ЭКСПЕРИМЕНТАЛЬНЫЕ ПОДХОДЫ К ДИАГНОСТИКЕ

ÑÅËÜÑÊÎÕÎÇßÉÑÒÂÅÍÍÀß ÁÈÎËÎÃÈß, 2014, ¹ 2, ñ. 20-30
ÓÄÊ 636.5:57.04:574.24:57.087
ÝÊÑÏÅÐÈÌÅÍÒÀËÜÍÛÅ ÏÎÄÕÎÄÛ Ê ÄÈÀÃÍÎÑÒÈÊÅ ÑÒÐÅÑÑÎÂ
 ÏÒÈÖÅÂÎÄÑÒÂÅ
(îáçîð)
À.Â. ÌÈÔÒÀÕÓÒÄÈÍÎÂ
Îáîáùåíû äàííûå ëèòåðàòóðû î ìåòîäàõ äèàãíîñòèêè ñòðåññîâ ó êóð â ëàáîðàòîðíûõ è
ïðîèçâîäñòâåííûõ óñëîâèÿõ.  íàñòîÿùåå âðåìÿ ðàçðàáîòàíû ìíîãî÷èñëåííûå ìåòîäû, ïîçâîëÿþùèå äèàãíîñòèðîâàòü ñîñòîÿíèå àäàïòàöèîííûõ ìåõàíèçìîâ è èíäèâèäóàëüíîé ñòðåññîâîé ÷óâñòâèòåëüíîñòè ó êóð.  òî æå âðåìÿ ñëåäóåò îòìåòèòü, ÷òî èñïîëüçîâàíèå òåõ èç ïðåäëîæåííûõ ïîäõîäîâ, êîòîðûå îñíîâàíû íà îïðåäåëåíèè áèîõèìè÷åñêèõ ìàðêåðîâ ñòðåññà â êðîâè, â ïðîèçâîäñòâåííûõ óñëîâèÿõ ïðåäñòàâëÿåò ñîáîé íåïðîñòóþ çàäà÷ó èç-çà íåîáõîäèìîñòè ïðèìåíÿòü ñïåöèàëüíîå îáîðóäîâàíèå, ïðèâëåêàòü âûñîêîêâàëèôèöèðîâàííûé ïåðñîíàë, à òàêæå âñëåäñòâèå íåîäíîçíà÷íîé èíòåðïðåòàöèè ðåçóëüòàòîâ. Âåñüìà èíôîðìàòèâíûì è ìåíåå ñëîæíûì ïðèåìîì ïðè äèàãíîñòèêå ñòðåññîâîãî ñîñòîÿíèÿ ó êóð ìîæåò áûòü èçó÷åíèå êîëè÷åñòâåííîãî ñîîòíîøåíèÿ ãåòåðîôèëîâ è ëèìôîöèòîâ â êðîâè. Âûñîêóþ äèàãíîñòè÷åñêóþ öåííîñòü â ñëó÷àå õðîíè÷åñêèõ ñòðåññîâ
èìåþò ïîêàçàòåëè îáùåãî îïåðåíèÿ.  äîïîëíåíèå ê íèì öåëåñîîáðàçíî ó÷èòûâàòü êîíöåíòðàöèþ
ñòðåññîâûõ ãîðìîíîâ â êðîâè è âðåìÿ òîíè÷åñêîé íåïîäâèæíîñòè êóð. Îäíàêî è â ýòèõ ñëó÷àÿõ
ïðîöåäóðû, êîòîðûì ïîäâåðãàåòñÿ ïòèöà, ñàìè ïî ñåáå ñëóæàò ôàêòîðàìè ñòðåññà, òîãäà êàê îäíî
èç âàæíåéøèõ óñëîâèé äîñòîâåðíîñòè ðåçóëüòàòîâ ïðè åãî äèàãíîñòèêå — ìèíèìàëüíîå âîçäåéñòâèå íà îáúåêò èññëåäîâàíèÿ. Äëÿ ýòèõ öåëåé íàìè ðàçðàáîòàí ñïîñîá, ïîçâîëÿþùèé äèàãíîñòèðîâàòü ñîñòîÿíèå ñòðåññà ó êóð ðîäèòåëüñêîãî ñòàäà ìÿñíîãî íàïðàâëåíèÿ ïðîäóêòèâíîñòè ñ ïîìîùüþ îïðåäåëåíèÿ êîðòèêîñòåðîíà â ïðîáàõ ïîìåòà. Ïðåäëàãàåìûé ñïîñîá ìîæåò áûòü èñïîëüçîâàí ïðè îöåíêå öåëåñîîáðàçíîñòè ïðèìåíåíèÿ ñõåì àíòèñòðåññîâîé òåðàïèè, äëÿ îïðåäåëåíèÿ îïòèìàëüíîé ïëîòíîñòè êîìïëåêòîâàíèÿ ïðîèçâîäñòâåííûõ ïëîùàäåé, äëÿ àíàëèçà ñòåïåíè âîçäåéñòâèÿ íà îðãàíèçì êóð òåõíîëîãè÷åñêèõ ôàêòîðîâ è âåòåðèíàðíûõ îáðàáîòîê.
Êëþ÷åâûå ñëîâà: ñòðåññ, êóðû, äèàãíîñòèêà ñòðåññîâ.
Ñòðåññû îòíîñÿòñÿ ê àêòóàëüíûì ïðîáëåìàì ïòèöåâîäñòâà (1).  ñîîòâåòñòâèè ñî ñëîæèâøåéñÿ êîíöåïöèåé, ñòðåññîðû ÷åðåç íåðâíóþ è ýíäîêðèííóþ ñèñòåìó âûçûâàþò ìîðôîëîãè÷åñêèå è ôóíêöèîíàëüíûå èçìåíåíèÿ â îðãàíàõ è òêàíÿõ, à òàêæå óñèëåííûé ñèíòåç è ñåêðåöèþ ãîðìîíîâ
àäàïòàöèè, óñèëèâàþùèõ ðåçèñòåíòíîñòü îðãàíèçìà ê âîçäåéñòâèþ íåãàòèâíûõ ôàêòîðîâ âíåøíåé ñðåäû è ñïîñîáñòâóþùèõ âîññòàíîâëåíèþ ïðîèçîøåäøèõ â íåì îáðàòèìûõ íàðóøåíèé (2-4).
Ñòðåññû íà÷èíàþòñÿ åùå ñ èíêóáàòîðà: ðàííÿÿ âûáîðêà èëè çàäåðæêà öûïëÿò â èíêóáàòîðå — ýòî ñèëüíûé ñòðåññ, ñïîñîáíûé ñóùåñòâåííî ñíèçèòü áóäóùóþ ïðîäóêòèâíîñòü ïòèöû (5). Ðàçëè÷íûå ñòðåññû ìîãóò íàðóøèòü ïðîöåññ ñîçðåâàíèÿ òîíêîãî êèøå÷íèêà â ïåðâûå íåñêîëüêî ñóòîê æèçíè öûïëåíêà (6, 7), âûðàæåííîå ñòðåññîâîå âîçäåéñòâèå îêàçûâàþò âàêöèíàöèÿ â èíêóáàòîðèè, òðàíñïîðòèðîâêà öûïëÿò â êîðïóñà,
ïîñàäêà ïðè îòñóòñòâèè âûðîâíåííîé ïîäñòèëêè, îïòèìàëüíîé òåìïåðàòóðû è âåíòèëÿöèè (5). Â äàëüíåéøåì äîáàâëÿþòñÿ êîðìîâûå ñòðåññû,
ñâÿçàííûå íå òîëüêî ñ ïåðåõîäîì öûïëåíêà îò æåëòêîâîãî ïèòàíèÿ ê ñóõîìó êîðìó, îòëè÷àþùåìóñÿ ïî ñîñòàâó îò æåëòêà, íî è ñ íåñáàëàíñèðîâàííîñòüþ ðàöèîíà ïî ïèòàòåëüíûì è áèîëîãè÷åñêè àêòèâíûì âåùåñòâàì.  ïðîöåññå ðîñòà è ðàçâèòèÿ íåèçáåæíû òåõíîëîãè÷åñêèå ñòðåññû,
ñâÿçàííûå ñ âàêöèíàöèÿìè, â ëåòíèå ìåñÿöû îñîáóþ ðîëü èãðàþò òåïëîâûå ñòðåññû, íàëè÷èå ìèêîòîêñèíîâ â êîðìàõ — îäèí èç îñíîâíûõ êîðìîâûõ ñòðåññîâ (8, 9).  òåõíîëîãèè ïðîèçâîäñòâà ïèùåâîãî è èíêóáàöèîííîãî ÿéöà âàæíåéøèì ñòðåññîì ñîïðîâîæäàåòñÿ âûõîä ïòèöû íà ïèê
ÿéöåíîñêîñòè (10). Îäíà èç ãëàâíûõ ïðîáëåì, ñâÿçàííûõ ñî ñòðåññàìè, —
ïîâûøåííàÿ ïîòðåáíîñòü â ïèòàòåëüíûõ è áèîëîãè÷åñêè àêòèâíûõ âåùåñòâàõ, íåîáõîäèìûõ äëÿ áîðüáû ñî ñòðåññàìè, â òî âðåìÿ êàê èõ ïîñòóï20
ëåíèå ñ êîðìîì óìåíüøàåòñÿ, ïîñêîëüêó â òàêîì ñîñòîÿíèè ïòèöà õóæå
ïîåäàåò êîðì (1, 11, 12).
Ìåòîäû äèàãíîñòèêà ñòðåññîâ ïòèöû âàæíû äëÿ ðåøåíèÿ êàê òåîðåòè÷åñêèõ, òàê è ïðàêòè÷åñêèõ çàäà÷. Ê îñíîâíûì èç íèõ îòíîñèòñÿ èçó÷åíèå ñòðåññîâ ó êóð â óñëîâèÿõ ïðîìûøëåííûõ òåõíîëîãèé è â ëàáîðàòîðíûõ ôèçèîëîãè÷åñêèõ îïûòàõ, à òàêæå îöåíêà ñòåïåíè âëèÿíèÿ âíåøíèõ ôàêòîðîâ íà ôèçèîëîãè÷åñêîå ñîñòîÿíèå ïòèöû. Ê ÷èñëó ïðàêòè÷åñêèõ çàäà÷ ìîæíî îòíåñòè êîíòðîëü çà ôèçèîëîãè÷åñêèì ñîñòîÿíèåì ïòèöû ïðè ðàçðàáîòêå íîâûõ ìåòîäîâ ïðîôèëàêòèêè ñòðåññîâ, ïðè ñîçäàíèè è
âûáîðå îïòèìàëüíûõ ñèñòåì ñîäåðæàíèÿ è êîðìëåíèÿ (13, 14).
 íàñòîÿùåå âðåìÿ åäèíûé ïîäõîä ê äèàãíîñòèêå ñòðåññîâ â ïòèöåâîäñòâå îòñóòñòâóåò (15). Ïðåäëàãàåìûå ìåòîäû èìåþò ñâîè ïðåèìóùåñòâà
è íåäîñòàòêè, ÷òî òðåáóåò îò èññëåäîâàòåëÿ î÷åíü îñòîðîæíîãî âûáîðà ìàðêåðîâ ñòðåññà (16). Ïðè èñïîëüçîâàíèè ëþáîãî ïîäõîäà ê èçó÷åíèþ ñòðåññîâ è ñòðåññîâîé ÷óâñòâèòåëüíîñòè îðãàíèçìà íåîáõîäèìî â îáÿçàòåëüíîì
ïîðÿäêå ó÷èòûâàòü âîçðàñò ïòèöû. Ïî äàííûì Â.È. Ôèñèíèíà ñ ñîàâò.
(13), àäàïòàöèîííî-êîìïåíñàòîðíûå ìåõàíèçìû ó öûïëÿò â ïðîöåññå ðîñòà
è ðàçâèòèÿ ïðåòåðïåâàþò ôàçîâûå èçìåíåíèÿ. Ïåðâàÿ ôàçà (âûñîêàÿ ÷óâñòâèòåëüíîñòü) îòìå÷àåòñÿ ó öûïëÿò â ðàííåì âîçðàñòå (îò 1 äî 90 ñóò), â
âîçðàñòå 90-120 ñóò íàñòóïàåò âòîðàÿ ôàçà (îòíîñèòåëüíàÿ ðåçèñòåíòíîñòü)
è òðåòüÿ ôàçà (ïîâòîðíîå ïîâûøåíèå ÷óâñòâèòåëüíîñòè) íà÷èíàåòñÿ ó öûïëÿò ñî 140-ñóòî÷íîãî âîçðàñòà (13).
Ê îáùèì ïîêàçàòåëÿì ñòðåññà, íå èìåþùèì êîëè÷åñòâåííîãî âûðàæåíèÿ, ìîæíî îòíåñòè àòðîôèþ òèìóñà è áóðñû ó ìîëîäîé ïòèöû, óâåëè÷åíèå ìàññû ïåðåäíåé äîëè ãèïîôèçà è íàäïî÷å÷íèêîâ, èñòîùåíèå õîëåñòåðèíà â íàäïî÷å÷íèêàõ, ïîâûøåíèå êîíöåíòðàöèè êîðòèêîñòåðîíà,
èíñóëèíà è ãëþêàãîíà â ïëàçìå êðîâè, óñèëåííûé ðàñõîä ãëþêîçû, ñíèæåíèå ïðèðîñòîâ æèâîé ìàññû è àòðîôèþ ìûøö, âûäåëåíèå â êðîâü ìàðêåðîâ îñòðûõ âîñïàëèòåëüíûõ ïðîöåññîâ, öèòîêèíîâ, íàðóøåíèå ðîñòà
õðÿùåé è êîñòåé, àíîðåêñèþ, ïîâûøåííóþ òåìïåðàòóðó òåëà. Èçáûòî÷íîå
îòëîæåíèå æèðà â áðþøíîé ïîëîñòè è àñöèò ñ÷èòàþò ìàðêåðíûìè ïðèçíàêàìè ñòðåññà ïðè èíòåíñèâíîì îòêîðìå öûïëÿò-áðîéëåðîâ (17).
Ïðîäóêòèâíîñòü êóð íå âñåãäà ñëóæèò îáúåêòèâíûì êðèòåðèåì íàëè÷èÿ ñòðåññà, òàê êàê êîìïåíñàòîðíûå ìåõàíèçìû îðãàíèçìà ïîçâîëÿþò
íåêîòîðîå âðåìÿ ïîääåðæèâàòü ãîìåîñòàç, ñîõðàíÿÿ îïðåäåëåííîå ñîñòîÿíèå çäîðîâüÿ è ïðîäóêòèâíîñòè, íî ïî ìåðå èñòîùåíèÿ ðåçåðâîâ àäàïòàöèîííûõ ñèñòåì íàáëþäàåòñÿ ñíèæåíèå ðåàêòèâíîñòè è ðåçèñòåíòíîñòè è
â ðåçóëüòàòå ïðîäóêòèâíîñòü ðåçêî ïàäàåò, ðàçâèâàþòñÿ áîëåçíè è ïðîèñõîäèò ìàññîâàÿ ãèáåëü êóð. Âûñîêàÿ ñìåðòíîñòü ìîæåò áûòü âàæíûì ïðèçíàêîì îñòðûõ è õðîíè÷åñêèõ ñòðåññîâ â ïòèöåâîäñòâå (18).
Óáåäèòåëüíî äîêàçàíà ýôôåêòèâíîñòü ìåòîäà, îñíîâàííîãî íà èçó÷åíèè ãîðìîíàëüíîãî ñòàòóñà, ìîðôîëîãè÷åñêèõ ïîêàçàòåëåé êðîâè, áàçîâûõ áèîõèìè÷åñêèõ êîíñòàíò îðãàíèçìà è ïîêàçàòåëåé ðåçèñòåíòíîñòè äëÿ
îïðåäåëåíèÿ ñòðåññîâîãî ñîñòîÿíèÿ êóð, è èñïîëüçîâàíèÿ îïèñàííîãî ïîäõîäà äëÿ îïðåäåëåíèÿ âîçäåéñòâèÿ òåõíîëîãè÷åñêèõ ôàêòîðîâ íà îðãàíèçì
ïòèö, à òàêæå ñîîòâåòñòâèÿ òåõíîëîãè÷åñêèõ ïàðàìåòðîâ ôèçèîëîãè÷åñêèì
ïîòðåáíîñòÿì êóð (2, 13, 14, 19-21).
Ñðåäè ñïîñîáîâ îïðåäåëåíèÿ êîëè÷åñòâåííûõ ïîêàçàòåëåé ñòðåññîâîãî ñîñòîÿíèÿ ó êóð ìîæíî âûäåëèòü ïðÿìûå, êîòîðûå îñíîâàíû íà
îöåíêå ñîäåðæàíèÿ ñòðåññîâûõ ãîðìîíîâ â êðîâè. Èçâåñòíû ìíîãî÷èñëåííûå èññëåäîâàíèÿ, äîêàçûâàþùèå ýôôåêòèâíîñòü íåïîñðåäñòâåííîãî èçìåðåíèÿ êîëè÷åñòâà îñíîâíûõ ãîðìîíîâ â ñèñòåìå ãèïîòàëàìóñ—ãèïî21
ôèç—íàäïî÷å÷íèêè, à òàêæå èõ ìåòàáîëèòîâ. Ïðè èíòåðïðåòàöèè ðåçóëüòàòîâ ïîäîáíûõ àíàëèçîâ íåîáõîäèìî ó÷èòûâàòü, ÷òî êîëè÷åñòâî ãîðìîíîâ
çàâèñèò îò ïðèðîäû è ñèëû ðàçäðàæèòåëÿ, äëèòåëüíîñòè åãî âîçäåéñòâèÿ,
âîçðàñòà è ãåíîòèïà ïòèöû (2).
Ïî äàííûì Â.È. Ôèñèíèíà è Í.À. Êðàâ÷åíêî (14), ó âçðîñëûõ êóð
ñîäåðæàíèå êàòåõîëàìèíîâ â íàäïî÷å÷íèêàõ è êðîâè è êîðòèêîñòåðîèäîâ
â êðîâè ìåíåå ïîäâåðæåíî èçìåíåíèÿì ïðè äåéñòâèè ðàçäðàæèòåëåé, ÷åì
ó öûïëÿò. Óêàçàííûé ôàêò îáúÿñíÿåòñÿ òåì, ÷òî ó âçðîñëîé ïòèöû ïðè
àäàïòàöèè ê íîâûì óñëîâèÿì îòìå÷àåòñÿ ìåíüøèé ðàñõîä êàòåõîëàìèíîâ è
êîðòèêîñòåðîèäîâ, îïèñàííîå ÿâëåíèå íå ñâÿçàíî ñ èñòîùåíèåì àäðåíîêîðòèêàëüíûõ ñòðóêòóð è ìîæåò áûòü ñëåäñòâèåì èçìåíåíèÿ ÷óâñòâèòåëüíîñòè ðåöåïòîðîâ. Ïðè ýòîì âàæíîå çíà÷åíèå èìååò èñõîäíîå ñîñòîÿíèå
íåéðîýíäîêðèííîé ñèñòåìû, êîòîðîå íåîáõîäèìî ó÷èòûâàòü ïðè àíàëèçå
ðåãèñòðèðóåìûõ äàííûõ (14).
Ïîïûòêè äèàãíîñòèðîâàòü ñòðåññ ïîñðåäñòâîì îïðåäåëåíèÿ êîëè÷åñòâà àäðåíîêîðòèêîòðîïíîãî ãîðìîíà (ÀÊÒÃ) â êðîâè íå íàøëè øèðîêîãî
ïðàêòè÷åñêîãî ïðèìåíåíèÿ, è ýòîò ïðèåì èñïîëüçóåòñÿ òîëüêî â êîìïëåêñíûõ ëàáîðàòîðíûõ èññëåäîâàíèÿõ (22).
Îïðåäåëåíèå äîôàìèíà â êðîâè — öåííûé ìåòîä äèàãíîñòèêè ñòðåññîâ è ñòðåññîâîé ÷óâñòâèòåëüíîñòè (23, 24). Â ìîçãå ïòèö, ïðîÿâëÿþùèõ
àãðåññèþ, îáíàðóæåíî áîëåå âûñîêîå ñîäåðæàíèå äîôàìèíà, ÷åì ó îñîáåé,
íå ñêëîííûõ ê äèâèàíòíîìó ïîâåäåíèþ (25).
Àíàëèç ñîäåðæàíèÿ àäðåíàëèíà è íîðàäðåíàëèíà â êðîâè â ïòèöåâîäñòâå èñïîëüçóåòñÿ äëÿ âûÿâëåíèÿ îñòðûõ òåõíîëîãè÷åñêèõ ñòðåññîâ, âûçâàííûõ ôàêòîðàìè, êîòîðûå õàðàêòåðèçóþòñÿ âíåçàïíîñòüþ è âûçûâàþò
âûñîêóþ ñìåðòíîñòü, à òàêæå ïðè äèàãíîñòèêå ïðè÷èí êàííèáàëèçìà (26,
27). Êîíöåíòðàöèÿ àäðåíàëèíà â ñûâîðîòêå êðîâè èñïîëüçóåòñÿ â êà÷åñòâå
ñåëåêöèîííîãî ïðèçíàêà ïðè ñîçäàíèè ëèíèé ïòèöû ñ âûñîêîé è íèçêîé
èíäóöèáåëüíîñòüþ ýòîãî ãîðìîíà, îïðåäåëÿþùåé ðàçíóþ ÷óâñòâèòåëüíîñòü
ê ñòðåññàì (28).
Èçìåíåíèå ñîäåðæàíèÿ êàòåõîëàìèíîâ è ýñòðîãåíîâ â òêàíÿõ íàäïî÷å÷íèêîâ ó êóð çàâèñèò îò âîçðàñòà, ïåðèîäà ðîñòà, ðàçâèòèÿ è ôèçèîëîãè÷åñêîãî ñîñòîÿíèÿ è èìååò âàæíîå ïðîãíîñòè÷åñêîå çíà÷åíèå ïðè îïðåäåëåíèè îïòèìàëüíûõ òåõíîëîãè÷åñêèõ ïàðàìåòðîâ äëÿ âûðàùèâàíèÿ è
ñîäåðæàíèÿ ïòèöû, à òàêæå ñîâåðøåíñòâîâàíèè íîðì ïèòàòåëüíûõ è áèîëîãè÷åñêè àêòèâíûõ âåùåñòâ, îáåñïå÷èâàþùèõ ïîòðåáíîñòè ìîëîäíÿêà è
êóð-íåñóøåê. Â.È. Ôèñèíèíûì è Í.À. Êðàâ÷åíêî (14) äîêàçàíî, ÷òî ñóùåñòâóþò òðè îñíîâíûå ôàçû, êîãäà ñîäåðæàíèå êàòåõîëàìèíîâ â íàäïî÷å÷íèêàõ ïîäâåðæåíî èçìåíåíèÿì. Ïåðâàÿ ñîâïàäàåò ñ ïåðèîäîì îáùåãî ðîñòà è ðàçâèòèÿ îðãàíèçìà, äëÿ íåå õàðàêòåðåí êîìïåíñàòîðíûé ñèíòåç êàòåõîëàìèíîâ, ïðè ýòîì ñîäåðæàíèå àäðåíàëèíà âûøå, ÷åì íîðàäðåíàëèíà.
Âî âòîðóþ ôàçó êîëè÷åñòâî ãîðìîíîâ â íàäïî÷å÷íèêàõ ñíèæàåòñÿ â ðåçóëüòàòå óñèëåíèÿ ñåêðåöèè àäðåíàëèíà. Ñîîòíîøåíèå ìåæäó àäðåíàëèíîì
è íîðàäðåíàëèíîì âûðàâíèâàåòñÿ. Âòîðàÿ ôàçà õàðàêòåðíà äëÿ ïåðèîäà
ïîäãîòîâêè è íà÷àëà ÿéöåêëàäêè ó ïòèöû.  òðåòüþ ôàçó ó âçðîñëûõ êóð
îñíîâíûì ãîðìîíîì â íàäïî÷å÷íèêàõ ñòàíîâèòñÿ íîðàäðåíàëèí (14). Óêàçàííûå ôèçèîëîãè÷åñêèå îñîáåííîñòè àêòèâàöèè àäðåíåðãè÷åñêîé ñèñòåìû îðãàíèçìà äîëæíû îáÿçàòåëüíî ó÷èòûâàòüñÿ ïðè èçó÷åíèè ñòðåññîâ è
ñòðåññîâîé ÷óâñòâèòåëüíîñòè êóð â óñëîâèÿõ ïðîìûøëåííîãî ñîäåðæàíèÿ.
Ñåðîòîíèí â îðãàíèçìå êóð âûïîëíÿåò ðàçíîîáðàçíûå ôèçèîëîãè÷åñêèå ôóíêöèè è âëèÿåò íà àäàïòàöèþ ê ôàêòîðàì ñîöèàëüíîé ñðåäû,
ñåêñóàëüíîå ïîâåäåíèå, ýìîöèè è âîñïðîèçâîäñòâî. Îïðåäåëåíèå â êðîâè
22
è ìîçãå ìåòàáîëèòîâ ñåðîòîíèíà, â îñíîâíîì 5-ãèäðîêñèèíäîëóêñóñíîé
êèñëîòû, èñïîëüçóåòñÿ äëÿ àíàëèçà ðàçëè÷íûõ ãåíåòè÷åñêèõ ëèíèé ïî ñòåïåíè àãðåññèè è ñêëîííîñòè ê äèâèàíòíîìó ïîâåäåíèþ (29, 30).
Îïðåäåëåíèå â êðîâè ãëþêîêîðòèêîèäíûõ ãîðìîíîâ — îáùåïðèçíàííûé ìåòîä ïîäòâåðæäåíèÿ àêòèâàöèè ñòðåññ-ðåàëèçóþùèõ ìåõàíèçìîâ
îðãàíèçìà. Â.È. Ôèñèíèíûì (21) èçó÷åí ãèäðîêîðòèçîí â êà÷åñòâå ìàðêåðà ñòðåññîâîãî ñîñòîÿíèÿ ó êóð. Ïî äàííûì ìíîãî÷èñëåííûõ èññëåäîâàòåëåé, êîðòèêîñòåðîí îòíîñèòñÿ ê íàèáîëåå ÷àñòî àíàëèçèðóåìûì ãîðìîíàì
ïðè äèàãíîñòèêå ñòðåññîâîãî ñîñòîÿíèÿ êóð, ïîñêîëüêó ýòî îñíîâíîé ãëþêîêîðòèêîèäíûé ãîðìîí ó ïòèöû (31-34). Ïðè îïðåäåëåíèè ãëþêîêîðòèêîèäíûõ ãîðìîíîâ ó ìîëîäíÿêà êóð äëÿ îöåíêè àêòèâàöèè ñòðåññ-ðåàëèçóþùèõ ìåõàíèçìîâ íåîáõîäèìî ó÷èòûâàòü, ÷òî ÷åì ñòàðøå îñîáè, ïîäâåðãøèåñÿ âîçäåéñòâèþ ñòðåññ-ôàêòîðà, òåì ìåíåå âûðàæåíî èçìåíåíèå êîëè÷åñòâà ãëþêîêîðòèêîèäîâ â êðîâè. Ýòè ðàçëè÷èÿ, ïî-âèäèìîìó, ìîæíî
ðàññìàòðèâàòü êàê ðåçóëüòàò ïîñòåïåííî ïðîãðåññèðóþùèõ ôèçèîëîãî-áèîõèìè÷åñêèõ èçìåíåíèé ãèïîòàëàìî-ãèïîôèçàðíî-íàäïî÷å÷íèêîâîé ñèñòåìû, îáóñëîâëèâàþùèõ áîëåå áûñòðîå âîçâðàùåíèå ñîäåðæàíèÿ ãëþêîêîðòèêîèäíûõ ãîðìîíîâ ê íîðìå (2).
Ê ãëàâíûì ïðîáëåìàì ïðè äèàãíîñòèêå ñòðåññîâ íà îñíîâå îïðåäåëåíèÿ êîíöåíòðàöèè ñòðåññîâûõ ãîðìîíîâ â êðîâè îòíîñèòñÿ àêòèâàöèÿ
ñòðåññ-ðåàëèçóþùèõ ìåõàíèçìîâ ïðè îòáîðå ïðîá êðîâè, ÷òî âåäåò ê èñêàæåíèþ ðåçóëüòàòîâ è íå ïîçâîëÿåò ïîëó÷àòü äîñòîâåðíûå äàííûå. J. Rushen
(35) è P. Moneva ñ ñîàâò. (36) óêàçûâàþò íà íåîäíîçíà÷íîñòü èíòåðïðåòàöèè ïîëó÷àåìûõ ðåçóëüòàòîâ è îòñóòñòâèå øèðîêîé íîðìàòèâíîé áàçû äëÿ
ñîïîñòàâëåíèÿ äàííûõ ïðè îïðåäåëåíèè ñòðåññîâûõ ãîðìîíîâ â êðîâè.
Íàøè èññëåäîâàíèÿ íåñêîëüêèõ ãðóïï ïðèçíàêîâ, èìåþùèõ ìàðêåðíîå çíà÷åíèå ïðè ñòðåññàõ, âûïîëíåííûå ñ èñïîëüçîâàíèåì äèñêðèìèíàíòíîãî àíàëèçà, ïîêàçàëè, ÷òî êîíöåíòðàöèÿ àäðåíàëèíà è êîðòèêîñòåðîíà ó êóð â ñîñòîÿíèè îòíîñèòåëüíîãî ïîêîÿ è ïîñëå àêòèâàöèè ñòðåññðåàëèçóþùèõ ìåõàíèçìîâ ìîæåò áûòü èñïîëüçîâàíà ëèøü â êà÷åñòâå óòî÷íÿþùåãî ïîêàçàòåëÿ ïðè èçó÷åíèè ñòðåññîâ è ñòðåññîâîé ÷óâñòâèòåëüíîñòè (37). Ñ öåëüþ èñêëþ÷åíèÿ äîïîëíèòåëüíîãî ñòðåññèðîâàíèÿ êóð ïðè
îòáîðå êðîâè è ïîëó÷åíèÿ íàèáîëåå îáúåêòèâíûõ äàííûõ ðåêîìåíäóåòñÿ
ïðèìåíÿòü êàòåòåðèçàöèþ êðóïíûõ êðîâåíîñíûõ ñîñóäîâ — ïîäêðûëüöîâîé âåíû, ïëå÷åâîé âåíû èëè àðòåðèè (36, 38, 39).
Ðàçëè÷íûå ìîäèôèêàöèè ìåòîäà îïðåäåëåíèÿ ñòðåññîâûõ ãîðìîíîâ
â êðîâè øèðîêî èñïîëüçóþòñÿ â ýêñïåðèìåíòàëüíûõ èññëåäîâàíèÿõ, íî â
óñëîâèÿõ ïðîìûøëåííîãî ïðîèçâîäñòâà èõ ïðèìåíåíèå çàòðóäíåíî èç-çà
òðóäîåìêîñòè îïðåäåëåíèÿ, ñëîæíîñòè èíòåðïðåòàöèè ðåçóëüòàòîâ è àêòèâàöèè ñòðåññ-ðåàëèçóþùèõ àäàïòàöèîííûõ ìåõàíèçìîâ îðãàíèçìà âñëåäñòâèå ïðîèçâîäèìûõ ìàíèïóëÿöèé è áîëåçíåííîñòè ïðîöåäóðû îòáîðà êðîâè
äëÿ ïòèöû (22, 40).
Ïðè èçó÷åíèè ñòðåññà è ñòðåññîâîé ÷óâñòâèòåëüíîñòè ó êóð øèðîêî
èñïîëüçóåòñÿ îäèí èç êîñâåííûõ ôèçèîëîãè÷åñêèõ òåñòîâ — äèíàìèêà ñîîòíîøåíèÿ ìåæäó ãåòåðîôèëàìè è ëèìôîöèòàìè (Ã/Ë) â ïåðèôåðè÷åñêîé
êðîâè. Óñòàíîâëåíî, ÷òî ïðè ðàçâèòèè ñîñòîÿíèÿ ñòðåññà ýòîò ïîêàçàòåëü
óâåëè÷èâàåòñÿ çà ñ÷åò àáîðòèâíîãî âûáðîñà íåçðåëûõ êëåòîê ãåòåðîôèëîâ
èç êîñòíîãî ìîçãà â êðîâÿíîå ðóñëî è ìèãðàöèè ëèìôîöèòîâ èç ïîñëåäíåãî â òêàíè. Èçìåíåíèÿ âåëè÷èíû ñîîòíîøåíèÿ Ã/Ë êîððåëèðóþò ñî ñäâèãîì êîíöåíòðàöèè êîðòèêîñòåðîíà â êðîâè ïòèöû è ïðîïîðöèîíàëüíû
ñòåïåíè äåéñòâèÿ ñòðåññ-ôàêòîðîâ ðàçëè÷íîé ïðèðîäû (16, 35, 41, 42). Ñîîòíîøåíèå Ã/Ë ìîæåò õàðàêòåðèçîâàòü òðè óðîâíÿ ôèçèîëîãè÷åñêîãî ñî23
ñòîÿíèÿ êóð: íèçêèé ñîîòâåòñòâóåò çíà÷åíèþ 0,2, ñðåäíèé — çíà÷åíèþ 0,5
è âûñîêèé — ìîæåò áûòü áîëåå 0,8 (41). Ê íåäîñòàòêàì òàêîãî ìåòîäà ñëåäóåò îòíåñòè òî, ÷òî ïðè âîçäåéñòâèè ñâåðõïîðîãîâûõ ðàçäðàæèòåëåé ñîîòíîøåíèå Ã/Ë íå âñåãäà èçìåíÿåòñÿ àäåêâàòíî ôèçèîëîãè÷åñêîìó ñîñòîÿíèþ ïòèöû, ïîýòîìó íå ìîæåò ïðèíèìàòüñÿ çà òî÷íûé ïîêàçàòåëü ñòðåññîâîãî âîçäåéñòâèÿ (36, 43, 44). Êðîìå òîãî, ïîäñ÷åò ôîðìåííûõ ýëåìåíòîâ â ìàçêàõ êðîâè ïòèöû — ïðîöåññ âåñüìà òðóäîåìêèé è òðåáóþùèé âûñîêîé êâàëèôèêàöèè ïåðñîíàëà. J. Post ñ ñîàâò. (45) óêàçûâàþò íà âîçìîæíîñòü èñïîëüçîâàíèÿ è òî÷íîñòü àâòîìàòè÷åñêèõ ãåìàòîëîãè÷åñêèõ àíàëèçàòîðîâ ïðè ïîäñ÷åòå ñîîòíîøåíèÿ Ã/Ë.
 êà÷åñòâå óòî÷íÿþùèõ ïîêàçàòåëåé ïðè äèàãíîñòèêå ñòðåññîâîãî
ñîñòîÿíèÿ êóð ïðèìåíÿþò îïðåäåëåíèå ÷èñëà ýîçèíîôèëîâ è ìîíîöèòîâ
â ìàçêàõ êðîâè. Â èññëåäîâàíèÿõ Â.È. Ôèñèíèíà ñ ñîàâò. (19), à òàêæå
M.H. Maxwell è G.M. Robertson (46) îïèñàíà ýîçèíîïåíèÿ, ñîïóòñòâóþùàÿ
ñòðåññîâîé ðåàêöèè ó êóð, ïðè ýòîì êîëè÷åñòâî ýîçèíîôèëîâ â îðãàíàõ è
òêàíÿõ óâåëè÷èâàåòñÿ, óêàçûâàÿ íà òî, ÷òî ýîçèíîôèëû â ïåðèîä ðàçâèòèÿ
ñòðåññà ïîêèäàþò ñîñóäèñòîå ðóñëî. Ïî äàííûì O. Altan ñ ñîàâò. (47), ñíèæåíèå ÷èñëà ìîíîöèòîâ è ýîçèíîôèëîâ ñâèäåòåëüñòâóåò îá îñòðîì ñòðåññå,
îäíàêî ýòîò ìåõàíèçì ðåàëèçóåòñÿ íå âñåãäà è íå ìîæåò íàäåæíî ìàðêèðîâàòü ñòðåññ (46).
Èçâåñòåí ïðèåì äèàãíîñòèêè ñòðåññà ïî ìèãðàöèîííîé àêòèâíîñòè
ëåéêîöèòîâ ïåðèôåðè÷åñêîé êðîâè íà îñíîâå êàïèëëÿðíîãî ìåòîäà â ìîäèôèêàöèè Þ.È. Çàáóäñêîãî ñ ñîàâò. (48). Ïî ìíåíèþ àâòîðîâ, ýòîò ïîäõîä èìååò ðÿä ïðåèìóùåñòâ ïî ñðàâíåíèþ ñ îïðåäåëåíèåì ñîîòíîøåíèÿ
Ã/Ë. Ñïîñîá ìåíåå òðóäîåìêèé, òàê êàê íå ïðåäïîëàãàåòñÿ ïðèìåíÿòü
èììåðñèîííóþ ìèêðîñêîïèþ, è áîëåå äîñòóïíûé äëÿ ïåðñîíàëà âåòåðèíàðíûõ ëàáîðàòîðèé ïòèöåôàáðèê, âëàäåþùåãî íàâûêàìè ïîñòàíîâêè ñåðîëîãè÷åñêèõ ðåàêöèé. Íåäîñòàòîê îïèñàííîãî ïîäõîäà â òîì, ÷òî îòáîð
êðîâè ïðåäñòàâëÿåò ñîáîé äîïîëíèòåëüíîå ñòðåññîâîå âîçäåéñòâèå, âëèÿþùåå íà ðåçóëüòàòû àíàëèçà (48).
 èññëåäîâàíèÿõ Â.È. Ôèñèíèíà ñ ñîàâò. (19) îáîñíîâàíà äèàãíîñòè÷åñêàÿ öåííîñòü ðåàêöèè îñåäàíèÿ ýðèòðîöèòîâ ïðè îïðåäåëåíèè ñòðåññîâîãî ñîñòîÿíèÿ ó êóð.  ïåðèîä ðàçâèòèÿ ñòðåññîâ ýòîò ïîêàçàòåëü ìîæåò
óâåëè÷èâàòüñÿ â íåñêîëüêî ðàç (19).
Ê ïîñëåäñòâèÿì ñòðåññîâ îòíîñèòñÿ óãíåòåíèå íåñïåöèôè÷åñêîé
ðåçèñòåíòíîñòè è èììóííîé ñèñòåìû. Â ðàáîòàõ Â.È. Ôèñèíèíà ñ ñîàâò.
(19) â êà÷åñòâå ìàðêåðà ñòðåññà èñïîëüçîâàí òèòð ëèçîöèìà, êîððåëèðóþùèé ñ êîëè÷åñòâîì öèòîêèíîâ è ãëþêîêîðòèêîèäíûõ ãîðìîíîâ. Èçâåñòíî, ÷òî öèòîêèíû èãðàþò âàæíóþ ðîëü âî âçàèìîäåéñòâèè èììóííîé è
ýíäîêðèííîé ñèñòåìû ïðè ðåàëèçàöèè ñòðåññîâ â îðãàíèçìå (49). Êëþ÷åâûì öèòîêèíîì, ñòèìóëèðóþùèì ñòðåññîâûå ðåàêöèè, ñ÷èòàåòñÿ ÈË-1β,
ñåêðåòèðóåìûé àêòèâèðîâàííûìè ìàêðîôàãàìè, ýíäîòåëèîöèòàìè, êåðàòèíîöèòàìè è äð. ÈË-1β äåéñòâóåò íà íåéòðîôèëû, ñïîñîáñòâóÿ õåìîòàêñèñó, àêòèâàöèè ìåòàáîëèçìà, âûõîäó èç êëåòîê ëèçîöèìà è ëàêòîôåððèíà. Ãëàâíûì ìåõàíèçìîì ÈË-1β-çàâèñèìîé èíäóêöèè ñòðåññà ñëóæèò àêòèâàöèÿ ñèíòåçà è ñåêðåöèè êîðòèêîòðîïèí-ðèëèçèíã-ôàêòîðà èç íåðâíûõ îêîí÷àíèé (20). S.-Y. Kang ñ ñîàâò. (50) îáíàðóæèëè ýôôåêòû ñòðåññà, âûðàçèâøèåñÿ â óñèëåíèè ýêñïðåññèè ïðîèíôëàìàòîðíûõ öèòîêèíîâ
â ïå÷åíè è ñåëåçåíêå ïòèöû, íà îñíîâå ÷åãî ðàçðàáàòûâàþòñÿ ñîîòâåòñòâóþùèå äèàãíîñòèêóìû.
Ñîñòîÿíèå ñòðàõà — îäíî èç ïðîÿâëåíèé ñòðåññà ó êóð, ïðè ýòîì
ôèçèîëîãè÷åñêèì èíäèêàòîðîì ñòðàõà, â ñâîþ î÷åðåäü, ñëóæèò âðåìÿ òî24
íè÷åñêîé íåïîäâèæíîñòè (51). Òîíè÷åñêàÿ íåïîäâèæíîñòü ìàëî èçó÷åíà è
ïðîÿâëÿåòñÿ â ñíèæåííîé ðåàêòèâíîñòè íà âíåøíèå âîçäåéñòâèÿ ïðè ïðèíóäèòåëüíîé ôèêñàöèè. Ïðîäîëæèòåëüíîñòü òîíè÷åñêîé íåïîäâèæíîñòè
ñâÿçàíà ñ àãðåññèâíûì ïîâåäåíèåì, ðàñêëåâîì è êàííèáàëèçìîì êóð â óñëîâèÿõ ïðîìûøëåííîãî ñîäåðæàíèÿ. Îíà ìîæåò áûòü êðèòåðèåì õðîíè÷åñêèõ ñòðåññîâ, èíäèâèäóàëüíîé ñòðåññîâîé ÷óâñòâèòåëüíîñòè è ñêëîííîñòè
ê àãðåññèâíîìó ïîâåäåíèþ (51-54).
Ê ïðèçíàêàì ñòðåññà ó êóð òàêæå îòíîñèòñÿ ïîòåðÿ ïåðüåâîãî ïîêðîâà.  ÷àñòíîñòè, ïî ñòåïåíè ïîòåðè ïåðà ñóäÿò î ñòðåññîâîì ñîñòîÿíèè
ïòèöû â ïåðèîä ïèêà ïðîäóêòèâíîñòè. Ïîòåðÿ ïåðà ñâÿçàíà ñ ðàñêëåâîì è
êàííèáàëèçìîì, à òàêæå ñ óâåëè÷åíèåì êîëè÷åñòâà êîðòèêîñòåðîíà â êðîâè (3, 37, 55, 56). Íàèáîëåå ñîâåðøåííûå ìåòîäû ïîçâîëÿþò êîìïëåêñíî
îöåíèâàòü ñîñòîÿíèå îïåðåíèÿ íà îñíîâàíèè ïîäñ÷åòà ïåðüåâ íà ðàçëè÷íûõ ó÷àñòêàõ òåëà ñ ïðåäñòàâëåíèåì ðåçóëüòàòîâ â áàëëàõ (57).
 íàñòîÿùåå âðåìÿ ïðîõîäèò àïðîáàöèþ ïðèåì äèàãíîñòèêè ñòðåññîâ, îñíîâàííîé íà âûÿâëåíèè áåëêîâ îñòðîé ôàçû âîñïàëåíèÿ. Ïåðñïåêòèâíûì ìàðêåðîì äëÿ ýòèõ öåëåé ïðåäñòàâëÿåòñÿ êèñëûé α1-ãëèêîïðîòåèí
(îðîçîìóêîèä) (58, 59).
Ïðèíöèïèàëüíî íîâûé ïîäõîä ê äèàãíîñòèêå è ïðîôèëàêòèêå ñòðåññîâ ïðåäëàãàþò Â.È. Ôèñèíèí è Ï. Ñóðàé (10). Ñîãëàñíî èõ äàííûì, óñòîé÷èâîñòü ê ñòðåññàì è êîððåêöèÿ ïîâðåæäåíèé íà ìîëåêóëÿðíîì óðîâíå
çàâèñÿò îò ôóíêöèîíàëüíîé àêòèâíîñòè âèòàãåíîâ, èãðàþùèõ âàæíåéøóþ
ðîëü â îáåñïå÷åíèè àäàïòàöèîííîé ñïîñîáíîñòè îðãàíèçìà. Âëèÿÿ íà ìåòàáîëèçì, îíè ó÷àñòâóþò â âîññòàíîâëåíèè íàðóøåííîãî ïðè ñòðåññå ãîìåîñòàçà è, êàê ñëåäñòâèå, çäîðîâüÿ è ïðîäóêòèâíîñòè ïòèöû. Âèòàãåíû êîíòðîëèðóþò ýêñïðåññèþ àíòèîêñèäàíòíûõ ôåðìåíòîâ ñóïåðîêñèääèñìóòàçû
è ãëóòàòèîíïåðîêñèäàçû, áåëêîâ òåïëîâîãî øîêà (øàïåðîíîâ), ñèðòóèíîâ
(ñåìåéñòâî ôåðìåíòîâ, êîòîðûå îêàçûâàþò çàùèòíîå äåéñòâèå â óñëîâèÿõ
ñòðåññà, ïðåäóïðåæäàÿ ïîâðåæäåíèÿ, âûçûâàåìûå ñâîáîäíûìè ðàäèêàëàìè),
ôåðìåíòîâ âòîðîé ôàçû äåòîêñèêàöèè ÷óæåðîäíûõ âåùåñòâ, à òàêæå ôàêòîðîâ ðîñòà è áåëêîâ, âîâëå÷åííûõ â ðåãóëÿöèþ ïðîöåññîâ ýíåðãåòè÷åñêîãî
îáìåíà è ïîääåðæàíèÿ êëåòî÷íîãî ãîìåîñòàçà êàëüöèÿ (10). Ïðè ñëàáîì
ñòðåññå îáðàçîâàíèå ñâîáîäíûõ ðàäèêàëîâ â êëåòêå íàõîäèòñÿ ïîä êîíòðîëåì àíòèîêñèäàíòíîé ñèñòåìû è ÷àñòîòà ïîâðåæäåíèé ÄÍÊ òàêîâà, ÷òî
ïîçâîëÿåò ñèðòóèíàì óñïåøíî îñóùåñòâëÿòü êàê ðåïàðàöèîííóþ, òàê è ðåãóëÿòîðíóþ ôóíêöèþ. Ñ óñèëåíèåì ñòðåññà êîëè÷åñòâî îáðàçóþùèõñÿ ñâîáîäíûõ ðàäèêàëîâ óâåëè÷èâàåòñÿ èç-çà ïðîãðåññèðóþùåãî ïîâðåæäåíèÿ
ìèòîõîíäðèé, â êîòîðûõ ïðîèñõîäÿò ïðîöåññû êëåòî÷íîãî äûõàíèÿ, è ðåïàðàöèîííàÿ ôóíêöèÿ ñèðòóèíîâ ñòàíîâèòñÿ ïðåîáëàäàþùåé. Êàê ñëåäñòâèå, â êëåòêå íàðóøàåòñÿ ìåòàáîëè÷åñêèé áàëàíñ, ÷òî íåãàòèâíî âëèÿåò íà
ðîñò, ðàçâèòèå, ïðîäóêòèâíûå è ðåïðîäóêòèâíûå êà÷åñòâà æèâîòíûõ è ïòèöû (4). Äèàãíîñòè÷åñêàÿ öåííîñòü äàííûõ î ïåðåêèñíîì îêèñëåíèè ëèïèäîâ ïðè ñòðåññå â íàñòîÿùåå âðåìÿ äîêàçàíà, â òî âðåìÿ êàê êîìïëåêñíîå
èññëåäîâàíèå ïðîäóêòîâ ïåðåêèñíîãî îêèñëåíèÿ, øàïåðîíîâ è ñèðòóèíîâ
ïðè àêòèâàöèè ñòðåññ-ðåàëèçóþùèõ ìåõàíèçìîâ îòêðûâàåò íîâûå, ïîêà
÷òî ìàëî èçó÷åííûå âîçìîæíîñòè äëÿ âûÿâëåíèÿ è ïðîôèëàêòèêè ñòðåññîâ â ïòèöåâîäñòâå.
Äëÿ òîãî ÷òîáû ïîëíîñòüþ èñêëþ÷èòü âíåøíåå âîçäåéñòâèå ïðè
äèàãíîñòèêå ñòðåññîâ è ñòðåññîâîé ÷óâñòâèòåëüíîñòè ó ïòèöû, ðàçðàáîòàíû íåèíâàçèâíûå ìåòîäû îïðåäåëåíèÿ ãîðìîíîâ â áèîëîãè÷åñêèõ ïðîáàõ.
Òàê, J. Downing (60) óêàçûâàåò íà âîçìîæíîñòü èñïîëüçîâàòü õèìè÷åñêèé
àíàëèç ÿéöà ïðè äèàãíîñòèêå ñòðåññîâ. Êàê èçâåñòíî, â ÿéöå íàêàïëèâàþòñÿ
25
ñòåðîèäíûå ãîðìîíû (60, 61), ïðè÷åì, ïî äàííûì F. Royo ñ ñîàâò. (62),
ïðèáëèçèòåëüíî 80 % êîðòèêîñòåðîíà íàõîäèòñÿ â æåëòêå è 20 % — â áåëêå. Îäíàêî îïðåäåëåíèå ñòðåññîâîãî ñîñòîÿíèÿ è ñòðåññîâîé ÷óâñòâèòåëüíîñòè êóð ïî ñîäåðæàíèþ êîðòèêîñòåðîíà â ÿéöå ïîêà ÷òî íå íàøëî ïðàêòè÷åñêîãî ïðèìåíåíèÿ: îäíîçíà÷íàÿ èíòåðïðåòàöèÿ ðåçóëüòàòîâ òàêîãî èññëåäîâàíèÿ çàòðóäíåíà, êðîìå òîãî, âðåìÿ, êîãäà ìåòîä ìîæåò ïðèìåíÿòüñÿ, îãðàíè÷åíî ñðîêîì ôîðìèðîâàíèÿ ÿéöà. Äëÿ äèàãíîñòèêè ñòðåññîâ ìëåêîïèòàþùèõ øèðîêî èñïîëüçóåòñÿ âûÿâëåíèå ãëþêîêîðòèêîèäíûõ ãîðìîíîâ â ôåêàëèÿõ (63).  íàñòîÿùåå âðåìÿ ðàçðàáàòûâàþòñÿ ñïîñîáû îöåíêè
ñòðåññîâîãî ñîñòîÿíèÿ ó êóð, îñíîâàííûå íà îïðåäåëåíèè êîëè÷åñòâà ãëþêîêîðòèêîèäîâ è èõ ìåòàáîëèòîâ ìåòîäàìè ðàäèîèììóíîëîãè÷åñêîãî àíàëèçà, ìàññ-ñïåêòðîìåòðèè è ãàçîæèäêîñòíîé õðîìàòîãðàôèè (64). Ïîäîáíûå ïîäõîäû ìîãóò óñïåøíî èñïîëüçîâàòüñÿ â ëàáîðàòîðíûõ ýêñïåðèìåíòàõ, îäíàêî â óñëîâèÿõ ïòèöåôàáðèê îíè òåõíè÷åñêè íåäîñòóïíû è î÷åíü
äîðîãè äëÿ ïëàíîâîãî ïðîèçâîäñòâåííîãî ïðèìåíåíèÿ.
Íàìè áûë ðàçðàáîòàí ìåòîä, ïîçâîëÿþùèé áåç íåïîñðåäñòâåííîãî
âîçäåéñòâèÿ íà îñîáåé äèàãíîñòèðîâàòü ñòðåññîâîå ñîñòîÿíèå ó êóð ðîäèòåëüñêîãî ñòàäà ìÿñíîãî íàïðàâëåíèÿ ïðîäóêòèâíîñòè, èñïîëüçóÿ êîðòèêîñòåðîí â êà÷åñòâå ñïåöèôè÷åñêîãî ìàðêåðà ñòðåññîâîé ðåàêöèè (65). Ñïîñîá îñíîâàí íà êîëè÷åñòâåííîì îïðåäåëåíèè êîðòèêîñòåðîíà â ïðîáàõ ïîìåòà, îòîáðàííûõ ÷åðåç 1-3 ÷ ïîñëå âîçäåéñòâèÿ ôàêòîðà, ïðåäïîëîæèòåëüíî âûçûâàþùåãî ñòðåññ. Ñîäåðæàíèå ãîðìîíà îöåíèâàëè ïîñëå ýêñòðàêöèè ýòèëîâûì ñïèðòîì, ïðèìåíÿÿ òâåðäîôàçíûé êîíêóðåíòíûé èììóíîôåðìåíòíûé àíàëèç, êîòîðûé äîñòóïåí äëÿ ëàáîðàòîðèé íà áîëüøèíñòâå ðîññèéñêèõ ïòèöåôàáðèê. Ýòîò ïðèåì âûñîêîñïåöèôè÷åí â îòíîøåíèè êîðòèêîñòåðîíà (100 %) ïðè íåçíà÷èòåëüíîé ïåðåêðåñòíîé ðåàêöèè ñ
ïðîãåñòåðîíîì (7,4 %), äèîêñèêîðòèêîñòåðîíîì (3,4 %), 11-äåãèäðîêîðòèêîñòåðîíîì (1,6 %) è äðóãèìè ñòåðîèäàìè (ìåíåå 0,3 %). Èììóíîôåðìåíòíûé àíàëèç êîðòèêîñòåðîíà õàðàêòåðèçóåòñÿ âûñîêîé ÷óâñòâèòåëüíîñòüþ ïðè øèðîêîì äèàïàçîíå îïðåäåëåíèÿ (îò 0 äî 240 íìîëü/ë). Êîðòèêîñòåðîí, êàê è äðóãèå ñòåðîèäû, óñòîé÷èâ âî âíåøíåé ñðåäå è õîðîøî
ðàñòâîðèì â îðãàíè÷åñêèõ ðàñòâîðèòåëÿõ. Ñîäåðæàíèå êîðòèêîñòåðîíà â
ïîìåòå áîëåå 50 íìîëü/ë (65) óêàçûâàåò íà àêòèâàöèþ ãèïîòàëàìî-ãèïîôèçàðíî-íàäïî÷å÷íèêîâîé ñèñòåìû è ñëóæèò èíäèêàòîðîì ñòðåññîâîãî ñîñòîÿíèÿ ó êóð ìÿñíîãî íàïðàâëåíèÿ ïðîäóêòèâíîñòè. Òàêèì îáðàçîì, ïðåäëîæåííûé íàìè ìåòîä ñïåöèôè÷åí, âûñîêî÷óâñòâèòåëåí è ïîëíîñòüþ èñêëþ÷àåò âîçäåéñòâèå íà èññëåäóåìûé îáúåêò, òî åñòü ñîîòâåòñòâóåò îäíîìó
èç âàæíåéøèõ óñëîâèé äîñòîâåðíîé äèàãíîñòèêè ñòðåññîâ. Îí ìîæåò èñïîëüçîâàòüñÿ ïðè îöåíêå öåëåñîîáðàçíîñòè ïðèìåíåíèÿ ñõåì àíòèñòðåññîâîé òåðàïèè, äëÿ îïðåäåëåíèÿ îïòèìàëüíîé ïëîòíîñòè êîìïëåêòîâàíèÿ
ïòèöåé ïðîèçâîäñòâåííûõ ïëîùàäåé, ñ öåëüþ àíàëèçà ñòåïåíè âîçäåéñòâèÿ
òåõíîëîãè÷åñêèõ ôàêòîðîâ è âåòåðèíàðíûõ îáðàáîòîê íà îðãàíèçì êóð.
Èòàê, â íàñòîÿùåå âðåìÿ ðàçðàáîòàíû ìíîãî÷èñëåííûå ìåòîäû, ïîçâîëÿþùèå äèàãíîñòèðîâàòü ñîñòîÿíèå àäàïòàöèîííûõ ìåõàíèçìîâ è èíäèâèäóàëüíîé ñòðåññîâîé ÷óâñòâèòåëüíîñòè ó êóð.  òî æå âðåìÿ ñëåäóåò
îòìåòèòü, ÷òî èñïîëüçîâàíèå òåõ èç ïðåäëîæåííûõ ïîäõîäîâ, êîòîðûå îñíîâàíû íà îïðåäåëåíèè áèîõèìè÷åñêèõ ìàðêåðîâ ñòðåññà â êðîâè, â ïðîèçâîäñòâåííûõ óñëîâèÿõ ïðåäñòàâëÿåò ñîáîé íåïðîñòóþ çàäà÷ó èç-çà íåîáõîäèìîñòè ïðèìåíÿòü ñïåöèàëüíîå îáîðóäîâàíèå, ïðèâëåêàòü âûñîêîêâàëèôèöèðîâàííûé ïåðñîíàë, à òàêæå âñëåäñòâèå íåîäíîçíà÷íîé èíòåðïðåòàöèè ðåçóëüòàòîâ. Âåñüìà èíôîðìàòèâíûì è ìåíåå ñëîæíûì ïðèåìîì ïðè
äèàãíîñòèêå ñòðåññîâîãî ñîñòîÿíèÿ ó êóð ìîæåò áûòü èçó÷åíèå êîëè÷åñò26
âåííîãî ñîîòíîøåíèÿ ãåòåðîôèëîâ è ëèìôîöèòîâ â êðîâè. Âûñîêóþ äèàãíîñòè÷åñêóþ öåííîñòü â ñëó÷àå õðîíè÷åñêèõ ñòðåññîâ èìåþò ïîêàçàòåëè
îáùåãî îïåðåíèÿ.  äîïîëíåíèå ê íèì öåëåñîîáðàçíî ó÷èòûâàòü êîíöåíòðàöèþ ñòðåññîâûõ ãîðìîíîâ â êðîâè è âðåìÿ òîíè÷åñêîé íåïîäâèæíîñòè
êóð. Îäíàêî è â ýòèõ ñëó÷àÿõ ïðîöåäóðû, êîòîðûì ïîäâåðãàåòñÿ ïòèöà, ñàìè ïî ñåáå ñëóæàò ôàêòîðàìè ñòðåññà, òîãäà êàê îäíî èç âàæíåéøèõ óñëîâèé äîñòîâåðíîñòè ðåçóëüòàòîâ ïðè åãî äèàãíîñòèêå — ìèíèìàëüíîå âîçäåéñòâèå íà îáúåêò èññëåäîâàíèÿ. Íàìè ðàçðàáîòàí íåèíâàçèâíûé ñïîñîá,
ïîçâîëÿþùèé âûÿâëÿòü ñîñòîÿíèå ñòðåññà ó êóð ðîäèòåëüñêîãî ñòàäà ìÿñíîãî íàïðàâëåíèÿ ïðîäóêòèâíîñòè ïî ñîäåðæàíèþ êîðòèêîñòåðîíà â ïðîáàõ ïîìåòà. Ýòîò ìåòîä ìîæåò èñïîëüçîâàòüñÿ ïðè îöåíêå öåëåñîîáðàçíîñòè ïðèìåíåíèÿ ñõåì àíòèñòðåññîâîé òåðàïèè, äëÿ îïðåäåëåíèÿ îïòèìàëüíîé ïëîòíîñòè êîìïëåêòîâàíèÿ ïòèöåé ïðîèçâîäñòâåííûõ ïëîùàäåé, à
òàêæå ñ öåëüþ àíàëèçà ñòåïåíè âîçäåéñòâèÿ òåõíîëîãè÷åñêèõ ôàêòîðîâ è
âåòåðèíàðíûõ îáðàáîòîê íà îðãàíèçì êóð.
ÔÃÁÎÓ ÂÏÎ Óðàëüñêàÿ ãîñóäàðñòâåííàÿ
àêàäåìèÿ âåòåðèíàðíîé ìåäèöèíû,
Ïîñòóïèëà â ðåäàêöèþ
3 îêòÿáðÿ 2012
457100 Ðîññèÿ, ×åëÿáèíñêàÿ îáë., ã. Òðîèöê, óë. Ãàãàðèíà, 13,
e-mail: nirugavm@mail.ru
EXPERIMENTAL APPROACHES TO STRESS DIAGNOSTICS IN POULTRY
(review)
A.V. Miftakhutdinov
Ural State Academy of Veterinary Medicine, 13, ul. Gagarina, Troitsk, Chelyabinsk Province, 457100 Russia,
e-mail nirugavm@mail.ru
Received October 3, 2012
Abstract
Special data concerning stresses in poultry and their diagnostics in the laboratories and at
the commercial farms are summarized. To estimate the adaptation mechanisms and individual stress
resistance in chickens, different procedures have been reported till now. But the methods in which
the blood biochemical stress markers must be analyzed are hard to use at commercial farms so far as
special equipment and qualified personnel are required. Besides, the biochemical data from these tests
are not reliable and accurate. Calculation of the blood heterophyls to lymphocytes ratio is considered as
easier and more informative, and the feathering is referred as the highly valuable indicator of chronic
stresses, especially if combined with the stress hormones contents in blood and the time of tonic immobility of chickens. Nevertheless, to evaluate these parameters, a chicken should be subject to the manipulations which are the stressors itself, while only the minimal impact ensures a reliable stress indication. Because of that we improved the method which allows diagnosing stress in parent flocks of the
meat chickens by corticosterone test in excrements. The method can be used to control anti-stress therapy, to optimize the chicken flock size and crowded conditions, and to analyze the impact of applied
technologies and veterinary treatments in poultry.
Keywords: stress, hens, chickens, diagnostics of stresses.
REFERENCES
1.
2.
3.
4.
5.
6.
7.
8.
9.
Fisinin V.I., Papazyan T., Surai P. Ptitsevodstvo, 2009, 8: 10-14.
Fisinin V.I., Kravchenko H.A. Doklady VASKHNIL, 1977, 7: 29-30.
Kavtarashvili A.Sh., Kolokol'nikova T.N. Physiology and productivity of poultry under stress
(review). Sel’skokhozyaistvennaya Biologiya [Agricultural Biology], 2010, 4: 25-37.
Surai P., Fisinin V.I. The modern anti-stress technologies in poultry: from antioxidants to vitagenes. Sel’skokhozyaistvennaya Biologiya [Agricultural Biology], 2012, 4: 3-12.
Fisinin V.I., Surai P. Ptitsa i ptitseprodukty, 2012, 3: 38-41.
Fisinin V.I., Surai P. Ptitsevodstvo, 2012, 2: 11-15.
Fisinin V.I., Surai P. Ptitsevodstvo, 2012, 3: 9-12.
Fisinin V.I., Surai P. Ptitsa i ptitseprodukty, 2012, 4: 36-40.
Surai P., Fisinin V.I. Materialy XVII Mezhdunarodnoi konferentsii VNAP «Innovatsionnye raz-
27
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
28
rabotki i ikh osvoenie v promyshlennom ptitsevodstve» [Proc. XVII Int. Conf. «Innovations and
their use in commercial poultry]. Sergiev Posad, 2012: 24-34.
Fisinin V.I., Surai P. Ptitsa i ptitseprodukty, 2011, 5: 23-27.
Fisinin V.I., Papazyan T.T., Surai P.F. Zhivotnovodstvo segodnya, 2009, 3: 62-67.
Fisinin V.I., Papazyan T.T., Surai P.F. Selenium in poultry nutrition. In: Current advances in
selenium research and applications. Taylor-Pickard, Wageningen Academic Publishers, The
Netherlands, 2008: 221-261.
Fisinin V.I., Mudryi I.N., Kravchenko N.A. Doklady VASKHNIL, 1975, 8: 26-29.
Fisinin V.I., Kravchenko N.A. Sel’skokhozyaistvennaya Biologiya [Agricultural Biology], 1979,
XIV(2): 191-194.
Dawkins M.S. Behavior as a tool in the assessment of animal welfare. Zoology, 2003, 106:
383-387.
Zabudskii Yu.I. Materialy Tret'ei mezhdunarodnoi irano-rossiiskoi konferentsii «Sel'skoe
khozyaistvo i prirodnye resursy» [Proc. 3d Int. Conf. «Iran—Russia: Agriculture and natural resources»]. Moscow, 2002: 134-135.
Brown K.L. Environmental control in poultry production. Oliver& Boyd, Edinburg, 1967: 101113.
Blokhuis H.J., Van Niekerk T.F., Bessei W., Elson A., Guemene D., Kjaer J.B., Levrino
G.A.M., Nicol C.J., Tauson R., Weeks C.A., De Weerd H.A.V. Welfare implications of changes
in production systems for laying hens. World’s Poult. Sci. J., 2007, 63(1): 101-114.
Fisinin V.I., Kravchenko N.A. Ptitsevodstvo, 1977, 7: 29-30.
Vozianov A.F., Butenko A.K., Zak K.P. Tsitokiny. Biologicheskie i protivovospalitel'nye svoistva
[Cytokines. Biological and anti-inflammatory properties]. Kiev, 1998.
Fisinin V.I. Vestnik sel'skokhozyaistvennoi nauki, 1976, 4: 81-85.
Mormede P., Andanson S., Auprein B., Beerda B., Guemene D., Mal-mkvist J.,
Manteca X., Manteuffel G., Prunet P., Van Reenen C.G., Richard S., Viessier J. Exploration of the hypothalamic-adrenal-axis as a tool to evaluate animal welfare. Physiol. Behav.,
2007, 92: 317-339.
Craig J.V., Muir W.M. Group selection for adaptation to multiple-hen cages: behavioral responses. Poultry Sci., 1996, 75: 1145-1155.
Goldstein D.S. Catecholamines and stress. Endocrine Regulations, 2003, 37: 69-80.
Holladay S.D., Edens F.W. Effect of caponization on central nervous system monoamines in
the Japanese quail. Comp. Biochem. Physiol., 1987, 86(3): 465-468.
Goldstein D.S. Plasma norepinephrine as an indicator of sympathetic neural activity in clinical
cardiology. Am. J. Cardiol., 1981, 48: 1147-1154.
Dillon J.E., Raleigh M.J., Mcguire M.T., Bergin-Pollack D., Yuwiler A. Plasma catecholamines
and social behavior in male vervet monkeys (Cercopithecus aethiops sabaeus). Physiol. Behav.,
1992, 51: 973-977.
Brown K.I., Nestor K.E. Implications of selection for high and low adrenal response to stress.
Poultry Sci., 1974, 53: 1297-1306.
Moffitt T.E., Brammer G.L., Caspi A., Fawcett J.P., Raleigh M., Yuwiler A., Silva P. Whole
blood serotonin relates to violence in an epidemiological study. Biological Psychiatry, 1998, 43:
446-457.
Lorrain D.S., Matuszewich L., Hull E.M. 8-OH-DPAT influences extracellular levels of serotonin and dopamine in the medial preoptic area of male rats. Brain Res., 1998, 790: 217-223.
Glick B. Antibody and gland studies in cortisone and ACTH-injected birds. J. Immunol., 1967,
98: 1076-1084.
Korte S.M., Beuving G., Ruesink W., Blockhuis H.J. Plasma catecholamine and corticosterone
levels during manual restraint in chicks from a high and low feather pecking line of laying hens.
Physiol. Behav., 1997, 62: 437-441.
Virden W.S., Thaxton J.P., Corzo A., Dozier W.A., Kidd M.T. Environment, well-being and
behavior evaluation of models using corticosterone and adrenocorticotropin to induce conditions mimicking physiological stress in commercial broilers. Poultry Sci., 2007, 86: 2485-2491.
Shini S., Kaiser P., Shini A., Bryden W.L. Biological response of chickens (Gallus gallus domesticus) induced by corticosterone and a bacterial endotoxin. Comp. Biochem. Physiol., 2008, 149:
324-333.
Rushen J. Problems associated with the interpretation of physiological data in the assessment of
animal welfare. Appl. Animal Behav. Sci., 1991, 28: 381-386.
Moneva P., Popova-Ralcheva S., Abadjieva D., Gudev D., Sredkova V. Poultry welfare assessment, is it possible to avoid handling-induced mental stress interference? Biotechnology in Animal Husbandry, 2009, 25(5-6): 1055-1062.
Miftakhutdinov A.V. Problemy biologii produktivnykh zhivotnykh, 2012, 1: 46-52.
Buyse J., Decuypere E., Darras V.M., Vleurick L.M., Kuhn E.R., Veldhuis J.D. Food depriva-
39.
40.
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
56.
57.
58.
59.
60.
61.
62.
tion and feeding of broiler chickens is associated with rapid and interdependent changes in the
somatotriphic and thyrotropic axes. Br. Poult. Sci., 2000, 41: 107-116.
De Jong I.C., Van Voorst A.S., Erkens J.H.F., Ehlhardt D.A., Blokhuis H.J. Determination of
the circadian rhythm in plasma corticosterone and catecholamine concentrations in growing
broiler breeders using intravenous cannulation. Physiol. Behav., 2001, 74: 299-304.
Savory C.J., Mann J. Is there a role for corticosterone in expression of abnormal behaviour in
restricted-fed fowls? Physiol. Behav., 1997, 62: 7-13.
Gross W.B., Siegel H.S. Evaluation of the heterophil/lymphocyte ratio as a measure of stress in
chickens. Avian Diseases, 1983, 27(4): 972-979.
Maxwell M.H., Hocking P.M., Robertson G.W. Differential leucocyte responses to various degrees of food restriction in broilers, turkeys and ducks. Br. Poult. Sci., 1992, 33: 177-187.
Edens F.W., Siegel H.S. Adrenal responses in high and low ACTH response lines of chickens
during acute heat stress. Gen. Comp. Endocrinol., 1975, 25: 64-73.
Maxwell M.H. Avian blood leucocyte responses to stress. World's Poult. Sci. J., 1993, 49: 34-43.
Post J., Rebel J.M., Huurne A.A. Physiological effect of elevated plasma corticosterone concentrations in broiler chickens, an alternative means by which to assess the physiological effects of
stress. Poultry Sci., 2003, 82: 1313-1318.
Maxwell M.H., Robertson G.M. The avian heterophil leucocyte: a review. World's Poult. Sci. J.,
1998, 54: 155-178.
Altan O., Altan A., Qabuk M., Bayraktar H. Effects of heat stress on some blood parameters in
broilers. Turkish J. Vet. Animal Sci., 2000, 24: 145-148.
Zabudskii Yu.I., Skutar' I.G. Veterinariya, 1993, 3: 44-46.
Felten S.Y., Madden K.S., Bellinger D.L., Kruszewska B., Moynihan J.A., Felten D.L. The
role of the sympathetic nervous system in the modulation of immune responses. Adv. Pharmacol., 1998, 42: 583-587.
Kang S.-Y., Ko Y.-H., Moon Y.-S., Sohn S.-H., Jang I.-S. Effects of the sombined stress induced by stocking density and feed restriction on hematological and cytokine parameters as
stress indicators in laying hens. Asian-Australasian Association of Animal Societies, 2011, 24(3):
414-420.
Jones R.B. Fear and adaptability in poultry: insights, implications and imperatives. World's
Poult. Sci. J., 1996, 52: 131-174.
Blokhuis H.J., Beuving G. Feather pecking and other characteristics in two lines of laying hen.
Proc. the 4th European Symposium on Poultry Welfare /C.J. Savory, B.O. Hughes (eds.). Universities Federation for Animal Welfare, Potters Bar, UK, 1993: 266.
Vestergaard K.S., Kruijtt J.P., Hogan J.A. Feather pecking and chronic fear in groups of red
jungle fowl: their relations to dustbathing, rearing environment and social stress. Animal Behav.,
1993, 62: 413-419.
Jones R.B., Blockhuis H.J., Beuving H.J. Open field and tonic immobility responses in domestic chicks of two genetic lines differing in their propensity to feather peck. Br. Poult. Sci., 1995,
36: 525-530.
Folsch D.W., Benelli A., Gozzoli L. Die Auswirkungen der Bodenund Batterihaltung mit
Unterschiedlicher Besatzdichte auf die Fortbewegung und das Gefieder von Legehennen. Proc.
6th European Poultry Conference. Hamburg, 1980: 160-168.
Shcherbatov V.I., Sidorenko L.I., Levchenko E.V., Pakhomova T.I., Kutovenko T.A., Dzholova
M.N. Sposob prognoza stressa u yaichnykh kur. Patent RF na izobretenie ¹ 2151502, A01K67/02.
Opubl. 27.06.2000, Byul. 2-2003 [Method to predict stress in egg hens. Patent of the Russian
Federation ¹ 2151502, A01K67/02. Published 27.06.2000, Bull. 2-2003].
Tauson R., Kjaer J., Maria G., Cepero R., Holm K. The creation of a common scoring system
for the integument and health of laying hens. In: Applied scoring of integument and health in laying hens. Specific Targeted Research Project. Swedish University of Agriculture Sciences, 2004:
70.
Grigor P.N., Hughes B.O., Appleby M.C. Social inhibition of movement in domestic hens.
Animal Behav., 1995, 49: 1381-1388.
Salamano G., Mellia E., Tarantola M., Gennero M.S., Doglione L., Schiavone A. Acute phase
proteins and heterophil:lymphocyte ratio in laying hens in different housing systems. Vet. Record, 2010, 167: 749-751.
Downing J. Non-invasive assessment of stress in commercial housing systems. A report for the Australian Egg Corporation Limited. Australian Egg Corporation Limited. AECL Publication No
US108A, 2012: 69.
Rettenbacher S., Mostl E., Groothuis T.G.G. Gestagens and glucocorticoids in chicken eggs.
Gen. Compar. Endocrinol., 2009, 164: 125-129.
Royo F., Mayo S., Carlsson H.E., Hau J. Egg corticosterone: a noninvasive measure of stress in
egg-laying birds. Journal of Avian Medicine and Surgery, 2008, 22: 310-314.
29
63. Rogovin K.A., Tupikin A.A., Randall Dzh.A., Kolosova I.E., Moshkin M.P. Zhurnal obshchei
biologii, 2006, 1(67): 37-52.
64. Möstl E., Rettenbacher S., Palme R. Measurement of corticosterone metabolites in birds droppings: an analytical approach. Annual New York Academy Science, 2005, 1046: 17-34.
65. Miftakhutdinov A.V. Sposob opredeleniya stressovogo sostoyaniya kur myasnogo napravleniya
produktivnosti. Polozhitel'noe reshenie na vydachu patenta na izobretenie po zayavke
¹ 2011120009 ot 18.05.2011 [Method to estimate stress in meat hens. Application for Innovation Certificate of the Russian Federation ¹ 2011120009, May 18, 2011].
30