БИОЛОГИЯ ВЗАИМООТНОШЕНИЙ ГРИБОВ РОДА Fusarium И

ÑÅËÜÑÊÎÕÎÇßÉÑÒÂÅÍÍÀß ÁÈÎËÎÃÈß, 2014, ¹ 3, ñ. 13-23
ÓÄÊ 632:57.047:[582.288+595.7
ÁÈÎËÎÃÈß ÂÇÀÈÌÎÎÒÍÎØÅÍÈÉ ÃÐÈÁÎÂ ÐÎÄÀ 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,
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Ðàáîòà ïîääåðæàíà ïðîåêòîì ÐÔÔÈ ¹ 12-04-00927.
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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)
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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. Â 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
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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
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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.
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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]
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, ovi-
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positor, 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.
Þáèëåéíûå äàòû
Ïðîôåññîðó Çàóðó Ìóçàäèë îãëû Ãàñàíîâó èñïîëíÿåòñÿ 75 ëåò
Äîêòîð ñåëüñêîõîçÿéñòâåííûõ íàóê, ïðîôåññîð Çàóð Ìóçàäèë îãëû Ãàñàíîâ ðîäèëñÿ â 1939 ãîäó
â ã. Ãÿíäæå Àçåðáàéäæàíñêîé Ðåñïóáëèêè. Â 1961 ãîäó
îêîí÷èë ôàêóëüòåò ïëîäîîâîùåâîäñòâà è âèíîãðàäàðñòâà Àçåðáàéäæàíñêîãî ñåëüñêîõîçÿéñòâåííîãî
èíñòèòóòà (ÀçÑÕÈ, ã. Ãÿíäæà).  òå÷åíèå äâóõ ëåò
ðàáîòàë ìëàäøèì íàó÷íûì ñîòðóäíèêîì (â ñåêòîðå
ýðîçèè, çàòåì â Èíñòèòóòå áîòàíèêè ÀÍ Àçåðáàéäæàíà, ã. Áàêó).  1967 ãîäó ïîñëå çàâåðøåíèÿ ó÷åáû â àñïèðàíòóðå ÀçÑÕÈ îí óñïåøíî çàùèòèë êàíäèäàòñêóþ äèññåðòàöèþ â Ãðóçèíñêîì èíñòèòóòå ñóáòðîïè÷åñêîãî õîçÿéñòâà (ã. Ñóõóìè). Ñ ýòîãî ïåðèîäà
îí íà÷àë íàó÷íî-ïåäàãîãè÷åñêóþ äåÿòåëüíîñòü íà êàôåäðå ïëîäîîâîùåâîäñòâà (íûíå ñàäîâîäñòâà) ÀçÑÕÈ,
ãäå ïðîäîëæàåò ðàáîòàòü âñå ýòî âðåìÿ. Â 1971 ãîäó
åìó ïðèñóæäåíî ó÷åíîå çâàíèå äîöåíòà, â 1982 ãîäó åãî èçáèðàþò çàâåäóþùèì
êàôåäðîé, â 1986 ãîäó — äåêàíîì ôàêóëüòåòà. Â 1991 ãîäó Çàóð Ìóçàäèë îãëû
Ãàñàíîâ çàùèòèë äîêòîðñêóþ äèññåðòàöèþ íà òåìó «Íàó÷íûå îñíîâû òåõíîëîãèè âîçäåëûâàíèÿ ñóáòðîïè÷åñêîé õóðìû â Àçåðáàéäæàíå».  òîì æå ãîäó îí áûë
èçáðàí çàâåäóþùèì êàôåäðîé ïëîäîâîäñòâà è ñóáòðîïè÷åñêèõ êóëüòóð, à ïîñëå
îáúåäèíåíèÿ íåñêîëüêèõ êàôåäð â îäíó — åå ïðîôåññîðîì. Îí òàêæå îñóùåñòâëÿåò íàó÷íîå ðóêîâîäñòâî îòðàñëåâîé èññëåäîâàòåëüñêîé ëàáîðàòîðèåé ïî ñîçäàíèþ ãåíîôîíäà è ðàçðàáîòêå òåõíîëîãèè âîçäåëûâàíèÿ ïëîäîâûõ, îâîùíûõ è
ñóáòðîïè÷åñêèõ ðàñòåíèé. Ñ 1998 ãîäà Çàóð Ìóçàäèë îãëû Ãàñàíîâ — ÷ëåí îäíîãî, çàòåì äâóõ äèññåðòàöèîííûõ ñîâåòîâ â Ðåñïóáëèêå Àçåðáàéäæàí, ïðåäñåäàòåëü íàó÷íîãî ñåìèíàðà îäíîãî èç ñîâåòîâ. Îí — ÷ëåí Ðåñïóáëèêàíñêîé êîìèññèè ïî ïëîäîâîäñòâó è âèíîãðàäàðñòâó ÔÀÎ, ÷ëåí íàó÷íî-òåõíè÷åñêîãî ñîâåòà è
Ýêñïåðòíîé êîìèññèè ïî îõðàíå ãåíåòè÷åñêèõ ðåñóðñîâ ïðè ÍÈÈ Ãåíåòè÷åñêèõ
ðåñóðñîâ Íàöèîíàëüíîé àêàäåìèè íàóê Àçåðáàéäæàíñêîé Ðåñïóáëèêè. Íàó÷íàÿ
äåÿòåëüíîñòü Ç.Ì. Ãàñàíîâà íàïðàâëåíà íà èññëåäîâàíèÿ ðàçëè÷íûõ àñïåêòîâ
ïëîäîâîäñòâà: ôèçèîëîãèè, áèîëîãèè ðîñòà è ðàçâèòèÿ, ýêîëîãèè, àãðîòåõíîëîãèè, ñîðòîèçó÷åíèÿ è äð.
Ç.Ì. Ãàñàíîâ — àâòîð îäíîé íàó÷íîé ìîíîãðàôèè, îêîëî 180 íàó÷íûõ
ñòàòåé, øåñòè ó÷åáíèêîâ, øåñòè ó÷åáíûõ ïîñîáèé, ïÿòè êíèã, îêîëî 15 íàó÷íî-ïóáëèöèñòè÷åñêèõ ñòàòåé è íàó÷íûé ðåäàêòîð øåñòè ó÷åáíèêîâ. Îí ïîäãîòîâèë âîñåìü êàíäèäàòîâ íàóê, 10 ìàãèñòðîâ ïëîäîâîäñòâà è îäíîãî äîêòîðà
íàóê, â íàñòîÿùåå âðåìÿ ðóêîâîäèò ñåìüþ äîêòîðàíòàìè è àñïèðàíòàìè. Ç.Ì.
Ãàñàíîâ — ÷ëåí ðåäàêöèîííîé êîëëåãèè ñáîðíèêà íàó÷íûõ òðóäîâ Âñåðîññèéñêîãî ÍÈÈ öâåòîâîäñòâà è ñóáòðîïè÷åñêèõ êóëüòóð «Ñóáòðîïè÷åñêîå è äåêîðàòèâíîå ñàäîâîäñòâî» (ã. Ñî÷è).
Í.À. Ñëåï÷åíêî
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