36
Êîýôôèöèåíò èíáðèäèíãà Fx ðàññ÷èòûâàëè ïî ôîðìóëå Ðàéòà â îáùåïðèíÿòîé ìîäèôèêàöèè À.Ä. Êèñëîâñêîãî ñ èñïîëüçîâàíèåì ðàçðàáîòàííîãî
âî Âñåðîññèéñêîì ÍÈÈ êîíåâîäñòâà ïðîãðàììíîãî îáåñïå÷åíèÿ ÈÏÑ
ÊÎÍÈ-3 ñ ó÷åòîì ðîäîñëîâíîé ïÿòè ïîêîëåíèé. Õàðàêòåð íîðìàëüíîñòè
ðàñïðåäåëåíèÿ ñòåïåíè ãîìîçèãîòíîñòè ìèêðîñàòåëëèòíûõ ëîêóñîâ ëîøàäåé îöåíèâàëè ïî àëãîðèòìó Ïèðñîíà è äðóãèì êðèòåðèÿì (13).
Ìàòåìàòè÷åñêóþ îáðàáîòêó äàííûõ ïðîâîäèëè ïðè ïîìîùè êîìïüþòåðíûõ ïðîãðàìì MS Excel è Statistica v. 6.0.
Ðåçóëüòàòû. Ó÷èòûâàÿ ðàçíóþ èíòåíñèâíîñòü îòáîðà, ãåíåòèêî-ïîïóëÿöèîííûé àíàëèç ïðîâîäèëè â òðåõ ãðóïïàõ ëîøàäåé (æåðåáöû, ìàòêè,
ìîëîäíÿê). Êðîìå òîãî, èíáðåäíûå ëîøàäè áûëè ïîäðàçäåëåíû íà ãðóïïû
â çàâèñèìîñòè îò êîýôôèöèåíòà èíáðèäèíãà Fx ñ èíòåðâàëîì 1,0 %. Äàííûå ïî ëîêóñó LEX 3, ëîêàëèçîâàííîìó â Õ-õðîìîñîìå, íå áûëè âêëþ÷åíû â îáðàáîòêó, ÷òîáû ïîëó÷èòü ñîïîñòàâèìûå äàííûå î ñòåïåíè ãîìî- è
ãåòåðîçèãîòíîñòè ëîøàäåé ðàçíîãî ïîëà.
Ïðè àíàëèçå ðîäîñëîâíûõ ó 1194 îðëîâñêèõ ðûñèñòûõ ëîøàäåé óñòàíîâëåíî, ÷òî òîëüêî 13,7 % æåðåáöîâ-ïðîèçâîäèòåëåé, 14,0 % ìàòîê è
20,6 % ìîëîäíÿêà áûëè ïîëó÷åíû ìåòîäîì íåðîäñòâåííîãî ñïàðèâàíèÿ,
èëè àóòáðèäèíãà (òàáë. 1), è íå èìåëè ïîâòîðÿþùèõñÿ êëè÷åê ïðåäêîâ â
ïÿòè ðÿäàõ ðîäîñëîâíîé. Â ñòðóêòóðå ïîãîëîâüÿ èíáðåäíûõ ëîøàäåé âñåõ
ïîëîâîçðàñòíûõ ãðóïï ÿâíî ïðåîáëàäàëè îñîáè ñ îòäàëåííûì èíáðèäèíãîì (Fx = 0,1-1,0 %). Ñ èñïîëüçîâàíèåì òàêîãî èíáðèäèíãà áûëî ñâÿçàíî
ïðîèñõîæäåíèå 422 îñîáåé, ÷òî â öåëîì ñîñòàâèëî 35,34 % îò âñåãî ïðîàíàëèçèðîâàííîãî ïîãîëîâüÿ (ñì. òàáë. 1). Ê ýòîé ãðóïïå îòíîñèëèñü 66
æåðåáöîâ-ïðîèçâîäèòåëåé (34,74 %), 261 ìàòêà (37,94 %) è ïî÷òè òðåòüÿ
÷àñòü ïîãîëîâüÿ ìîëîäíÿêà (30,06 %).
1. Ðàñïðåäåëåíèå æèâîòíûõ (%) ñ ðàçíûì êîýôôèöèåíòîì èíáðèäèíãà Fx ïî
ïîëîâîçðàñòíûì ãðóïïàì íà îñíîâàíèè àíàëèçà ðîäîñëîâíûõ ó èññëåäîâàííûõ ëîøàäåé (Equus caballus) îðëîâñêîé ðûñèñòîé ïîðîäû (14 âåäóùèõ õîçÿéñòâ èç ðåãèîíîâ Ðîññèè, 2010-2013 ãîäû)
Ãðóïïà
ëîøàäåé
n
Æåðåáöû
190
Êîáûëû
688
Ìîëîäíÿê 316
Âñåãî
1194
0
13,68
13,95
20,57
15,66
Fx(%) ïî Ðàéòó
0,1-1,0 1,1-2,0 2,1-3,0 3,1-4,0 4,1-5,0 5,1-6,0 6,1-7,0 7,1 è áîëåå
34,74
37,94
30,06
35,34
21,58
18,17
24,05
20,27
6,84
6,69
12,66
8,29
14,74
13,08
6,33
11,56
4,74
5,38
2,53
4,52
1,05
0,87
1,27
1,01
1,58
1,89
0,95
1,59
1,05
2,03
1,58
1,76
Ïðàêòè÷åñêè ïÿòàÿ ÷àñòü ñîâðåìåííîãî ïëåìåííîãî ñîñòàâà îðëîâñêèõ ðûñàêîâ (20,27 %) èìåëà êîýôôèöèåíò èíáðèäèíãà â ïðåäåëàõ 1,12,0 % ïî Ðàéòó, è â èõ ðîäîñëîâíûõ ïîâòîðÿëèñü êëè÷êè ïðåäêîâ â III-IV
è II-V ðÿäàõ ðîäîñëîâíîé èëè âñòðå÷àëñÿ êîìïëåêñíûé èíáðèäèíã â ñòåïåíè IV-IV è â áîëåå îòäàëåííûõ ñòåïåíÿõ ðîäñòâà. Ïðè ýòîì ðàçëè÷èÿ
ìåæäó ãðóïïàìè ëîøàäåé îêàçàëèñü íåñóùåñòâåííûìè
Äîëÿ æèâîòíûõ ñ êîýôôèöèåíòîì èíáðèäèíãà 2,1-3,0 % áûëà ñóùåñòâåííî íèæå (ñì. òàáë. 1), ïîâûøàÿñü, òåì íå ìåíåå, â ãðóïïå ñ Fx 3,14,0 % ó ëîøàäåé, â ðîäîñëîâíûõ êîòîðûõ ïîâòîðÿëèñü êëè÷êè ïðåäêîâ â
III-IV è II-V ðÿäàõ èëè âñòðå÷àëñÿ êîìïëåêñíûé èíáðèäèíã â ñòåïåíè IVIV è V-V. Ïðè ïîñëåäóþùåì ïîâûøåíèè êîýôôèöèåíòà èíáðèäèíãà ÷èñëî
îñîáåé ñ ñîîòâåòñòâóþùèìè çíà÷åíèÿìè Fx ðåçêî ñíèæàëîñü (ñì. òàáë. 1), è
òîëüêî 1,76 % ëîøàäåé ýòîé ïîðîäû áûëè ïîëó÷åíû ñ èñïîëüçîâàíèåì
êîìïëåêñíîãî èíáðèäèíãà 7,1 % è áîëåå. Ñðåäíèé êîýôôèöèåíò èíáðèäèíãà ïî âñåìó îáñëåäîâàííîìó ïîãîëîâüþ ñîñòàâèë 1,61 % ïî Ðàéòó, òî
åñòü ïðàêòè÷åñêè íå óâåëè÷èëñÿ çà ÷åòûðå ïîñëåäíèõ äåñÿòèëåòèÿ (14).
Ïðè ýòîì ñðåäíèé êîýôôèöèåíò èíáðèäèíãà ïðîäóöèðóþùèõ æåðåáöîâ è
37
ìàòîê ðàâíÿëñÿ ñîîòâåòñòâåííî 1,65 è 1,67 %, òîãäà êàê ó ðûñèñòîãî ìîëîäíÿêà îí îêàçàëñÿ äàæå íèæå — 1,45 %.
Àíàëèç ñòðóêòóðû ïîãîëîâüÿ ëîøàäåé îðëîâñêîé ðûñèñòîé ïîðîäû ïî ñðåäíåé ñòåïåíè ãîìîçèãîòíîñòè ïîêàçàë, ÷òî âî âñåõ ïîëîâîçðàñòíûõ ãðóïïàõ (íåçàâèñèìî îò ðàçäåëåíèÿ ïî Fx) è â öåëîì ïî ïîïóëÿöèè îíà ñîîòâåòñòâóåò íîðìàëüíîìó ðàñïðåäåëåíèþ (Ð > 0,999) ïðè ïîëîæèòåëüíîì êîýôôèöèåíòå àñèììåòðèè As = 0,186 è íåáîëüøîì îòðèöàòåëüíîì ýêñöåññå Ex = −0,039. Ïðè ýòîì ñàìàÿ ìíîãî÷èñëåííàÿ ãðóïïà
èç 260 ëîøàäåé (21,78 %) îáðàçîâàëà ìîäàëüíûé êëàññ ñî ñòåïåíüþ ãîìîçèãîòíîñòè 31,25 %.
Âàæíî îòìåòèòü, ÷òî ñàìûé áîëüøîé ðàçáðîñ ïî ïîêàçàòåëÿì ñòåïåíè ãîìîçèãîòíîñòè áûë çàðåãèñòðèðîâàí â ãðóïïå ëîøàäåé ñ óìåðåííûì
èíáðèäèíãîì (Fx = 0,1-1,0 %).  ýòîé ãðóïïå îòìå÷àëè øèðîêèé èíòåðâàë
êîëåáàíèé — îò 0 äî 75,00 %. Ñòåïåíü ãîìîçèãîòíîñòè àóòáðåäíûõ ëîøàäåé òàêæå âàðüèðîâàëà â áîëüøîì äèàïàçîíå — îò 0 (100 % ãåòåðîçèãîòû)
äî 56,25 %, òî åñòü äîñòèãàëà çíà÷åíèé ãîðàçäî âûøå è íèæå ñðåäíåé ïî
ïîïóëÿöèè. Â ïðåäûäóùèõ èññëåäîâàíèÿõ òàêàÿ æå çàêîíîìåðíîñòü áûëà
îïèñàíà ó ëîøàäåé ÷èñòîêðîâíîé âåðõîâîé ïîðîäû (10).
Ó æåðåáöîâ-ïðîèçâîäèòåëåé ïðè óâåëè÷åíèè êîýôôèöèåíòà èíáðèäèíãà íå îòìå÷àëè çàìåòíîãî ðîñòà ñòåïåíè ãîìîçèãîòíîñòè, è àíàëèçèðóåìûé ïîêàçàòåëü ìåíÿëñÿ çèãçàãîîáðàçíî (òàáë. 2.). Ïðè ýòîì âûñîêèå
êîýôôèöèåíòû èíáðèäèíãà (ïîðÿäêà Fx = 6,01 % è áîëåå) áûëè âûÿâëåíû
òîëüêî ó ïÿòè ïðîèçâîäèòåëåé, äâîå èç êîòîðûõ èìåëè äîñòàòî÷íî âûñîêóþ ñòåïåíü ãåòåðîçèãîòíîñòè — ïðèìåðíî 75,00 % (ïðè ñðåäíåì ïîêàçàòåëå ïî ïðîòåñòèðîâàííûì îðëîâñêèì ðûñàêàì 68,54 %). Íàèáîëåå ãîìîçèãîòíûì ñðåäè ïðîèçâîäèòåëåé îêàçàëñÿ æåðåáåö Èëâà÷ 2005 ãîäà ðîæäåíèÿ
îò Ëèñòîïàäà è Èçëó÷èíû, êîòîðûé áûë èíáðèäèðîâàí íà æåðåáöà Ïåðâà÷à â ñòåïåíè III, IV-IV è êîáûëó Èçëó÷èíó â ñòåïåíè IV-IV (Fx = 3,13 %).
Ñðåäíÿÿ ñòåïåíü ãåòåðîçèãîòíîñòè ó æåðåáöîâ ñîñòàâèëà 68,40 % è
îêàçàëàñü äàæå íåñêîëüêî âûøå, ÷åì ó ìàòîê (68,10 %). Ñàìóþ âûñîêóþ
ãåòåðîçèãîòíîñòü ìèêðîñàòåëëèòíûõ ëîêóñîâ çàðåãèñòðèðîâàëè ó ìîëîäíÿêà (69,58 %), íî â öåëîì ðàçëè÷èÿ ìåæäó ãðóïïàìè áûëè íåäîñòîâåðíûìè.
Ñòåïåíü ãîìîçèãîòíîñòè ïî ìèêðîñàòåëëèòíûì ëîêóñàì (Ñà, %) â ïîëîâîçðàñòíûõ ãðóïïàõ ñ ðàçíûì êîýôôèöèåíòîì èíáðèäèíãà (F x , %) ó èññëåäîâàííûõ ëîøàäåé (Equus caballus) îðëîâñêîé ðûñèñòîé ïîðîäû:  — æåðåáöû, „ — êîáûëû, z — ìîëîäíÿê, ▲ — âñåãî ïî ïîðîäå
(n = 1194, 14 âåäóùèõ õîçÿéñòâ èç ðåãèîíîâ Ðîññèè, 2010-2013 ãîäû).
Ïî ñðàâíåíèþ ñ àóòáðåäíîé ãðóïïîé êîáûë ó èíáðåäíûõ îñîáåé
ñòåïåíü ãîìîçèãîòíîñòè ïî ìåðå óâåëè÷åíèÿ êîýôôèöèåíòà èíáðèäèíãà
38
òàêæå ìåíÿëàñü íåñóùåñòâåííî è óâåëè÷èâàëàñü òîëüêî ïðè Fx > 6,10 %
è áîëåå (ðèñ.).
 ãðóïïå ðûñèñòîãî ìîëîäíÿêà ìàêñèìàëüíàÿ ñòåïåíü ãîìîçèãîòíîñòè íå ïðåâûøàëà 62,50 %, íàáëþäàëàñü ìåíåå âûðàæåííàÿ èíäèâèäóàëüíàÿ âàðèàáåëüíîñòü ïî ñòåïåíè ãåòåðîçèãîòíîñòè ìèêðîñàòåëëèòíûõ ëîêóñîâ (òàáë. 2), ïðè ýòîì íå áûëî âûÿâëåíî ïîëíîñòüþ ãåòåðîçèãîòíûõ æèâîòíûõ. Ïðàêòè÷åñêè 40 % ëîøàäåé ýòîé ãðóïïû õàðàêòåðèçîâàëèñü ñòåïåíüþ ãîìîçèãîòíîñòè 25,00-31,25 %, à êîýôôèöèåíò èíáðèäèíãà 5,10 % è
áîëåå èìåëè âñåãî 3,80 % ìîëîäûõ ðûñàêîâ (ñì. òàáë. 1).
2. Ðàñïðåäåëåíèå ïî ñòåïåíè ãîìîçèãîòíîñòè ìèêðîñàòåëëèòíûõ ëîêóñîâ â
ãðóïïàõ ñ ðàçíûì êîýôôèöèåíòîì èíáðèäèíãà Fx ó èññëåäîâàííûõ ëîøàäåé
(Equus caballus) îðëîâñêîé ðûñèñòîé ïîðîäû (14 âåäóùèõ õîçÿéñòâ èç ðåãèîíîâ Ðîññèè, 2010-2013 ãîäû)
Còåïåíü ãîìîçèãîòíîñòè Ñà, %
0,00
6,25
12,50
18,75
25,00
31,25
37,50
43,75
50,00
56,25
62,50
68,75
75,00
81,25
Âñåãî:
÷èñëî, ãîë.
äîëÿ, %
Cañð.
±m
0
Âñåãî,
Ãðóïïà ïî êîýôôèöèåíòó èíáðèäèíãà Fx(%)
0,1-1,0 1,1-2,0 2,1-3,0 3,1-4,0 4,1-5,0 5,1-6,0 6,1-7,0 7,1 è áîëåå ãîë.
1
7
9
25
38
41
29
25
7
5
0
0
0
0
1
14
34
51
73
90
65
58
23
7
4
1
1
0
0
5
6
29
57
51
46
25
13
6
4
0
0
0
0
5
5
15
19
23
12
11
3
5
0
0
1
0
0
1
3
21
22
36
19
19
9
7
0
1
0
0
0
1
5
5
8
11
10
5
5
4
0
0
0
0
0
0
0
4
2
2
2
2
0
0
0
0
0
0
0
1
0
0
3
2
4
6
2
0
1
0
0
0
0
1
0
2
2
4
6
4
1
0
1
0
0
0
2
35
62
152
224
260
193
155
63
34
10
2
2
0
187
15,66
30,30
0,008
422
35,34
30,85
0,006
242
20,27
31,92
0,007
99
8,29
30,37
0,013
138
11,56
33,11
0,010
54
4,52
32,87
0,017
12
1,01
29,17
0,028
19
1,59
37,83
0,028
21
1,76
34,82
0,026
1194
100
31,46
0,003
Ïðîàíàëèçèðîâàâ èçìåíåíèÿ ñòåïåíè ãîìîçèãîòíîñòè ïî ìåðå ïîâûøåíèÿ êîýôôèöèåíòà èíáðèäèíãà, óñòàíîâèëè, ÷òî âî âñåõ ïîëîâîçðàñòíûõ ãðóïïàõ ëîøàäåé è ïî âñåìó ïîãîëîâüþ â öåëîì ïðîñëåæèâàåòñÿ íåáîëüøàÿ ïîëîæèòåëüíàÿ äèíàìèêà åå ðîñòà (ñì. ðèñ.). Ïðè íàëè÷èè íåêîòîðûõ êîëåáàíèé ñðåäíåé ñòåïåíè ãîìîçèãîòíîñòè ó èíáðåäíûõ ëîøàäåé
ðàçíûõ ïîëîâîçðàñòíûõ ãðóïï â ïðåäåëàõ 28,60-39,62 % ýòîò âàæíûé ïîêàçàòåëü èìåë ÿâíóþ òåíäåíöèþ ê óâåëè÷åíèþ ïðè áëèçêîðîäñòâåííûõ ñïàðèâàíèÿõ.
Êîýôôèöèåíò êîððåëÿöèè (r ) ìåæäó êîýôôèöèåíòîì èíáðèäèíãà
è ñòåïåíüþ ãîìîçèãîòíîñòè ïî ëîêóñàì ìèêðîñàòåëëèòîâ ÄÍÊ îêàçàëñÿ
ïîëîæèòåëüíûì, íî íåçíà÷èòåëüíûì ïî âåëè÷èíå (0,079).  òî æå âðåìÿ
ðåãðåññèîííûé àíàëèç âûÿâèë ñóùåñòâåííóþ çàâèñèìîñòü ìåæäó ýòèìè
ïîêàçàòåëÿìè — R y/x = 0,546.
Âî âñåõ èññëåäîâàííûõ ãðóïïàõ ëîøàäåé è â öåëîì ó ïðåäñòàâèòåëåé ïîðîäû îòìå÷àëè íåêîòîðîå çàêîíîìåðíîå ñíèæåíèå ñòåïåíè ãîìîçèãîòíîñòè ðûñàêîâ â ãðóïïå ñ êîýôôèöèåíòîì èíáðèäèíãà (Fx) 5,1-6,0 %, à
òàêæå 7,1 % è áîëåå, ÷òî ìîæåò áûòü îáóñëîâëåíî åãî êîìïëåêñíûì õàðàêòåðîì è ñâÿçàíî ñ àëãîðèòìîì âû÷èñëåíèÿ âåëè÷èíû Fx ïî ôîðìóëå Ðàéòà.
 öåëîì èçìåíåíèÿ ñòåïåíè ãîìîçèãîòíîñòè âî âñåõ ñðàâíèâàåìûõ ïîëîâîçðàñòíûõ ãðóïïàõ ëîøàäåé íîñèëè îäíîíàïðàâëåííûé õàðàêòåð, ïðè
ýòîì ðàçëè÷èÿ ìåæäó ãðóïïàìè áûëè ñòàòèñòè÷åñêè íåäîñòîâåðíûìè.
Ìîíèòîðèíã óðîâíÿ èíáðèäèíãà â ïîðîäàõ è ïîïóëÿöèÿõ ñëóæèò
âàæíîé ñîñòàâíîé ÷àñòüþ ñåëåêöèîííûõ ïðîãðàìì, ïðè ýòîì òîëüêî ïðèìåíåíèå ãåíîòèïèðîâàíèÿ æèâîòíûõ ïîçâîëÿåò ïîëó÷èòü íàäåæíóþ èí39
ôîðìàöèþ î âëèÿíèè ñåëåêöèè íà ôàêòè÷åñêóþ ñòåïåíü ãîìîçèãîòíîñòè
îñîáåé (15, 16).
Òàêèì îáðàçîì, ñòåïåíü ãîìîçèãîòíîñòè ó àóòáðåäíûõ è óìåðåííî
èíáðåäíûõ ëîøàäåé îðëîâñêîé ðûñèñòîé ïîðîäû ìîæåò âàðüèðîâàòü â
î÷åíü øèðîêîì èíòåðâàëå (îò 0 äî 75,00 %). Êîýôôèöèåíò èíáðèäèíãà,
ðàññ÷èòàííûé ïî ôîðìóëå Ðàéòà, õàðàêòåðèçóåò ëèøü âåðîÿòíîñòü ïåðåõîäà ÷àñòè ãåíîâ â ãîìîçèãîòíîå ñîñòîÿíèå, îäíàêî ïðè ýòîì íå ó÷èòûâàåòñÿ
ñòîëü ñóùåñòâåííûé ïàðàìåòð, êàê ñòåïåíü ãîìîçèãîòíîñòè ñàìîãî ïðåäêà,
ïîòîìñòâî êîòîðîãî áûëî èñïîëüçîâàíî â áëèçêîðîäñòâåííûõ ñêðåùèâàíèÿõ ñ öåëüþ ïîâûøåíèÿ ãåíåòè÷åñêîãî ñõîäñòâà ñ íèì. Ïðèìåíåíèå óìåðåííîãî êîìïëåêñíîãî èíáðèäèíãà ïðàêòè÷åñêè äàåò î÷åíü áîëüøîé ðàçáðîñ ãîìîçèãîòíîñòè è ìîæåò ðàññìàòðèâàòüñÿ êàê àóòáðåäíîå ðàçâåäåíèå.
Ïîýòîìó î÷åíü ÷àñòî ðàññ÷èòàííûé êîýôôèöèåíò èíáðèäèíãà íå ñîîòâåòñòâóåò ôàêòè÷åñêîé ãîìîçèãîòíîñòè æèâîòíûõ, äëÿ îïðåäåëåíèÿ êîòîðîé
ìîãóò áûòü èñïîëüçîâàíû ìèêðîñàòåëëèòíûå ëîêóñû è äðóãèå ÄÍÊ-ìàðêåðû, õàðàêòåðèçóþùèåñÿ âûñîêèì ïîëèìîðôèçìîì.
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ÃÍÓ Âñåðîññèéñêèé ÍÈÈ êîíåâîäñòâà
Ðîññåëüõîçàêàäåìèè,
391105 Ðîññèÿ, Ðÿçàíñêàÿ îáë., Ðûáíîâñêèé ð-í, ïîñ. Äèâîâî,
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Ïîñòóïèëà â ðåäàêöèþ
31 ìàðòà 2014 ãîäà
e-mail: [email protected], [email protected]
INBREEDING AND HOMOZYGOSITY IN ORLOV TROTTER BREED
L.A. Khrabrova, N.V. Blokhina, A.V. Ust’yantseva
All-Russian Research Institute of Horse Breeding, Russian Academy of Agricultural Sciences, pos. Divovo, Rybnovskii
Region, Ryazan Province, 391105 Russia, e-mail [email protected], [email protected]
Received March 31, 2014
doi: 10.15389/agrobiology.2014.4.35eng
Abstract
Evaluation of animals for the pedigree does not track the actual similarity of probands with
ancestors, including that when inbreeding is used. The total number of marker genes defined in
horses already reached several hundred and enables to control a significant part of the genome of
animals and use genetic information in practical breeding. Previously, we showed that Thoroughbred
horse degree of homozygosity on microsatellite loci is not directly based on the degree of inbreeding,
and the largest scale variability of this indicator is observed in the group of inbred horses. In the present work we investigated the effects of inbreeding in Orlov Trotter breed on an individual level. The
analysis was carried out of the relationship between the degree of homozygosity of 16 microsatellite
loci (AHT4, AHT5, ASB2, ASB17, ASB23, CA425, HMS1, HMS2, HMS3, HMS6, HMS7, HTG4,
HTG6, HTG7, HTG10, VHL20) and an indicator of inbreeding in 1194 individuals from 14 leading
horse-breeding studs and farms in Russia, including 19 stallions, 688 mares and 316 foals. Data on
he LEX loci 3, localized in X-chromosome, were not taken into account to obtain comparable results on the degree of homo-/heterozygosity in horses of different sex. It has been determined that
the degree of homozygosity in Orlov Trotters varies within a wide range (from 0 up to 75.00 %) and
has a normal distribution with a positive coefficient of asymmetry As = 0.186 and a small negative
excess Ex = −0.039. The most wide variability on homozygosity was indicated in the horses of mild
inbreeding (Fx = 0.1-1.0 %). The homozygosity in outbred horses also varied significantly, from 0
(100 % heterozygosity) to 56.30 %, being much more and less than an average index for the population. Previously, we showed the similar trend in the Thoroughbred horses. The average level of inbreeding of the tested horses amounted to 1.61 % by Wright`s inbreeding coefficient (Fx) in 5 generations in pedigrees and in fact remained unchanged over the past four decades. The average inbreeding coefficient in sires and mares reached 1.65 % and 1.67 %, respectively, while in young
stock was 1.45 % only. Dynamics of changes in homozygosity with increase of inbreeding coefficient
in all sex and age groups of compared horses had unidirectional trend of slowly increasing. A significant rise of the homozygosity degree was noted only at inbreeding coefficient of 6.10 % and more,
on the margin of inbred depression for economically valuable traits. The coefficient of correlation
between the degree of homozygosity of microsatellite loci and the level of inbreeding was 0.079 with
the regression coefficient of 0.546. In all groups, it has been found an evident decline in the degree
of homozygosity in Orlov Trotter horses when the inbreeding coefficient reached 5.1-6.0 % due to
complex inbreeding, with two or more ancestors used.
Keywords: homozygosity, inbreeding, microsatellite DNA, Orlov Trotter breed.
Íàó÷íûå ñîáðàíèÿ
10TH EUROPEAN CONFERENCE OF PRCISION AGRICULTURE (ECPA)
«EFFICIENT RESOURCES MANAGEMENT UNDER CHANGING GLOBAL CONDITIONS»
(12-16 èþëÿ 2015 ãîäà, Èçðàèëü)
Îðãàíèçàòîð: ARO (Agricultural Research Organization) Ìèíèñòåðñòâà ñåëüñêîãî õîçÿéñòâà è
ðàçâèòèÿ ñåëüñêîõîçÿéñòâåííîãî ïðîèçâîäñòâà Èçðàèëÿ.
Òåìàòèêà êîíôåðåíöèè: ïðåöèçèîííîå ñåëüñêîå õîçÿéñòâî êàê ñðåäñòâî ýôôåêòèâíîãî óïðàâëåíèÿ ðåñóðñàìè â óñëîâèÿõ ãëîáàëüíûõ èçìåíåíèé.
ARO — êðóïíåéøèé ñåëüñêîõîçÿéñòâåííûé èññëåäîâàòåëüñêèé öåíòð â Èçðàèëå,
îáúåäèíÿþùèé øåñòü èíñòèòóòîâ (ïî ðàñòåíèåâîäñòâó, æèâîòíîâîäñòâó, çàùèòå ðàñòåíèé,
ïî÷âåííûì, âîäíûì è ýêîëîãè÷åñêèì èññëåäîâàíèÿì, ïåðåðàáîòêå è ïèùåâûì ïðîäóêòàì), à
òàêæå ÷åòûðå îïûòíûõ ñòàíöèè â ðàçíûõ ðàéîíàõ ñòðàíû. Ãåíáàíê Èçðàèëÿ ïî ñåëüñêîõîçÿéñòâåííûì êóëüòóðàì òàêæå ðàñïîëîæåí â ýòîì öåíòðå. Íà ôîíå ðàçâèòèÿ âñåõ íàïðàâëåíèé
èññëåäîâàíèé îñîáîå âíèìàíèå óäåëÿåòñÿ ïðîáëåìàì âåäåíèÿ ñåëüñêîãî õîçÿéñòâà â àðèäíûõ
çîíàõ, ÷òî ïîçâîëÿåò Èçðàèëþ áûòü ìèðîâûì ëèäåðîì ïî âûõîäó ïðîäóêöèè â ïîäîáíûõ óñëîâèÿõ. Ïðåäëàãàåìûå òåõíîëîãèè è íàó÷íûå ðàçðàáîòêè èçðàèëüñêèõ ñïåöèàëèñòîâ ñïîñîáíû âíåñòè ñóùåñòâåííûé âêëàä â óëó÷øåíèå ñèòóàöèè íà ïðîäîâîëüñòâåííîì ðûíêå è ïðèâëåêàþò ê ñåáå ïðèñòàëüíîå âíèìàíèå ó÷åíûõ è ñïåöèàëèñòîâ âî âñåì ìèðå.
Êîíòàêòû è èíôîðìàöèÿ: http://www.ispag.org/Events/9thECPA/
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