ÑÅËÜÑÊÎÕÎÇßÉÑÒÂÅÍÍÀß ÁÈÎËÎÃÈß, 2014, ¹ 3, ñ. 13-23 ÓÄÊ 632:57.047:[582.288+595.7 ÁÈÎËÎÃÈß ÂÇÀÈÌÎÎÒÍÎØÅÍÈÉ ÃÐÈÁΠÐÎÄÀ Fusarium È ÍÀÑÅÊÎÌÛÕ∗ (îáçîð) Ò.Þ. ÃÀÃÊÀÅÂÀ, È.Â. ØÀÌØÅÂ, Î.Ï. ÃÀÂÐÈËÎÂÀ, Î.Ã. ÑÅËÈÖÊÀß Ïðåäñòàâèòåëè ðîäà Fusarium õàðàêòåðèçóþòñÿ âûñîêîé ìåòàáîëè÷åñêîé àêòèâíîñòüþ è àäàïòàöèîííîé ïëàñòè÷íîñòüþ. Îíè ñïîñîáíû çàñåëÿòü ðàñòèòåëüíûå ñóáñòðàòû, ïðèíèìàòü àêòèâíîå ó÷àñòèå â ðàçëîæåíèè îðãàíè÷åñêèõ îñòàòêîâ è ïî÷âîîáðàçîâàòåëüíîì ïðîöåññå, à òàêæå ïîðàæàòü íàñåêîìûõ è ìëåêîïèòàþùèõ. Ãðèáû ðîäà Fusarium è íàñåêîìûå ñîñóùåñòâóþò âî ìíîãèõ áèîòîïàõ. Òàê, K. O’Donnell ñ ñîàâò. (2012) ïðè ìîëåêóëÿðíî-ãåíåòè÷åñêîé èäåíòèôèêàöèè 140 èçîëÿòîâ ãðèáîâ, âûäåëåííûõ èç íàñåêîìûõ, âûÿâèëè êàê ìèíèìóì 23 âèäà èëè êîìïëåêñà âèäîâ ðîäà Fusarium. Ñâÿçü ìåæäó âèäîâûì ñîñòàâîì ôóçàðèåâûõ ãðèáîâ è ñèñòåìàòè÷åñêèì ïîëîæåíèåì íàñåêîìûõ-õîçÿåâ ïîêà íå óñòàíîâëåíà. Êîýâîëþöèÿ ýòèõ ãðóïï îðãàíèçìîâ ïðèâåëà ê âîçíèêíîâåíèþ ðàçëè÷íûõ ôîðì âçàèìîîòíîøåíèé ìåæäó íèìè. Àíòàãîíèñòè÷åñêàÿ ôîðìà âçàèìîäåéñòâèÿ îêàçûâàåò îäíîíàïðàâëåííûé ýôôåêò è ìîæåò èìåòü ëåòàëüíûå è íåëåòàëüíûå ïîñëåäñòâèÿ äëÿ íàñåêîìûõ.  îáîèõ ñëó÷àÿõ îíà ðåàëèçóåòñÿ ñ ó÷àñòèåì ëåòó÷èõ (íàïðèìåð, ðåïåëëåíòû) è íåëåòó÷èõ (íàïðèìåð, ìèêîòîêñèíû) âòîðè÷íûõ ìåòàáîëèòîâ ãðèáîâ. Ëåòó÷èå îðãàíè÷åñêèå ñîåäèíåíèÿ (ËÎÑ) ñëóæàò ñèãíàëüíûìè ìîëåêóëàìè (èíôîõåìèêè èëè ñåìèîõåìèêè) è ïðîÿâëÿþò ïî îòíîøåíèþ ê íàñåêîìûì ñâîéñòâà ôåðîìîíîâ, àëëîìîíîâ, êàéðîìîíîâ è äð. Èçó÷åíèå ËÎÑ ó ãðèáîâ ðîäà Fusarium áûëî íà÷àòî â êîíöå 1990-õ ãîäîâ, ïóáëèêàöèé ïî ýòîé òåìå ïîêà íåìíîãî. Óñòàíîâëåíî, ÷òî ËÎÑ Fusarium âêëþ÷àþò áîëüøîå ÷èñëî êîìïîíåíòîâ, îòíîñÿùèõñÿ ê ðàçëè÷íûì ãðóïïàì ñîåäèíåíèé. Ìåõàíèçì ïðîíèêíîâåíèÿ ýíòîìîïàòîãåííûõ ïðåäñòàâèòåëåé ðîäà Fusarium â îðãàíèçì õîçÿèíà ïðèíöèïèàëüíî îòëè÷àåòñÿ îò òàêîâîãî ó äðóãèõ ãðóïï ãðèáîâ è ïðîèñõîäèò ÷åðåç åñòåñòâåííûå îòâåðñòèÿ (ðîòîâûå îðãàíû, äûõàëüöà, ÿéöåêëàä) èëè ðàíû, à íå ÷åðåç êóòèêóëó íàñåêîìîãî. Ñèìáèîòè÷åñêèå âçàèìîäåéñòâèÿ ìåæäó âèäàìè ðîäà Fusarium è íàñåêîìûìè õàðàêòåðèçóþòñÿ áîëüøèì ðàçíîîáðàçèåì è âêëþ÷àþò êàê âçàèìîâûãîäíîå ñîñóùåñòâîâàíèå (ìóòóàëèçì), òàê è êîììåíñàëèçì. Èçâåñòíà ðîëü íàñåêîìûõ â ðàñïðîñòðàíåíèè ñïîð ãðèáîâ Fusarium íà íîâûå íåçàñåëåííûå ñóáñòðàòû. Ïîêàçàí àòòðàêòèâíûé ýôôåêò ëåòó÷èõ ìåòàáîëèòîâ ãðèáîâ äëÿ íàñåêîìûõ.  ðÿäå ñëó÷àåâ ôóçàðèåâûå ãðèáû ñëóæàò èñòî÷íèêîì âåùåñòâ, æèçíåííî íåîáõîäèìûõ íàñåêîìûì äëÿ íîðìàëüíîãî ðàçâèòèÿ. Ðàçëè÷íûå ôîðìû âçàèìîîòíîøåíèé ìåæäó ãðèáàìè ðîäà Fusarium è íàñåêîìûìè îáåñïå÷èâàþò ñîñóùåñòâîâàíèå ýòèõ îðãàíèçìîâ â óñòîé÷èâûõ ñàìîðåãóëèðóþùèõñÿ ýêîñèñòåìàõ. Îáñóæäàþòñÿ âîçìîæíûå âçàèìîäåéñòâèÿ ãðèáîâ ðîäà Fusarium è âðåäèòåëåé çàïàñîâ ïðè èñïîëüçîâàíèè èìè çåðíîâîê çëàêîâûõ â êà÷åñòâå îáùåãî ïèùåâîãî ñóáñòðàòà. Êëþ÷åâûå ñëîâà: ãðèáû, Fusarium, íàñåêîìûå, àíòàãîíèñòè÷åñêèå è ñèìáèîòè÷åñêèå âçàèìîîòíîøåíèÿ. Íàñåêîìûå è ãðèáû ïðåäñòàâëÿþò äâå íàèáîëåå ìíîãî÷èñëåííûå è ðàçíîîáðàçíûå ãðóïïû ýóêàðèîò, êîòîðûå ñîñóùåñòâóþò âî ìíîãèõ áèîòîïàõ è âñòóïàþò âî âçàèìîîòíîøåíèÿ, îõâàòûâàþùèå ðàçëè÷íûå òðîôè÷åñêèå óðîâíè. Íàêîïëåíà îáøèðíàÿ ëèòåðàòóðà ïî ðàçëè÷íûì àñïåêòàì âçàèìîäåéñòâèé îòäåëüíûõ ãðóïï ãðèáîâ ñ íàñåêîìûìè, ïðè ýòîì îñîáîå âíèìàíèå óäåëÿåòñÿ áàçèäèîìèöåòàì è ýíòîìîïàòîãåííûì ìèêðîìèöåòàì. Ïðåäñòàâèòåëè ðîäà Fusarium õàðàêòåðèçóþòñÿ âûñîêîé ìåòàáîëè÷åñêîé àêòèâíîñòüþ è àäàïòàöèîííîé ïëàñòè÷íîñòüþ, ÷òî ïîçâîëèëî èì îñâîèòü ìíîãèå ýêîëîãè÷åñêèå íèøè. Îíè ñïîñîáíû çàñåëÿòü ðàñòèòåëüíûå ñóáñòðàòû, ïðèíèìàòü àêòèâíîå ó÷àñòèå â ðàçëîæåíèè îðãàíè÷åñêèõ îñòàòêîâ è ïî÷âîîáðàçîâàòåëüíîì ïðîöåññå, à òàêæå ïîðàæàòü íàñåêîìûõ è ìëåêîïèòàþùèõ. Øèðîêîå ðàñïðîñòðàíåíèå, çíà÷èòåëüíàÿ âðåäîíîñíîñòü è ñïîñîáíîñòü ê îáðàçîâàíèþ âòîðè÷íûõ ìåòàáîëèòîâ, òîêñè÷íûõ äëÿ òåïëîêðîâíûõ, ïîääåðæèâàþò óñòîé÷èâûé èíòåðåñ èññëåäîâàòåëåé ê ýòîé ãðóïïå ãðèáîâ. Ïðèñóòñòâèå ãðèáîâ ðîäà Fusarium íà íàñåêîìûõ èçâåñòíî äàâíî, ∗ Ðàáîòà ïîääåðæàíà ïðîåêòîì ÐÔÔÈ ¹ 12-04-00927. 13 íî òîëüêî â ïîñëåäíåå âðåìÿ ñ èñïîëüçîâàíèåì ìîëåêóëÿðíî-ãåíåòè÷åñêèõ ìåòîäîâ óäàëîñü îáúåêòèâíî îöåíèòü èõ âèäîâîå ðàçíîîáðàçèå. Íàïðèìåð, K. O’Donnell ñ ñîàâò. (1), ïðîâåäÿ èäåíòèôèêàöèþ 140 èçîëÿòîâ ãðèáîâ, âûäåëåííûõ èç íàñåêîìûõ, âûÿâèëè êàê ìèíèìóì 23 âèäà èëè êîìïëåêñà âèäîâ ðîäà Fusarium. Ñóùåñòâóåò åäèíñòâåííûé îáçîð, â êîòîðîì áûëè îáîáùåíû ðåçóëüòàòû èññëåäîâàíèé ïî ðàçíûì àñïåêòàì âçàèìîäåéñòâèé ôóçàðèåâûõ ãðèáîâ è íàñåêîìûõ (2). Ìíîãèå ðàáîòû, óïîìÿíóòûå â ýòîé ïóáëèêàöèè, â íàñòîÿùåå âðåìÿ ñëîæíî îáñóæäàòü â ñâÿçè ñ èñïîëüçîâàíèåì ðàçëè÷íûõ òàêñîíîìè÷åñêèõ ñèñòåì äëÿ îïðåäåëåíèÿ âèäîâ ðîäà Fusarium. Êðîìå òîãî, àâòîðû ñêîíöåíòðèðîâàëè îñíîâíîå âíèìàíèå íà âîïðîñàõ ïàòîãåíåçà âèäîâ ðîäà Fusarium ïî îòíîøåíèþ ê íàñåêîìûì. È íàêîíåö, çà ïîñëåäíèå äâà äåñÿòèëåòèÿ ïîÿâèëîñü ìíîãî ïóáëèêàöèé, ïîñâÿùåííûõ ýòîé ïðîáëåìå, â òîì ÷èñëå ïî òàêèì èíòåíñèâíî ðàçâèâàþùèìñÿ íàïðàâëåíèÿì, êàê ëåòó÷èå îðãàíè÷åñêèå ñîåäèíåíèÿ (ËÎÑ).  îáùåì âèäå âçàèìîäåéñòâèÿ ìåæäó ãðèáàìè ðîäà Fusarium è íàñåêîìûìè ìîãóò áûòü îõàðàêòåðèçîâàíû êàê àíòàãîíèçì è ñèìáèîç.  ñâîþ î÷åðåäü, àíòàãîíèñòè÷åñêèå âçàèìîîòíîøåíèÿ äåëÿòñÿ íà äâå êàòåãîðèè â çàâèñèìîñòè îò èõ ïîñëåäñòâèé äëÿ íàñåêîìîãî: ëåòàëüíûå è íåëåòàëüíûå. Ìíîãèå ôóçàðèåâûå ãðèáû ìîãóò âûçûâàòü ãèáåëü íàñåêîìûõ è/èëè êîëîíèçèðîâàòü ïîãèáøèå îñîáè. Îäíàêî íå âî âñåõ îïèñàííûõ ñëó÷àÿõ óäàåòñÿ äîñòîâåðíî îòäåëèòü áåçóñëîâíî ýíòîìîïàòîãåííûå âèäû ãðèáîâ, çàðàæåíèå êîòîðûìè ïðèâîäèò ê ãèáåëè íàñåêîìûõ, îò ñàïðîòðîôíûõ. Ñâÿçü ìåæäó âèäîâûì ñîñòàâîì ôóçàðèåâûõ ãðèáîâ è ñèñòåìàòè÷åñêèì ïîëîæåíèåì íàñåêîìûõ-õîçÿåâ ïîêà íå óñòàíîâëåíà. Òàê, Fusarium semitectum, èçîëèðîâàííûé èç ìåðòâûõ îñîáåé áàõ÷åâîé (õëîïêîâîé) òëè Aphis gossypii, âûçûâàë ãèáåëü æåëòîãî ÷àéíîãî òðèïñà Scirtothrips dorsalis, îðàíæåðåéíîãî ïðîçðà÷íîãî êëåùà Polyphagotarsonemus latus, òëè Ceratavacuna lanigera, áåëîêðûëêè Aleyrodicus disperses, òàáà÷íîé áåëîêðûëêè Bemisia tabaci è êëåùà Aceria guerroronis (3). Îäíàêî çàðàæåíèå ýòèì ãðèáîì íå ïðèâîäèëî ê ãèáåëè ãóñåíèö âðåäíûõ ÷åøóåêðûëûõ è ëè÷èíîê áîæüåé êîðîâêè Menochilus sexmaculatus, à òàêæå õèùíîãî êëåùà Amblysius ovalis, ëè÷èíî÷íîãî ïàðàçèòîèäà Goniozus nephantidis è ãóñåíèö òóòîâîãî øåëêîïðÿäà Bombix mori. Êðîìå òîãî, îí íå âëèÿë íà ðîñò íàâîçíîãî ÷åðâÿ Eisenia foetida è åãî ñïîñîáíîñòü îáðàçîâûâàòü êîìïîñò. Ïîêàçàíà ïàòîãåííîñòü ãðèáà F. verticillioides, èçîëèðîâàííîãî èç ëè÷èíîê è èìàãî ãèãàíòñêîé ñàðàí÷è Tropidacris collaris, êîòîðàÿ îáèòàåò â Öåíòðàëüíîé Àìåðèêå (4). Èç òðóïîâ ëè÷èíîê ïøåíè÷íîãî ñòåáëåâîãî ïèëèëüùèêà Cephus cinctus áûëè âûäåëåíû ðàçëè÷íûå âèäû ãðèáîâ Fusarium, â òîì ÷èñëå F. pseudograminearum, F. culmorum, F. avenaceum, F. equiseti è F. acuminatum (5). Èçîëÿòû Fusarium spp. âûçûâàëè ãèáåëü êàê òîïèêàëüíî îáðàáîòàííûõ äèàïàóçèðóþùèõ ëè÷èíîê â ëàáîðàòîðíûõ îïûòàõ, òàê è ðàçâèâàþùèõñÿ ëè÷èíîê, êîòîðûå ïèòàëèñü íà èíôèöèðîâàííûõ ðàñòåíèÿõ ïøåíèöû â óñëîâèÿõ òåïëèöû. Õàðàêòåð êîððåëÿöèè ìåæäó ñòåïåíüþ îáåñöâå÷èâàíèÿ èíòåãóìåíòà, ðîñòîì ãèô ãðèáîâ è ñìåðòíîñòüþ ëè÷èíîê ïèëèëüùèêà ñâèäåòåëüñòâîâàë î òîì, ÷òî èçîëÿòû Fusarium spp. àêòèâíî èíôèöèðîâàëè ëè÷èíîê è âûçûâàëè èõ ãèáåëü, à íå êîëîíèçèðîâàëè òðóïû êàê âòîðè÷íûå ïàòîãåíû. Ãðèá F. graminearum áûë âûäåëåí èç èìàãî æóêà áëåñòÿíêè ÷åòûðåõòî÷å÷íîé Glischrochilus quadrisignatus, ëè÷èíîê äëèííîóñîé áëîøêè Diabrotica longicornis, òðèïñîâ è êóçíå÷èêîâ (6-10). Ò.Ê. Êàëüâèø (11) èçîëèðîâàë F. sporotrichiella var. poae è F. semitectum èç ãóñåíèö íåïàðíîãî øåëêîïðÿäà Lymantria dispar â Ñèáèðè è Êàçàõñòàíå. Â.À. Ñìèðíîâ (12) 14 âûäåëèë F. larvarum èç òëåé Adelges piceae íà ïèõòå áàëüçàìè÷åñêîé â Êàíàäå, è èçîëÿòû ãðèáà âûçûâàëè çíà÷èòåëüíóþ ñìåðòíîñòü òëè.  ïîëåâûõ ýêñïåðèìåíòàõ îòìå÷àëè êðàéíå âûñîêóþ (100 %) ñìåðòíîñòü êîðè÷íåâîé öèêàäêè Nilaparvata lugens ïðè èíîêóëÿöèè íàñåêîìûõ ãðèáîì F. oxysporum (13, 14). Èíîêóëÿöèÿ ýòèìè èçîëÿòàìè F. oxysporum ðèñà, õëîï÷àòíèêà èëè òîìàòà íå ïðèâîäèëà ê ðàçâèòèþ çàáîëåâàíèÿ ðàñòåíèé (15). Ñëåäóåò äîáàâèòü, ÷òî ìåõàíèçì ïðîíèêíîâåíèÿ ýíòîìîïàòîãåííûõ ïðåäñòàâèòåëåé ðîäà Fusarium â îðãàíèçì õîçÿèíà ïðèíöèïèàëüíî îòëè÷àåòñÿ îò òàêîâîãî ó äðóãèõ ãðóïï ãðèáîâ è ïðîèñõîäèò ÷åðåç åñòåñòâåííûå îòâåðñòèÿ (ðîòîâûå îðãàíû, äûõàëüöà, ÿéöåêëàä) èëè ðàíû, à íå ÷åðåç êóòèêóëó íàñåêîìîãî (16, 17). Íåëåòàëüíûå àíòàãîíèñòè÷åñêèå âçàèìîîòíîøåíèÿ ìåæäó âèäàìè Fusarium è íàñåêîìûìè ñâÿçàíû ñ ïðèñóòñòâèåì ó ãðèáîâ ËÎÑ. Ýòè âåùåñòâà ñëóæàò ñèãíàëüíûìè ìîëåêóëàìè (èíôîõåìèêè èëè ñåìèîõåìèêè) è ïðîÿâëÿþò ïî îòíîøåíèþ ê íàñåêîìûì ñâîéñòâà ôåðîìîíîâ, àëëîìîíîâ, êàéðîìîíîâ è äð. Íàñåêîìûå ìîãóò âîñïðèíèìàòü èõ êàê àòòðàêòèâíûå, ðåïåëëåíòíûå, äåòåððåíòíûå èëè íåéòðàëüíûå (18). Èçó÷åíèå ËÎÑ ãðèáîâ ðîäà Fusarium áûëî íà÷àòî â êîíöå 1990-õ ãîäîâ, ïóáëèêàöèé ïî ýòîé òåìå ïîêà íåìíîãî. Óñòàíîâëåíî, ÷òî ËÎÑ Fusarium âêëþ÷àþò áîëüøîå ÷èñëî êîìïîíåíòîâ, îòíîñÿùèõñÿ ê ðàçëè÷íûì ãðóïïàì ñîåäèíåíèé (19-24). Âòîðè÷íûå ìåòàáîëèòû ãðèáîâ (â òîì ÷èñëå ËÎÑ) èãðàþò âàæíóþ ðîëü â èõ æèçíåäåÿòåëüíîñòè. ËÎÑ ãðèáîâ ìîãóò âûïîëíÿòü ôóíêöèþ ðåïåëëåíòîâ, ñèãíàëèçèðóÿ íàñåêîìûì î «íåáëàãîïðèÿòíîì» ïèùåâîì ñóáñòðàòå (àíòàãîíèñòè÷åñêèå âçàèìîîòíîøåíèÿ). Íàïðèìåð, ãóñåíèöû áàáî÷êè-îãíåâêè Eldana saccharina èçáèðàòåëüíî ðåàãèðîâàëè íà çåðíà êóêóðóçû, èíîêóëèðîâàííûå èçîëÿòàìè ãðèáîâ F. sacchari, F. pseudonymagai è F. proliferatum. Âûÿâëåíî ðåïåëëåíòíîå âëèÿíèå íà èìàãî èçîëÿòîâ F. sacchari.  òî æå âðåìÿ ââåäåíèå òàêèõ ãðèáîâ â ðàöèîí ãóñåíèö îêàçûâàëî íåãàòèâíûé ýôôåêò íà èõ ðàçâèòèå (25). Ñëåäóåò îòìåòèòü, ÷òî òî÷íûé õèìè÷åñêèé ñîñòàâ ðåïåëëåíòíûõ ËÎÑ ôóçàðèåâûõ ãðèáîâ ïîêà íå óñòàíîâëåí. Èçâåñòíî, ÷òî ãðèáû ðîäà Fusarium ñëóæàò ïðîäóöåíòàìè âòîðè÷íûõ ìåòàáîëèòîâ, âûñîêîòîêñè÷íûõ äëÿ òåïëîêðîâíûõ îðãàíèçìîâ.  íåêîòîðûõ ñëó÷àÿõ ïðèñóòñòâèå ýòèõ ñîåäèíåíèé îêàçûâàåò ïîõîæåå äåéñòâèå è íà íàñåêîìûõ, ÷òî ìîæíî ðàññìàòðèâàòü â êà÷åñòâå åùå îäíîãî ïðîÿâëåíèÿ àíòàãîíèñòè÷åñêèõ îòíîøåíèé ìåæäó íèìè è ãðèáàìè ïðè èñïîëüçîâàíèè îäíîãî ïèùåâîãî ñóáñòðàòà (26). Íàïðèìåð, ðàçâèòèå æóêîâ àìáàðíîãî äîëãîíîñèêà Sitophilus granarium íà çåðíîâêàõ, çàðàæåííûõ ôóçàðèåâûìè ãðèáàìè, ïðèâîäèëî ê óâåëè÷åíèþ èõ ñìåðòíîñòè è ñíèæåíèþ ðåïðîäóêòèâíîãî ïîòåíöèàëà (27). Ïîêàçàíî, ÷òî T-2 òîêñèí îêàçûâàë ñëàáîå òîêñè÷åñêîå äåéñòâèå íà æóêîâ ìàëîãî ìó÷íîãî õðóùàêà Tribolium confusum (28). Ñèìáèîòè÷åñêèå îòíîøåíèÿ ìåæäó âèäàìè ðîäà Fusarium è íàñåêîìûìè õàðàêòåðèçóþòñÿ áîëüøèì ðàçíîîáðàçèåì è âêëþ÷àþò êàê âçàèìîâûãîäíîå ñîñóùåñòâîâàíèå (ìóòóàëèçì), òàê è êîììåíñàëèçì. Êëàññè÷åñêèé ïðèìåð ñèìáèîòè÷åñêèõ îòíîøåíèé ìåæäó ãðèáàìè è íàñåêîìûìè — ó÷àñòèå ïîñëåäíèõ â ðàñïðîñòðàíåíèè ñïîð ãðèáîâ (ýíòîìîõîðèÿ). Äëÿ ïðåäñòàâèòåëåé ðîäà Fusarium õàðàêòåðíî ôîðìèðîâàíèå äâóõ òèïîâ ñïîð áåñïîëîãî ðàçìíîæåíèÿ — ìàêðî- è ìèêðîêîíèäèé. Áûñòðîå è èçáûòî÷íî áîëüøîå êîëè÷åñòâî îáðàçóþùèõñÿ êîíèäèé (r-ñòðàòåãèÿ) ðàññ÷èòàíî íà èõ íèçêóþ âûæèâàåìîñòü â îêðóæàþùåé ñðåäå. Çàõâàò è ïåðåíîñ ñïîð ãðèáà íàñåêîìûì âîçìîæåí êàê ïðè ñëó÷àéíîì ñîïðèêîñíîâåíèè åãî òåëà ñî ñïîðîíîñÿùèìè ñòðóêòóðàìè ãðèáà, òàê è ïðè 15 ïîåäàíèè ãðèáíîãî ìèöåëèÿ âìåñòå ñ ïèùåé è âûäåëåíèè íå ïîääàþùèõñÿ ïåðåâàðèâàíèþ ñïîð ñ ôåêàëèÿìè. Õîðîøî èçâåñòåí ôàêò ïåðåíîñà íàñåêîìûìè-îïûëèòåëÿìè ñïîð ãðèáîâ âìåñòå ñ ïûëüöîé ðàñòåíèé. Íàïðèìåð, çíà÷èòåëüíûå ýêîíîìè÷åñêèå ïîòåðè ñâÿçàíû ñ ãíèëüþ ôèã, âûçûâàåìîé ôóçàðèåâûìè ãðèáàìè (29). Ñïîðû ãðèáîâ F. verticillioides è F. solani ðàñïðîñòðàíÿþòñÿ õàëüöèäîèäíûì íàåçäíèêîì Blastophaga psenes — ñïåöèàëèçèðîâàííûì îïûëèòåëåì èíæèðîâûõ äåðåâüåâ (30). Ïîêàçàíî, ÷òî ïðîõîæäåíèå ñïîð ÷åðåç êèøå÷íèê êîìíàòíîé ìóõè Musca domestica íå âëèÿëî íà ìîðôîëîãèþ è æèçíåñïîñîáíîñòü Fusarium spp. è íåêîòîðûõ äðóãèõ ãðèáîâ (31). F. verticillioides âûäåëÿëè èç ôåêàëèé íàñåêîìûõ, à òàêæå ÷àñòåé êóêóðóçû, ïîâðåæäåííîé õëîïêîâîé ñîâêîé Helicoverpa armigera è êóêóðóçíûì ìîòûëüêîì Ostrinia nubilalis (32). Ñèìáèîòè÷åñêèå ñâÿçè ìåæäó ãðèáîì è íàñåêîìûì-îïûëèòåëåì îñóùåñòâëÿþòñÿ ñ ó÷àñòèåì ðàçëè÷íûõ ìåõàíèçìîâ. Ãðèáû ìîãóò îáðàçîâûâàòü ñïåöèôè÷åñêèé ïî îêðàñêå ìèöåëèé, ñîçäàþùèé äîïîëíèòåëüíûé îïòè÷åñêèé ñòèìóë äëÿ íàñåêîìûõ (ïðèìåðû äëÿ ôóçàðèåâûõ ãðèáîâ ïîêà îòñóòñòâóþò). Íåêîòîðûå ôèòîïàòîãåííûå ãðèáû âûäåëÿþò ëåòó÷èå õèìè÷åñêèå àòòðàêòàíòû è ïèòàòåëüíûå âåùåñòâà (óãëåâîäû), ïðèâëåêàþùèå íàñåêîìûõ.  ñâîþ î÷åðåäü, ðàñòèòåëüíîÿäíûå íàñåêîìûå, ïîâðåæäàÿ ðàñòåíèå, îáëåã÷àþò äîñòóï ôèòîïàòîãåííûõ ãðèáîâ ê åãî òêàíÿì. Êðîìå òîãî, ïàòîãåííûå ãðèáû ìîãóò âëèÿòü íà êà÷åñòâî è êîëè÷åñòâî ïèòàòåëüíûõ âåùåñòâ, äîñòóïíûõ â ðàñòèòåëüíûõ òêàíÿõ è íåêòàðå, îêàçûâàÿ òåì ñàìûì ïðÿìîå èëè êîñâåííîå äåéñòâèå íà ïîâåäåíèå è ðàçâèòèå íàñåêîìûõ.  îòëè÷èå îò ðåïåëëåíòíûõ ËÎÑ ôóçàðèåâûõ ãðèáîâ, êîòîðûå ñëóæàò íîñèòåëÿìè èíôîðìàöèîííûõ ñèãíàëîâ ïðè àíòàãîíèñòè÷åñêèõ îòíîøåíèÿõ Fusarium ñ íàñåêîìûìè, õèìè÷åñêèé ñîñòàâ íåêîòîðûõ àòòðàêòèâíûõ ËÎÑ óæå èçâåñòåí. Æóê-áëåñòÿíêà Carpophilus humeralis ñëóæèò âåêòîðîì ïðè ðàñïðîñòðàíåíèè ãðèáà F. verticillioides (= F. moniliforme), êîòîðûé âûçûâàåò çàáîëåâàíèå ïî÷àòêîâ êóêóðóçû è ïðîäóöèðóåò ìèêîòîêñèíû ôóìîíèçèíû (33). Ñðåäè ëåòó÷èõ ñîåäèíåíèé, âûäåëÿåìûõ F. verticillioides, áûëè îáíàðóæåíû ïÿòü ñïèðòîâ (ýòàíîë, 1-ïðîïàíîë, 2-ìåòèë-1-ïðîïàíîë, 3-ìåòèë-1-áóòàíîë è 2-ìåòèë-1-áóòàíîë), àöåòàëüäåãèä è ýòèëàöåòàò, ïÿòü ôåíîëüíûõ êîìïîíåíòîâ, à òàêæå ñåðèÿ ïîêà íå èäåíòèôèöèðîâàííûõ ñåñêâèòåðïåíîâûõ óãëåâîäîðîäîâ è îäíî íåèçâåñòíîå âåùåñòâî, âåðîÿòíî, 10-óãëåðîäíûé êåòîíîí (21). Íàáëþäåíèÿ, ïðîâåäåííûå â îëüôàêòîìåòðå, ïîêàçàëè, ÷òî àòòðàêòèâíûé ýôôåêò ó æóêîâ ñâÿçàí ñ ïðèñóòñòâèåì â ïðèìàíêå ñïèðòîâ, àöåòàëüäåãèäà è ýòèëàöåòàòà, à íå ôåíîëîâ (èñïîëüçîâàëèñü òàêæå ñèíòåòè÷åñêèå àíàëîãè ýòèõ âåùåñòâ). Ïîêàçàíî, ÷òî ãðèáû âûäåëÿþò ËÎÑ, êîòîðûå âîñïðèíèìàþòñÿ êàê àòòðàêòàíòû æóêàìè, ïîâðåæäàþùèìè õðàíÿùååñÿ çåðíî (34) è ñóõîôðóêòû (35), à òàêæå íåñêîëüêèìè âèäàìè ÷åøóåêðûëûõ (36). Ïèòàíèå çåðíàìè êóêóðóçû, çàðàæåííîé èçîëÿòàìè F. pseudonymagai, ïðèâîäèëî ê óâåëè÷åíèþ ÷èñëà ãóñåíèö áàáî÷êè-îãíåâêè Eldana saccharina, äîñòèãøèõ îêóêëèâàíèÿ, è ê çàìåòíîìó óâåëè÷åíèþ èõ ðàçìåðîâ ïî ñðàâíåíèþ ñ âàðèàíòàìè, ãäå îíè ïèòàëèñü íåçàðàæåííûìè èëè çàðàæåííûìè äðóãèìè âèäàìè ãðèáîâ çåðíàìè. Ñàìûé áëàãîïðèÿòíûé äëÿ ãóñåíèö èçîëÿò âûçûâàë è íàèáîëüøèé àòòðàêòèâíûé ýôôåêò ïî îòíîøåíèþ ê áàáî÷êàì (25).  ñâîþ î÷åðåäü, ïèòàþùèåñÿ íàñåêîìûå ñïîñîáñòâóþò ïåðåíîñó ñïîð ãðèáîâ íà íåçàðàæåííûå çåðíîâêè. Òî åñòü ËÎÑ ãðèáîâ, ðàçâèâàþùèõñÿ íà ðàñòèòåëüíîì ìàòåðèàëå, ìîãóò äàâàòü íàñåêîìûì èíôîðìàöèþ î åãî ïðèãîäíîñòè äëÿ ïèòàíèÿ è îòêëàäêè ÿèö. Ãðèáíûå ËÎÑ òàêæå èñïîëüçóþòñÿ õèùíûìè è ïàðàçèòè÷åñêèìè 16 íàñåêîìûìè â êà÷åñòâå îðèåíòèðà äëÿ îáíàðóæåíèÿ æåðòâû èëè õîçÿèíà.  ñèñòåìå àìáàðíûé äîëãîíîñèê—ãðèá—ïàðàçèòîèä çàñåëåíèå çåðíà äîëãîíîñèêîì âåäåò ê ðîñòó ïëåñåíè, êîòîðàÿ îáðàçóåò ëîêàëüíûå ãîðÿ÷èå ïÿòíà, âûçûâàÿ âûñîêóþ ñìåðòíîñòü íàñåêîìîãî.  îëüôàêòîìåòðå çàïëåñíåâåëîå çåðíî è ïîêðûòûå ïëåñåíüþ ôåêàëèè íàñåêîìîãî îòïóãèâàëè ñàìîê ëè÷èíî÷íîãî ïàðàçèòîèäà Lariophagus distinguendus. Îñíîâíûì ËÎÑ, âûäåëÿþùèìñÿ èç çàïëåñíåâåëûõ ñóáñòðàòîâ, îêàçàëñÿ 1-îêòåí-3-îë (èëè ãðèáíîé ñïèðò), â ìåíüøåì êîëè÷åñòâå îáíàðóæåíû 3-îêòàíîí è 3-îêòàíîë. Ïðè âûñîêèõ äîçàõ (> 300 íã) 1-îêòåí-3-îë ñëóæèë ðåïåëëåíòîì äëÿ L. distinguendus, íî íèçêèå äîçû (30 íã) îêàçàëèñü íåéòðàëüíûìè. Èññëåäîâàòåëè ïðåäïîëàãàþò, ÷òî íåïðèâëåêàòåëüíîñòü ïëåñåíè äëÿ ïàðàçèòîèäà äàåò åìó âîçìîæíîñòü èçáåãàòü ïðîñòðàíñòâ, ãäå ñêëàäûâàþòñÿ íåáëàãîïðèÿòíûå äëÿ æèçíåäåÿòåëüíîñòè õîçÿèíà óñëîâèÿ (37). Êðîìå âûäåëåíèÿ ñîáñòâåííûõ ËÎÑ, âèäû ðîäà Fusarium èçâåñòíû â êà÷åñòâå èíäóêòîðîâ ëåòó÷èõ âòîðè÷íûõ ìåòàáîëèòîâ, êîòîðûå ðàñòåíèÿ îáðàçóþò â îòâåò íà ïîâðåæäàþùåå äåéñòâèå ãðèáîâ (38-42). Îïèñàíû ïðèìåðû, êîãäà ïàòîãåí ñòèìóëèðóåò ðàñòåíèå âûäåëÿòü ËÎÑ, ïðèâëåêàòåëüíûå äëÿ íàñåêîìîãî — ïåðåíîñ÷èêà ñïîð ãðèáà (18). Ðàçëè÷íûå àñïåêòû èíäóöèðóåìûõ ôèòîôàãàìè ðåàêöèé ðàñòåíèé, â òîì ÷èñëå ñ ó÷àñòèåì ôèòîïàòîãåííûõ ãðèáîâ, ðàññìàòðèâàþòñÿ â îïóáëèêîâàííîé íåäàâíî ìîíîãðàôèè (43). ßðêèì ïðèìåðîì ìóòóàëèçìà ñëóæàò îòíîøåíèÿ ãðèáîâ Fusarium ñ æóêàìè-äðåâîòî÷öàìè ïîäñåìåéñòâà Scolytinae (Curculionidae), êîòîðûå ïåðåíîñÿò èõ ñïîðû. Èçâåñòíî, ÷òî ãðèáû óìåþò ñèíòåçèðîâàòü íåçàìåíèìûå ñòåðèíû, âõîäÿùèå â ñîñòàâ êëåòî÷íûõ ìåìáðàí è ñëóæàùèå ïðåäøåñòâåííèêàìè ñòåðîèäíûõ ãîðìîíîâ, æèçíåííî íåîáõîäèìûõ äëÿ íîðìàëüíîãî ðàçâèòèÿ íàñåêîìûõ (44).  2009 ãîäó ïîÿâèëîñü ñîîáùåíèå î âûÿâëåíèè íîâîãî âðåäîíîñíîãî çàáîëåâàíèÿ äåðåâüåâ (àâîêàäî, äóáà âå÷íîçåëåíîãî, êëåíà ÿñåíåëèñòíîãî è äð.) â Êàëèôîðíèè è â Èçðàèëå, êîòîðîå âûçûâàåòñÿ ñèìáèîòè÷åñêèì êîìïëåêñîì æóê Euwallacea sp.—ãðèá Fusarium sp. Ñïîðû ãðèáà ðàñïðîñòðàíÿþò ñàìêè æóêà. Èíîêóëÿöèÿ îñóùåñòâëÿåòñÿ âî âðåìÿ ÿéöåêëàäêè â íèøàõ ãàëåðåé, êîòîðûå ñàìêè âûãðûçàþò âíóòðè äðåâåñèíû, è â äàëüíåéøåì ìèöåëèé ãðèáà ñëóæèò åäèíñòâåííûì ïèòàòåëüíûì ñóáñòðàòîì ëè÷èíîê (45). Ðàñïðîñòðàíåíèå Fusarium â ýòîì ñëó÷àå ïðèâîäèò ê âîçíèêíîâåíèþ ôóçàðèîçíîé ñóõîâåðøèííîñòè äðåâåñíûõ ðàñòåíèé. Íàïðèìåð, â áîòàíè÷åñêèõ ñàäàõ Êàëèôîðíèè èç 335 âèäîâ äðåâåñíûõ ðàñòåíèé ó 207 (62 %) áûëè îáíàðóæåíû ïðèçíàêè ïîâðåæäåíèÿ æóêîì, ó 54 % ïîâðåæäåííûõ ðàñòåíèé âûÿâëåíû ñëåäû ãðèáíîé èíôåêöèè (46). Êîìïëåêñ ãðèáîâ F. solani òåñíî ñâÿçàí ñ æóêàìè-äðåâîòî÷öàìè Xyleborus ferrugineus. Ñàìêè îòêëàäûâàþò ÿéöà íà ãðèáû, è ëè÷èíêè ïîëó÷àþò ýðãîñòåðîë, íåîáõîäèìûé äëÿ îêóêëèâàíèÿ (47). Ïèòàíèå çåðíîì ðèñà è ïøåíèöû, çàðàæåííûì F. moniliforme, ñòèìóëèðîâàëî ðîñò ó ëè÷èíîê áóëàâîóñîãî ìó÷íîãî õðóùàêà Tribolium castaneum (48). Ïðè ñêàðìëèâàíèè ëè÷èíêàì ìàëîãî ìó÷íîãî õðóùàêà T. confusum çåðíîïðîäóêòîâ, çàðàæåííûõ F. roseum var. graminearum, ÷èñëåííîñòü íàñåêîìûõ óâåëè÷èâàëàñü (49). Îáà âèäà õðóùàêà ìîãóò ðàññìàòðèâàòüñÿ â êà÷åñòâå ïîòåíöèàëüíûõ ïåðåíîñ÷èêîâ ñïîð ôóçàðèåâûõ ãðèáîâ. Ïîâðåæäåíèÿ ðàñòåíèé íàñåêîìûìè ñïîñîáñòâóþò ïðîíèêíîâåíèþ ãðèáîâ â òêàíü è ðàçâèòèþ çàáîëåâàíèÿ. Óñòàíîâëåíà ñâÿçü ìåæäó F. oxysporum (âûçûâàåò óâÿäàíèå ëþöåðíû), F. roseum Link (ýòèîëîãè÷åñêèé ôàêòîð êðàñíîé êîðíåâîé ãíèëè êëåâåðà) è ïëîäîâûì êîìàðèêîì Bradysia sp., ëè÷èíêè êîòîðîãî ïîâðåæäàþò êîðíè, îñíîâàíèÿ ñòåáëåé è ïîáåãè ðàñòå17 íèé (14). Íàëè÷èå ýòèõ íàñåêîìûõ âûçûâàëî âñïûøêè çàáîëåâàíèé, äîñòîâåðíî óñèëèâàëèñü ïðîÿâëåíèÿ ôóçàðèîçíûõ óâÿäàíèé è ãíèëè êîðíåé ðàñòåíèé. Õîðîøî èçâåñòåíà çàâèñèìîñòü ðàçâèòèÿ ôóçàðèîçà ïî÷àòêîâ îò ïîâðåæäåíèÿ êóêóðóçíûì ìîòûëüêîì è õëîïêîâîé ñîâêîé (50, 51). Îñîáè îðàíæåâîãî çëàêîâîãî êîìàðèêà Sitodiplosis mosellana, ïðåäâàðèòåëüíî çàðàæåííûå ãðèáîì F. graminearum, âûçûâàëè ôóçàðèîç êîëîñà (52, 53). Ïðè èíôèöèðîâàíèè ðàñòåíèé êóêóðóçû ãðèáîì F. verticillioides óñèëèâàåòñÿ èõ ïîðàæåíèå áàáî÷êîé-îãíåâêîé Eldana saccharina. Òàêèå ðàñòåíèÿ îêàçàëèñü ïðèâëåêàòåëüíåå äëÿ ÿéöåêëàäêè, ÷åì çäîðîâûå, à ãóñåíèöû íà íèõ ëó÷øå âûæèâàëè è ðàçâèâàëèñü (54, 55). Ïðèìåðîì îäíîñòîðîííèõ ñèìáèîòè÷åñêèõ âçàèìîîòíîøåíèé (êîììåíñàëèçì) ìîæåò áûòü ðàñïðîñòðàíåíèå F. poae íàñåêîìûìè, íàïðèìåð òðèïñàìè (Limothrips), òëÿìè (Sitobion), êëåùàìè (Siteroptes graminum, S. avenae). Âåðîÿòíî, ñëàäêîâàòûé çàïàõ, õàðàêòåðíûé äëÿ ìíîãèõ êóëüòóð ýòîãî ãðèáà, ïðèâëåêàåò ÷ëåíèñòîíîãèõ (8, 56, 57). Ó÷èòûâàÿ ñëàáóþ ïàòîãåííîñòü, õàðàêòåðíóþ äëÿ øòàììîâ ýòîãî âèäà, è øèðîêîå ïîâñåìåñòíîå ðàñïðîñòðàíåíèå íà çåðíîâûõ êóëüòóðàõ (58), ëîãè÷íî ïðåäïîëîæèòü ó÷àñòèå íàñåêîìûõ êàê ôàêòîðà ïåðåäà÷è èíôåêöèè îò îäíîãî ðàñòåíèÿ íà äðóãîå èëè ñ îäíîé åãî ÷àñòè íà äðóãóþ. Îáùàÿ êàðòèíà áèîöåíîòè÷åñêèõ ñâÿçåé ìåæäó ãðèáàìè è íàñåêîìûìè áûâàåò äîñòàòî÷íî ñëîæíîé, îñîáåííî åñëè îíè èñïîëüçóþò îäèí è òîò æå ïèùåâîé ñóáñòðàò. Õîðîøèì ïðèìåðîì ñëóæèò ñèñòåìà «çåðíîâêè çëàêîâ—ãðèáû Fusarium—çåðíîâûå äîëãîíîñèêè». Ãðèáû Fusarium è çåðíîâûå æóêè (íàïðèìåð, ðèñîâûé äîëãîíîñèê Sitophilus zeamais) ïðè áëàãîïðèÿòíûõ óñëîâèÿõ çàñåëÿþò çåðíîâêè çëàêîâ êàê â ïîëåâûõ óñëîâèÿõ, òàê è â ïåðèîä õðàíåíèÿ è ñîñóùåñòâóþò äëèòåëüíîå âðåìÿ. Áèîöåíîòè÷åñêèå ñâÿçè ìåæäó ãðèáàìè ðîäà Fusarium è çåðíîâûìè æóêàìè. Àíòàãîíèñòè÷åñêèå îòíîøåíèÿ ìåæäó ãðèáàìè è æóêàìè, âèäèìî, îïðåäåëÿþòñÿ óòèëèçàöèåé îáùåãî è î÷åíü ñïåöèôè÷åñêîãî ïèùåâîãî ñóá18 ñòðàòà — çåðíîâîê çëàêîâûõ êóëüòóð è îêàçûâàþò îäíîíàïðàâëåííîå äåéñòâèå (ðèñ.). Ìåõàíèçìû òàêèõ îòíîøåíèé ìîãóò áûòü ñâÿçàíû ñ âûäåëåíèåì ãðèáàìè ðåïåëëåíòíûõ è/èëè òîêñè÷íûõ ñîåäèíåíèé, ÿâëÿþùèõñÿ ÷àñòüþ ïðîäóöèðóåìûõ èìè âòîðè÷íûõ ìåòàáîëèòîâ (59). Ïðè÷åì â ïåðâîì ñëó÷àå âçàèìîäåéñòâèå ìåæäó ãðèáàìè è æóêàìè — äèñòàíòíîå, èìåþùåå ôîðìó îëüôàêòîðíîãî ñòèìóëà, âî âòîðîì — êîíòàêòíîå. Âåðîÿòíûé ðåçóëüòàò ïîäîáíîãî àíòàãîíèçìà — èçáåãàíèå æóêàìè çåðíîâîê, çàðàæåííûõ ãðèáîì. Ïîòðåáëåíèå çåðíîâîê, ñîäåðæàùèõ òîêñè÷íûå ìåòàáîëèòû ãðèáà, ìîæåò ïðèâîäèòü ê ñíèæåíèþ æèçíåñïîñîáíîñòè è ïëîäîâèòîñòè íàñåêîìûõ, à òàêæå ê óõóäøåíèþ óñëîâèé ðàçâèòèÿ ïîòîìñòâà. Íåëüçÿ èñêëþ÷èòü, ÷òî æóêè, çàñåëèâøèå çåðíîâêè (íàïðèìåð, ëè÷èíêè), âûäåëÿþò âåùåñòâà, èìåþùèå ôóíãèöèäíûå ñâîéñòâà è ïîäàâëÿþùèå ðàçâèòèå ãðèáîâ, êàê ýòî èçâåñòíî ó íåêîòîðûõ äðóãèõ íàñåêîìûõ (60). Ñèìáèîòè÷åñêèå îòíîøåíèÿ ïðåäïîëàãàþò âçàèìîâûãîäíûé ðåçóëüòàò äëÿ îáîèõ ïàðòíåðîâ. Íàïðèìåð, ïîä âîçäåéñòâèåì íàñåêîìûõ óâåëè÷èâàåòñÿ âëàæíîñòü è ïîâûøàåòñÿ òåìïåðàòóðà õðàíÿùåãîñÿ çåðíà è ñîçäàþòñÿ áëàãîïðèÿòíûå óñëîâèÿ äëÿ ðàçâèòèÿ ôóçàðèåâûõ ãðèáîâ è èõ ìåòàáîëè÷åñêîé àêòèâíîñòè (61). Êðîìå òîãî, âðåäèòåëåé çàïàñîâ, êîòîðûå ïèòàþòñÿ çåðíîì è ïðèñóòñòâóþùèìè â íåì ãðèáàìè, ìîæíî ðàññìàòðèâàòü â êà÷åñòâå ïîòåíöèàëüíûõ ïåðåíîñ÷èêîâ ïðîïàãóë ãðèáà íà íåçàñåëåííûå ñóáñòðàòû.  ñâîþ î÷åðåäü, ãðèáû ïðîäóöèðóþò àòòðàêòèâíûå ëåòó÷èå ìåòàáîëèòû è íåòîêñè÷íûå ïèòàòåëüíûå ñîåäèíåíèÿ (íàïðèìåð, ñòåðîèäíûå ãîðìîíû), îáëåã÷àþùèå íàñåêîìûì ïîèñê ïèùè èëè íåîáõîäèìûå äëÿ èõ ðàçâèòèÿ. Èíòåðåñíî îòìåòèòü, ÷òî äåéñòâèå ýòèõ ìåõàíèçìîâ áîëåå âûðàæåíî ó ñëàáîïàòîãåííûõ âèäîâ ðîäà Fusarium.  íàøèõ îïûòàõ îò÷åòëèâàÿ òåíäåíöèÿ àòòðàêòèâíîãî äåéñòâèÿ áûëà âûÿâëåíà ó øòàììîâ ñëàáîïàòîãåííûõ âèäîâ ôóçàðèåâûõ ãðèáîâ F. langsethiae è F. ðîàå â îòíîøåíèè ðèñîâîãî äîëãîíîñèêà (62, 63). Î÷åâèäíûé ðåçóëüòàò òàêîãî âçàèìîâûãîäíîãî ïàðòíåðñòâà — ðàñøèðåíèå àðåàëîâ âèäîâ (èëè ïîääåðæàíèå èõ ãðàíèö â ïðåäåëàõ îïòèìóìà), óâåëè÷åíèå ïëîòíîñòè ïîïóëÿöèé è ïîâûøåíèå æèçíåñïîñîáíîñòè. Òàêèì îáðàçîì, êîýâîëþöèÿ ãðèáîâ ðîäà Fusarium è íàñåêîìûõ ïðèâåëà ê ôîðìèðîâàíèþ ðàçíîîáðàçíûõ ôîðì âçàèìîîòíîøåíèé, êîòîðûå â êîíå÷íîì ðåçóëüòàòå îáåñïå÷èâàþò ñîñóùåñòâîâàíèå ýòèõ îðãàíèçìîâ â óñòîé÷èâûõ ñàìîðåãóëèðóþùèõñÿ ýêîñèñòåìàõ. Àíòàãîíèñòè÷åñêèå âçàèìîîòíîøåíèÿ îêàçûâàþò îäíîíàïðàâëåííîå äåéñòâèå è ìîãóò èìåòü ëåòàëüíûå è íåëåòàëüíûå ïîñëåäñòâèÿ äëÿ íàñåêîìûõ.  îáîèõ ñëó÷àÿõ ýòà ôîðìà âçàèìîäåéñòâèÿ îñóùåñòâëÿåòñÿ ñ ïîìîùüþ ëåòó÷èõ (íàïðèìåð, ðåïåëëåíòû) è íåëåòó÷èõ (íàïðèìåð, ìèêîòîêñèíû) âòîðè÷íûõ ìåòàáîëèòîâ ãðèáîâ. Ñèìáèîòè÷åñêèå âçàèìîäåéñòâèÿ ìåæäó âèäàìè ðîäà Fusarium è íàñåêîìûìè õàðàêòåðèçóþòñÿ áîëüøèì ðàçíîîáðàçèåì è âêëþ÷àþò êàê âçàèìîâûãîäíîå ñîñóùåñòâîâàíèå (ìóòóàëèçì), òàê è êîììåíñàëèçì. Èçâåñòíà ðîëü íàñåêîìûõ â ðàñïðîñòðàíåíèè ñïîð ãðèáîâ Fusarium íà íåçàñåëåííûå ñóáñòðàòû. Ïîêàçàí àòòðàêòèâíûé ýôôåêò ëåòó÷èõ ìåòàáîëèòîâ ãðèáîâ, êîòîðûå ìîãóò âûïîëíÿòü ôóíêöèè ñèãíàëüíûõ ìîëåêóë ïðè âçàèìîäåéñòâèÿõ ãðèáîâ ñ äðóãèìè îðãàíèçìàìè.  ðÿäå ñëó÷àåâ ôóçàðèåâûå ãðèáû ñëóæàò èñòî÷íèêîì âåùåñòâ, æèçíåííî íåîáõîäèìûõ äëÿ íîðìàëüíîãî ðàçâèòèÿ íàñåêîìûõ. ÃÍÓ Âñåðîññèéñêèé ÍÈÈ çàùèòû ðàñòåíèé Ðîññåëüõîçàêàäåìèè, Ïîñòóïèëà â ðåäàêöèþ 26 íîÿáðÿ 2013 ãîäà 196608 Ðîññèÿ, ã. Ñàíêò-Ïåòåðáóðã—Ïóøêèí, ø. Ïîäáåëüñêîãî, 3, e-mail: [email protected], [email protected], [email protected], [email protected] 19 BIOLOGICAL RELATIONSHIPS BETWEEN Fusarium FUNGI AND INSECTS (review) T.Yu. Gagkaeva, I.V. Shamshev, O.P. Gavrilova, O.G. Selitskaya All-Russian Research Institute of Plant Protection, Russian Academy of Agricultural Sciences, 3, sh. Podbelskogo, St. Petersburg—Pushkin, 196608 Russia, e-mail [email protected], [email protected], [email protected], [email protected] Received November 26, 2013 Supported by Russian Foundation for Basic Research Abstract Fusarium fungi are characterized by high metabolic activity and adaptability. They colonize plant substrates, participate in the destruction of organic materials and in soil formation, and can be harmful to insects and mammalians. Fusarium fungi and insects coexist in different biotopes. Using molecular methods to classify 140 fungi samples, isolated from insects, K. O’Donnell et al. (2012) identified at least 23 strains or strain complexes as the Fusarium species. A host range specificity of Fusarium fungi to insects is no still found. The co-evolution of these groups of organisms has resulted in the emergence of various forms of interactions between them. The antagonistic forms have unilateral action, which can lead to lethal and non-lethal effects to insects. In both cases, this interaction involves volatile (e.g., repellent) and nonvolatile (e.g., mycotoxins) secondary metabolites of fungi. Volatile organic compounds (VOCs) are the signal molecules (infochemicals, semiochemicals) which act as the insect pheromones, allomones, kairomones, etc. The research of Fusarium fungi VOC was initiated in the late 1990s, and there are few scientific publications on the subject for the time being. The Fusarium VOCs were shown to consist of a large number of components from the different groups of chemical compounds. The mechanisms, due to which the infestation of entomopathogenic Fusarium fungi into host insects occurs, differ fundamentally from those observed in other fungi: the Fusarium fungi enter the insect’s body through natural orifices, such as mouthparts, spiracles, ovipositor, or wounds, but not through the cuticle. Symbiotic interactions between Fusarium fungi and insects are very diverse and include both mutually beneficial co-existence (mutualism) and forms that could be described as commensalism. The role of insects in spreading of Fusarium spores on new substrates is well documented. An attractive effect of biologically active volatile metabolites of fungi has been shown. In a number of cases Fusarium fungi are a source of substances that are essential to maintain normal growth and development of some insects. Obviously, the various forms of relationships between Fusarium fungi and insects are an important evolutionary factor. Diverse forms of interactions between Fusarium fungi and insects allow forming their sustainable self-regulating ecosystems. Special attention is paid to possible interactions of Fusarium fungi and insect pests in case of their sharing cereal grains as an alimentary substrate. Keywords: Fusarium fungi, insects, relationships, antagonistic and symbiotic interactions. 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