ÑÅËÜÑÊÎÕÎÇßÉÑÒÂÅÍÍÀß ÁÈÎËÎÃÈß, 2014, ¹ 4, ñ. 30-34 ÓÄÊ 579.64:579.61/.62:579.26 doi: 10.15389/agrobiology.2014.4.30rus ÈÇÓ×ÅÍÈÅ ÏÎÏÓËßÖÈÉ ÏÀÒÎÃÅÍÍÛÕ ËÈÑÒÅÐÈÉ Â ÎÁÚÅÊÒÀÕ ÎÊÐÓÆÀÞÙÅÉ ÑÐÅÄÛ Ñ ÈÑÏÎËÜÇÎÂÀÍÈÅÌ ÑÊÀÍÈÐÓÞÙÅÉ ÝËÅÊÒÐÎÍÍÎÉ ÌÈÊÐÎÑÊÎÏÈÈ È.Á. ÏÀÂËÎÂÀ, Ä.À. ÁÀÍÍÈÊÎÂÀ, À.Á. ÊÎÍÎÍÅÍÊÎ Â îòíîøåíèè ðÿäà èíôåêöèîííûõ çàáîëåâàíèé, âûçûâàåìûõ ïàòîãåííûìè è óñëîâíîïàòîãåííûìè áàêòåðèÿìè, ïðîèñõîäèò ïîñòåïåííûé ïåðåñìîòð ìåñòà âîçáóäèòåëÿ â ðàçëè÷íûõ ýêîñèñòåìàõ, àêòóàëüíûìè îñòàþòñÿ âîïðîñû ðàñïðîñòðàíåíèÿ, àäàïòàöèè, ñïîñîáíîñòè èõ æèçíåííûõ ôîðì ê ðàçâèòèþ â ïîïóëÿöèÿõ â ñðåäå îáèòàíèÿ. Èçâåñòíî, ÷òî ëèñòåðèè óñòîé÷èâû ê âíåøíèì ôàêòîðàì, ïðè ýòîì ñóùåñòâåííî ó÷àñòèëèñü ñëó÷àè ïåðåäà÷è ëèñòåðèîçà ñ ðàñòèòåëüíûìè ïðîäóêòàìè. Îäíàêî äàííûå îá ýêîëîãè÷åñêèõ àñïåêòàõ ñóùåñòâîâàíèÿ âîçáóäèòåëÿ ëèñòåðèîçà îãðàíè÷åíû. Îäíî èç ïåðñïåêòèâíûõ íàïðàâëåíèé â èçó÷åíèè ýêîëîãèè ïàòîãåííûõ áàêòåðèé ― ïîïóëÿöèîííûé ïîäõîä, îñíîâàííûé íà èññëåäîâàíèè êîëîíèé áàêòåðèé, à íå îòäåëüíîé êëåòêè.  ñòàòüå ïðèâåäåíû äàííûå ýêñïåðèìåíòàëüíûõ èññëåäîâàíèé, ïðîâåäåííûõ ñ ïðèìåíåíèåì ìåòîäà ñêàíèðóþùåé ýëåêòðîííîé ìèêðîñêîïèè, ïîçâîëÿþùèì àíàëèçèðîâàòü êîëîíèè áàêòåðèé áåç íàðóøåíèÿ èõ åñòåñòâåííîé àðõèòåêòîíèêè. Âûïîëíåíî ñðàâíåíèå ìîðôîëîãè÷åñêèõ îñîáåííîñòåé ïîïóëÿöèé ëèñòåðèé â øèðîêîì äèàïàçîíå òåìïåðàòóð (îò −18 äî 37 °Ñ) â ðàçëè÷íûõ îáúåêòàõ ñðåäû îáèòàíèÿ, òàêèõ êàê ìîëî÷íûå è ìÿñíûå ïðîäóêòû, êîðìà, âîäà, çåëåíûå ðàñòåíèÿ. Ìîðôîëîãè÷åñêàÿ ïëàñòè÷íîñòü êëåòîê ëèñòåðèé è îñîáåííîñòè ñóùåñòâîâàíèÿ ïîïóëÿöèé ëèñòåðèé â îêðóæàþùåé ñðåäå ïðîäåìîíñòðèðîâàíû íà ïðèâåäåííûõ ñêàíîãðàììàõ. Ïîêàçàíî, ÷òî ëèñòåðèè àäàïòèðóþòñÿ ê ìåíÿþùèìñÿ óñëîâèÿì ñóùåñòâîâàíèÿ, ïðèîáðåòàÿ ñïîñîáíîñòü ïåðåõîäèòü ê ãåòåðîìîðôèçìó ñ ðàçëè÷íûìè ïðîÿâëåíèÿìè L-òðàíñôîðìàöèè, ÷òî îáåñïå÷èâàåò èì äëèòåëüíóþ ïåðñèñòåíöèþ â îêðóæàþùåé ñðåäå. Ïîëó÷åííûå äàííûå ïîçâîëèëè âûÿâèòü îñíîâíûå çàêîíîìåðíîñòè àäàïòàöèîííûõ ìåõàíèçìîâ âûæèâàåìîñòè ó ýòèõ áàêòåðèé. Ê òàêèì ìåõàíèçìàì, â ÷àñòíîñòè, îòíîñèòñÿ ãåòåðîìîðôèçì ñ ðàçëè÷íûìè ïðîÿâëåíèÿìè L-òðàíñôîðìàöèè; ñïîñîáíîñòü ê ôîðìèðîâàíèþ ìèêðîêîëîíèé è êîëîíèé ïî òèïó ìíîãîêëåòî÷íîãî îðãàíèçìà; öåëîñòíîñòü ïîïóëÿöèè â ïðîöåññå ðàçâèòèÿ çà ñ÷åò âíóòðèïîïóëÿöèîííûõ ñâÿçåé; ñïîñîáíîñòü ïîïóëÿöèè ðåàãèðîâàòü íà âîçäåéñòâèå ñðåäû îáèòàíèÿ. Êëþ÷åâûå ñëîâà: ïàòîãåííûå áàêòåðèè, ïîïóëÿöèè ëèñòåðèé, ãåòåðîìîðôèçì, Lòðàíñôîðìàöèÿ, îêðóæàþùàÿ ñðåäà, ñàïðîôèòèçì, ñêàíèðóþùàÿ ýëåêòðîííàÿ ìèêðîñêîïèÿ. Íàêîïëåíèå äàííûõ î ñàïðîôèòè÷åñêîì ñóùåñòâîâàíèè ïàòîãåííûõ ìèêðîîðãàíèçìîâ ïðèâåëî ê ïåðåñìîòðó ìåñòà âîçáóäèòåëåé ðÿäà èíôåêöèîííûõ çàáîëåâàíèé â ðàçëè÷íûõ ýêîñèñòåìàõ (1-4).  ïîñëåäíåå äåñÿòèëåòèå ñïåöèàëèñòû ïî ïèùåâûì òîêñèêîèíôåêöèÿì âñå áîëüøå âíèìàíèÿ óäåëÿþò âîçáóäèòåëþ ëèñòåðèîçà, ñëó÷àè ïåðåäà÷è êîòîðîãî ñ ðàñòèòåëüíûìè ïðîäóêòàìè ñóùåñòâåííî ó÷àñòèëèñü. Íåñìîòðÿ íà çíà÷èòåëüíîå ÷èñëî ïóáëèêàöèé ïî ýïèçîîòîëîãèè, ýïèäåìèîëîãèè, ìîëåêóëÿðíîé áèîëîãèè ïàòîãåíà, ôàðìàêîëîãèè, èììóíîëîãèè, ãåíåòèêå è êëèíè÷åñêèì îñîáåííîñòÿì çàáîëåâàíèÿ, ýêîëîãèÿ âîçáóäèòåëÿ ëèñòåðèîçà èçó÷åíà ìàëî (1, 5, 6).  òî æå âðåìÿ èçâåñòíî, ÷òî ëèñòåðèè äîñòàòî÷íî óñòîé÷èâû ê íåáëàãîïðèÿòíûì ôàêòîðàì, ìîãóò ðàñòè ïðè íèçêèõ çíà÷åíèÿõ ðÍ, âûñîêîé êîíöåíòðàöèè NaCl, ïðîÿâëÿþò ñâîéñòâà ìèêðîàýðîôèëîâ è ïñèõðîôèëîâ. Êàê òèïè÷íûå ïðåäñòàâèòåëè ñàïðîíîçîâ ëèñòåðèè ñïîñîáíû ðàçìíîæàòüñÿ â ïî÷âå, ïåðåìåùàòüñÿ ñ ãðóíòîâûìè âîäàìè è ïîäíèìàòüñÿ â ñëîè ñ âåãåòèðóþùèìè ïîáåãàìè îäíîëåòíèõ ðàñòåíèé, â òîì ÷èñëå êîðìîâûõ. Âïåðâûå âîïðîñ î ñóùåñòâîâàíèè ïðèðîäíûõ ðåçåðâóàðîâ âîçáóäèòåëåé èíôåêöèé âî âíåøíåé ñðåäå ïîäíÿë Â.È. Òåðñêèõ (4).  íàñòîÿùåå âðåìÿ ïðèçíàåòñÿ, ÷òî èçó÷àòü ôóíäàìåíòàëüíûå è ïðèêëàäíûå àñïåêòû ýòîé ïðîáëåìû íåîáõîäèìî ñ øèðîêèõ áèîëîãè÷åñêèõ ïîçèöèé íà îñíîâå ïîïóëÿöèîííî-ýêîëîãè÷åñêèõ ïîäõîäîâ (7, 8). Àäàïòàöèÿ ïàòîãåíîâ ñîïðîâîæäàåòñÿ ãåòåðîìîðôèçìîì è èçìåíåíèåì ôèçèîëîãèè êëåòîê, ñïîñîáñòâóþùèì óìåíüøåíèþ ýíåðãèè ðîñòà (â ÷àñòíîñòè, ñíèæåíèåì ôåðìåíòà30 òèâíîé àêòèâíîñòè â îòíîøåíèè ðÿäà ñóáñòðàòîâ) (9), ÷òî ÷àñòî îñëîæíÿåò èäåíòèôèêàöèþ èçîëÿòîâ. Ïðè èçó÷åíèè æèçíåííûõ ôîðì ëèñòåðèé â ðàçíûõ ñðåäàõ ìû èñïîëüçîâàëè 48-÷àñîâóþ êóëüòóðó Listeria innocua (10) â S-ôîðìå ñ òèïè÷íûìè êóëüòóðàëüíî-áèîõèìè÷åñêèìè ñâîéñòâàì. Âçàèìîäåéñòâèå ëèñòåðèé ñ ðàñòåíèÿìè èññëåäîâàëè ïî àâòîðñêîé ìåòîäèêå ìîäåëèðîâàíèÿ ïðîíèêíîâåíèÿ ëèñòåðèé ÷åðåç êîðíåâóþ ñèñòåìó â âåãåòàòèâíûå îðãàíû ñ ïðèìåíåíèåì ìåòîäà ñêàíèðóþùåé ýëåêòðîííîé ìèêðîñêîïèè (ÑÝÌ) è îöåíêîé äèíàìèêè âûæèâàåìîñòè áàêòåðèé (10). CÝÌ ïðîâîäèëè íà ýëåêòðîííîì ìèêðîñêîïå Hitachi-800 ñî ñêàíèðóþùåé ïðèñòàâêîé Hitachi-8010 (ßïîíèÿ) ïðè óñêîðÿþùåì íàïðÿæåíèè 75 ê è èíñòðóìåíòàëüíîì óâåëè÷åíèè ½1-10 òûñ. (10). Ïèùåâûå ïðîäóêòû â ñòåðèëüíûõ åìêîñòÿõ êîíòàìèíèðîâàëè âçâåñüþ ëèñòåðèé (5½108 ÊÎÅ/ìë) è îñòàâëÿëè ïðè êîìíàòíîé òåìïåðàòóðå íà 3 ÷, çàòåì ïîìåùàëè â ðàçíûå òåìïåðàòóðíûå óñëîâèÿ (18-22 °Ñ íà 3 ñóò, 4 °Ñ íà 3 ñóò, −8 è −18 °Ñ íà 10 ñóò), ïîñëå ÷åãî îòáèðàëè ïðîáû äëÿ áàêòåðèîëîãè÷åñêîãî àíàëèçà è ÑÝÌ. Î ìîðôîëîãè÷åñêîé ïëàñòè÷íîñòè ëèñòåðèé (ðèñ. 1) ñâèäåòåëüñòâîâàëî íàëè÷èå îâîèäíûõ êëåòîê (òèïè÷íû äëÿ S-êîëîíèé), â ðàçíûõ óñëîâèÿõ ïðåâðàùàþùèõñÿ â ïàëî÷êîâèäíûå, â êëåòêè ïðîòîïëàñòíîãî òèïà, íèòåâèäíûå, ìåëêèå îêðóãëûå (L-ôîðìû), èçâèòûå ôîðìû, ÷òî ïðèñóùå áàêòåðèÿì íà ñòàäèÿõ L-òðàíñôîðìàöèè. Øèðîêàÿ ýêîëîãè÷åñêàÿ òîëåðàíòíîñòü, ñîïðîâîæäàþùàÿñÿ èçìåíåíèåì ìîðôîëîãè÷åñêèõ îñîáåííîñòåé ïîïóëÿöèé, çàòðóäíÿåò ïåðâè÷íóþ èíäèêàöèþ è èäåíòèôèêàöèþ ëèñòåðèé â îáúåêòàõ âåòåðèíàðíî-ñàíèòàðíîãî íàäçîðà. Ïðè ïîíèæåííûõ òåìïåðàòóðàõ, óãíåòàþùèõ áîëüøèíñòâî ìåçîôèëüíûõ ïàòîãåíîâ, ëèñòåðèè íå âñòðå÷àþò êîíêóðåíöèè è ìîãóò àêòèâíî ðàçìíîæàòüñÿ (11). Ïî äàííûì Â.Ñ. Çóåâà ñ ñîàâò. (12), ïðè òåìïåðàòóðå 4-6 °Ñ ó ñàëüìîíåëë â âîäå íàáëþäàåòñÿ L-òðàíñôîðìàöèÿ (12). Ìû îòìå÷àëè (ðèñ. 2, À), ÷òî ëèñòåðèè â âîäå â êîëîíèÿõ è ìèêðîêîëîíèÿõ ñîõðàíÿëè òèïè÷íóþ ïàëî÷êîâèäíóþ ôîðìó è áûëè îáúåäèíåíû ìåæêëåòî÷íûì ìàòðèêñîì, ôîðìèðóþùèì ïîêðîâû, ÷òî ñâèäåòåëüñòâîâàëî î íàëè÷èè ïîëíîöåííîé êëåòî÷íîé ñòåíêè. Îòðèöàòåëüíûå òåìïåðàòóðû (îò −8 äî −18 °Ñ) íå âëèÿëè íà ìîðôîëîãèþ è æèçíåñïîñîáíîñòü ïîïóëÿöèé ëèñòåðèé (13, 14). Îíè ïîääåðæèâàëè æèçíåñïîñîáíîñòü ïðè íèçêèõ ïîëîæèòåëüíûõ è äàæå îòðèöàòåëüíûõ òåìïåðàòóðàõ, ñóùåñòâóÿ â êîëîíèÿõ è ìèêðîêîëîíèÿõ ñ ïîâåðõíîñòíîé áèîïëåíêîé (ñì. ðèñ. 2, Á, Â), îáðàçîâàííîé âíåêëåòî÷íûìè ïðîäóêòàìè, àíàëîãè÷íûìè êîìïîíåíòàì êëåòî÷íîé ñòåíêè, ñèíòåçèðóåìûì è ñåêðåòèðóåìûì ïîëíîöåííîé áàêòåðèåé (15). Èçâåñòíî, ÷òî ó ýíòåðîïàòîãåííûõ áàêòåðèé îñíîâó áèîïëåíêè ñîñòàâëÿþò àäãåçèíû áåëêîâîé è ïîëèñàõàðèäíîé ïðèðîäû (16). Ñòðàòåãèÿ âûæèâàíèÿ ïîïóëÿöèé ëèñòåðèé â ãîëîäíîé ñðåäå (âîäîïðîâîäíàÿ âîäà) ïðè ðàçíûõ òåìïåðàòóðàõ îñíîâàíà íà ãåòåðîìîðôèçìå ñ ðàçëè÷íûìè ïðîÿâëåíèÿìè L-òðàíñôîðìàöèè, ÷òî ðàñøèðÿåò âîçìîæíîñòè ïîääåðæàíèÿ èõ ÷èñëåííîñòè è öèðêóëÿöèè â ìåæýïèäåìèîëîãè÷åñêèå ïåðèîäû (13). Ñâîéñòâî ëèñòåðèé íå òîëüêî âûæèâàòü, íî è ðàçìíîæàòüñÿ ïðè ïîíèæåííûõ òåìïåðàòóðàõ îáóñëîâëåíî àäàïòàöèîííûìè ìåõàíèçìàìè, ïðèñóùèìè ïñèõðîôèëàì.  ïåðâóþ î÷åðåäü, ýòî òàê íàçûâàåìàÿ ãîìåîâÿçêîñòíàÿ àäàïòàöèÿ — èçìåíåíèå ñîñòàâà æèðíûõ êèñëîò â ëèïèäàõ ìåìáðàí â ñîîòâåòñòâèè ñ òåìïåðàòóðîé îêðóæàþùåé ñðåäû. Ïðè ïîíèæåíèè òåìïåðàòóðû ñîäåðæàíèå íåíàñûùåííûõ æèðíûõ êèñëîò ïîâûøàåòñÿ, ÷åì ïðåäîòâðàùàåòñÿ çàìåðçàíèå ëèïèäîâ è èçáûòî÷íàÿ æåñòêîñòü ìåìáðàí. Ê äðóãèì ïîäîáíûì ôàêòîðàì îòíîñèòñÿ ñïîñîáíîñòü êîìïåíñèðîâàòü ìàëóþ ýôôåêòèâíîñòü ðèáîñîì ïðè íèçêèõ òåìïåðàòóðàõ èõ äîïîëíè31 òåëüíûì îáðàçîâàíèåì, ÷òî ïðîÿâëÿåòñÿ â óâåëè÷åíèè ñîäåðæàíèÿ ÐÍÊ. Êðîìå òîãî, àäàïòàöèÿ ïñèõðîôèëîâ ñâÿçàíà ñ êîíôîðìàöèîííûìè èçìåíåíèÿìè áåëêîâ ïîä äåéñòâèåì òåðìîëàáèëüíûõ ôåðìåíòîâ (17-19). Âñå ÷àùå ðåãèñòðèðóþò âñïûøêè ëèñòåðèîçà ïðè óïîòðåáëåíèè â ïèùó ìÿñíûõ è ìîëî÷íûõ ïðîäóêòîâ (6, 20, 21). Ìû ïîêàçàëè, ÷òî ïîñëå êîíòàìèíàöèè ëèñòåðèÿìè èõ êîëîíèåîáðàçóþùàÿ àêòèâíîñòü ïðè 4 °Ñ çà 7 ñóò â ñòåðèëèçîâàííîì ìîëîêå óâåëè÷èâàëàñü íà ÷åòûðå ïîðÿäêà, â êèñëîìîëî÷íûõ ïðîäóêòàõ — íà òðè, â ñûðàõ — íà äâà (22). Ñëåäîâàòåëüíî, íàëè÷èå äàæå íåáîëüøîãî ÷èñëà ëèñòåðèé ñïîñîáíî ïðèâåñòè ê áóðíîìó ðàçâèòèþ ïîïóëÿöèè äî îïàñíîãî äëÿ çäîðîâüÿ êîëè÷åñòâà.  óñëîâèÿõ ïîíèæåííûõ òåìïåðàòóð áûëî óñòàíîâëåíî, ÷òî, êàê è â âîäíîé ñðåäå, îäíà ÷àñòü ïîïóëÿöèè ñóùåñòâóåò â êîëîíèÿõ è ìèêðîêîëîíèÿõ ñ ìåæêëåòî÷íûì ìàòðèêñîì, ôîðìèðóþùèì ïîêðîâû (áèîïëåíêè), äðóãàÿ ― íàõîäèòñÿ íà ðàçíûõ ñòàäèÿõ ãåòåðîìîðôèçìà, ÷òî ñâèäåòåëüñòâóåò î ôåíîòèïè÷åñêîé ãåòåðîãåííîñòè ïîïóëÿöèé (ðèñ. 3, À, Á, Â). Èçó÷åíèå ìîðôîëîãèè è îñîáåííîñòåé ðàçâèòèÿ ëèñòåðèé â ñòåáëÿõ êîðìîâîãî îâñà ñîðòà Óëîâ (10) ïîçâîëèëî êîíñòàòèðîâàòü, ÷òî ýòè ìèêðîîðãàíèçìû èç ïî÷âû ÷åðåç êîðíåâóþ ñèñòåìó ïðîíèêàþò âíóòðü âåãåòàòèâíûõ îðãàíîâ. Òàê, â ãîìîãåíàòå êîðíåé ÷èñëî ëèñòåðèé áûëî âûøå, ÷åì â ãîìîãåíàòå ëèñòîâûõ ïëàñòèíîê. Âíóòðè ëèñòîâûõ ïëàñòèíîê íà âñåõ ñðîêàõ íàáëþäåíèÿ áûë âûÿâëåí ãåòåðîìîðôèçì ëèñòåðèé ñ äåôåêòíîé êëåòî÷íîé ñòåíêîé (ðèñ. 4, À). Èç ìîðôîëîãè÷åñêèõ âàðèàíòîâ îòìå÷àëè áàêòåðèè ñ èçìåíåííîé ôîðìîé, íàðóøåííûì ìåõàíèçìîì äåëåíèÿ, ñ ïîòåðåé ñïîñîáíîñòè ê àäãåçèè è êîëîíèçàöèè, ÷òî, êàê èçâåñòíî, âåäåò ê óòðàòå ïàòîãåííûõ ñâîéñòâ, îáðàçîâàíèå êëåòîê ïðîòîïëàñòíîãî òèïà. Ïðè áëàãîïðèÿòíûõ óñëîâèÿõ ãåòåðîìîðôíûå êëåòêè ðåâåðñèðîâàëè â èñõîäíóþ ôîðìó ñ ñîõðàíåíèåì ìîðôîëîãè÷åñêèõ è ôèçèîëîãè÷åñêèõ ñâîéñòâ (10). Ñòðàòåãèÿ âûæèâàíèÿ ïîïóëÿöèé ëèñòåðèé â ðàñòåíèè îïðåäåëÿåòñÿ êîìïëåêñîì åãî ìåõàíèçìîâ çàùèòû, äåéñòâèå êîòîðûõ âûçûâàåò ðàçðóøåíèå êëåòî÷íîé ñòåíêè, ÷òî ïðèâîäèò ê ðàçëè÷íûì ïðîÿâëåíèÿì L-òðàíñôîðìàöèè è îáðàçîâàíèþ íåñòàáèëüíûõ L-ôîðì, ñïîñîáíûõ ê ðåâåðñèè â èñõîäíîå ñîñòîÿíèå. Ýòîò ïðîöåññ ðàñøèðÿåò âîçìîæíîñòè âîçáóäèòåëÿ ïîääåðæèâàòü ÷èñëåííîñòü è íàêàïëèâàòüñÿ â çåëåíûõ êîðìàõ (23). Êîíòàìèíàöèè êîðìîâ ëèñòåðèÿìè òàêæå ñïîñîáñòâóåò àäãåçèÿ è êîëîíèçàöèÿ òâåðäûõ ÷àñòèö êîðìîâ (24). Ñ èñïîëüçîâàíèåì ÑÝÌ áûëî ïîêàçàíî, ÷òî ïðè ïîíèæåííûõ òåìïåðàòóðàõ è ïîâûøåííîé âëàæíîñòè â òå÷åíèå 24 ÷ êëåòêè ïðèêðåïëÿëèñü ê ïîâåðõíîñòè äðîáëåíîãî çåðíà ñ ïîñëåäóþùèì ôîðìèðîâàíèåì ìèêðîêîëîíèé è êîëîíèé (ñì. ðèñ. 4, Á), òî åñòü ïðîèñõîäèëî àêòèâíîå ðàçìíîæåíèå ëèñòåðèé. Ïðè ïîâûøåíèè òåìïåðàòóðû àäãåçèÿ è êîëîíèçàöèÿ çàìåäëÿëèñü âñëåäñòâèå ïñèõðîôèëüíîñòè ýòèõ áàêòåðèé. Òàêèì îáðàçîì, ïîïóëÿöèè ëèñòåðèé â ðàçíûõ îáúåêòàõ (âîäà, çåëåíûå ðàñòåíèÿ, çåðíî, ïðîäóêòû ïèòàíèÿ) â äèàïàçîíå òåìïåðàòóð îò −18 äî +37 °Ñ àäàïòèðóþòñÿ ê ìåíÿþùèìñÿ óñëîâèÿì, ïðîÿâëÿÿ ãåòåðîìîðôèçì ñ ðàçëè÷íûìè ýëåìåíòàìè L-òðàíñôîðìàöèè, ÷òî îáåñïå÷èâàåò äëèòåëüíóþ ïåðñèñòåíöèþ. L-òðàíñôîðìàöèÿ ñóùåñòâåííî çàòðóäíÿåò èíäèêàöèþ ëèñòåðèé, òàê êàê íåðåäêî âèäèìîãî ðîñòà ïðè êóëüòèâèðîâàíèè òàêèõ áàêòåðèé íà ïèòàòåëüíûõ ñðåäàõ íå íàáëþäàåòñÿ. Ñîáñòâåííûå èññëåäîâàíèÿ è àíàëèç äàííûõ ëèòåðàòóðû ïîçâîëèëè íàì íàçâàòü îñíîâíûå àäàïòàöèîííûå ìåõàíèçìû, îáåñïå÷èâàþùèå ñîõðàíåíèå æèçíåñïîñîáíîñòè ó ëèñòåðèé. Ýòî ôåíîòèïè÷åñêàÿ ãåòåðîãåííîñòü (ãåòåðîìîðôèçì ñ ðàçëè÷íûìè ïðîÿâëåíèÿìè L-òðàíñôîðìàöèè); âîçìîæíîñòü ôîðìèðîâàòü ìèêðîêîëîíèè è êîëîíèè ïî òèïó ìíîãîêëåòî÷íîãî 32 îðãàíèçìà; öåëîñòíîñòü ïîïóëÿöèè çà ñ÷åò âíóòðèïîïóëÿöèîííûõ ñâÿçåé, îòñóòñòâóþùèõ ó åäèíè÷íûõ êëåòîê, è åå ñïîñîáíîñòü ðåàãèðîâàòü íà âîçäåéñòâèå ñðåäû îáèòàíèÿ (9). Èç-çà ñïîñîáíîñòè íåñòàáèëüíûõ L-ôîðì èíèöèèðîâàòü íîâûå êîëîíèé ïîñðåäñòâîì ðåâåðñèè â èñõîäíóþ ôîðìó Â.Ä. Òèìàêîâ è Ã.ß. Êîãàí íàçâàëè ïðîöåññ L-òðàíñôîðìàöèè «òðàíçèòíîé ñèñòåìîé» (25). Òàêîé àäàïòàöèîííûé ìåõàíèçì, ïðîÿâëÿþùèéñÿ èçìåíåíèåì áèîõèìè÷åñêèõ è ìîðôîëîãè÷åñêèõ ñâîéñòâ â ïîïóëÿöèÿõ, ñïîñîáñòâóåò âûæèâàíèþ âèäà áàêòåðèé è ñîçäàåò îïàñíîñòü âîçíèêíîâåíèÿ èíôåêöèîííîãî î÷àãà (9). Èòàê, ñ ïðèìåíåíèåì íåèíâàçèâíîé ñêàíèðóþùåé ýëåêòðîííîé ìèêðîñêîïèè èçó÷åí ãåòåðîìîðôèçì, âîçäåéñòâèå àáèîòè÷åñêèõ è áèîòè÷åñêèõ ôàêòîðîâ ñðåäû íà ïîïóëÿöèþ, ñïîñîáíîñòü ê ðåâåðñèè, ïðîöåññû àäãåçèè è êîëîíèçàöèè è ïð. ó ëèñòåðèé, ÷òî äàëî îáúåêòèâíóþ èíôîðìàöèþ îá èõ ïîïóëÿöèÿõ â ñðåäå îáèòàíèÿ è ðàñøèðèëî ïðåäñòàâëåíèå î ñòðóêòóðíî-ôóíêöèîíàëüíûõ îñîáåííîñòÿõ ïàòîãåííûõ îðãàíèçìîâ â öåëîì.  øèðîêîì ñìûñëå ýòî ïîçâîëÿåò ïî-íîâîìó ïîäîéòè êî ìíîãèì ïðàêòè÷åñêèì ïðîáëåìàì âîçäåéñòâèÿ íà âîçáóäèòåëåé êàê â ñðåäå îáèòàíèÿ, òàê è â ìàêðîîðãàíèçìå, â òîì ÷èñëå îáîñíîâàòü ïîäáîð íîâûõ äåçèíôèöèðóþùèõ ñðåäñòâ è ëåêàðñòâåííûõ ïðåïàðàòîâ. ÃÍÓ Âñåðîññèéñêèé ÍÈÈ âåòåðèíàðíîé ñàíèòàðèè, ãèãèåíû è ýêîëîãèè Ðîññåëüõîçàêàäåìèè, Ïîñòóïèëà â ðåäàêöèþ 20 ìàðòà 2013 ãîäà 123022 Ðîññèÿ, ã. Ìîñêâà, Çâåíèãîðîäñêîå ø., 5, e-mail: sanmicrobio@mail.ru EXPERIMENTAL STUDY OF POPULATIONS OF PATHOGENIC Listeria IN ENVIRONMENT USING SCANNING ELECTRON MICROSCOPY I.B. Pavlova, D.A. Bannikova, A.B. Kononenko All-Russian Research Institute of Veterinary Sanitation, Hygiene and Ecology, Russian Academy of Agricultural Sciences, 5, Zvenigorodskoe sh., Moscow, 123022 Russia, e-mail sanmicrobio@mail.ru Received March 20, 2013 doi: 10.15389/agrobiology.2014.4.30eng Abstract Due to accumulated data, the ideas on existance and the state of pathogenic and opportunistic bacteria in different ecosystems gradually change, while distribution, adaptation, growth and development of bacterial forms in populations in different media and objects remain relevant. Listeria are known to be resistant to external factors, and the frequency of their transmission through plant production increses significantly. At that, the data concerning ecological aspects of living cycle of the pathogen are still limited. Ecology of the bacteria is one of the substantial sections of biology concerning development, survival and interactions of the microorganisms population and the environment. One of the perspective trends in studying ecology of the pathogenic bacteria is approach on population level being based on a study of bacterial colonies, and not a separate cell. The date of the experiments are presented in the paper being carried out with using scanning electron microscopy method resulting in ability to study bacterial colonier without violation of their natural architectonics. The morphological features of Listeria populations in the different objects of inhabitance medium, e.g. milk and meat products, feeds, water, green plants, in a broad temperature rang (from −18 up to 37 °Ñ) were examined. Morphological plasticity of Listeria cells and the peculiarities of Listeria populatios existence in the environment were demonstrated on the presental scanogramms. It was shown that Listeria adapted to the changing conditions of existence acquiring the ability to transform into heteromorphism with the different manifestations of L-transformation resulting in their prolonged persistence in the environment. The obtained data allowed to reveal the main conformities of the survival rate of those bacteria. Specifically, heteromorphism concerns such mechanism with the different manifestations of L-transformation, as well as the ability to form the microcolonies and colonies by a type of multicellular organism, integrity of a population during a process owing to the interpopulations bonds, ability of a population to react onto influence of inhabitance medium. The data etended our ideas on the structure and functions of pathogenic organism. 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