Фотосинтез и продуктивность: роль температурного фактора
реклама
ÑÅËÜÑÊÎÕÎÇßÉÑÒÂÅÍÍÀß ÁÈÎËÎÃÈß, 2014, ¹ 5, ñ. 88-95 Ôîòîñèíòåç è ïðîäóêòèâíîñòü: ðîëü òåìïåðàòóðíîãî ôàêòîðà ÓÄÊ 633.1:581.132.1:58.056 doi: 10.15389/agrobiology.2014.5.88rus ÑÂßÇÜ ÌÅÆÄÓ ÂÅËÈ×ÈÍÎÉ ÕËÎÐÎÔÈËËÜÍÎÃÎ ÔÎÒÎÑÈÍÒÅÒÈ×ÅÑÊÎÃÎ ÏÎÒÅÍÖÈÀËÀ È ÓÐÎÆÀÉÍÎÑÒÜÞ ÎÇÈÌÎÉ ÏØÅÍÈÖÛ (Triticum aestivum L.) ÏÐÈ ÏÎÂÛØÅÍÍÛÕ ÒÅÌÏÅÐÀÒÓÐÀÕ Ã.À. ÏÐßÄÊÈÍÀ, Î.Î. ÑÒÀÑÈÊ, Ë.Í. ÌÈÕÀËÜÑÊÀß, Â.Â. ØÂÀÐÒÀÓ Äëÿ ðàçðàáîòêè êðèòåðèåâ ñåëåêöèè íà âûñîêóþ ïðîäóêòèâíîñòü è ìîäåëåé ïðîãíîçèðîâàíèÿ óðîæàéíîñòè â àãðîöåíîçàõ âàæíîå çíà÷åíèå èìååò âçàèìîñâÿçü ìåæäó ïîêàçàòåëÿìè ôîòîñèíòåòè÷åñêîé àêòèâíîñòè è ïðîäóêòèâíîñòüþ êóëüòóð. Ïðè îïòèìàëüíûõ ïîãîäíûõ óñëîâèÿõ íàèáîëåå òåñíî ñ âåëè÷èíîé óðîæàÿ êîððåëèðóåò õëîðîôèëëüíûé ôîòîñèíòåòè÷åñêèé ïîòåíöèàë (ÕëÔÏ), õàðàêòåðèçóþùèé ñóììàðíîå êîëè÷åñòâî õëîðîôèëëà â íàäçåìíîé ÷àñòè ðàñòåíèé (èëè ëèñòüÿõ) â ðàñ÷åòå íà åäèíèöó ïëîùàäè ïîñåâà çà âåãåòàöèþ èëè åå îïðåäåëåííûé ïåðèîä (Ò.Ì. Øàä÷èíà ñ ñîàâò., 2007; E. Kutasy ñ ñîàâò., 2005). Ñîâðåìåííûå ãëîáàëüíûå êëèìàòè÷åñêèå èçìåíåíèÿ ïðåæäå âñåãî ìîãóò ñêàçûâàòüñÿ íà ÷óâñòâèòåëüíûõ ê âûñîêèì òåìïåðàòóðàì êóëüòóðàõ, ê êîòîðûì, â ÷àñòíîñòè, îòíîñèòñÿ îçèìàÿ ìÿãêàÿ ïøåíèöà. Ïðè ýòîì âçàèìîñâÿçü ôîòîñèíòåòè÷åñêèõ ïîêàçàòåëåé ñ óðîæàéíîñòüþ ìîæåò áûòü íàðóøåíà.  óñëîâèÿõ ïîëåâîãî îïûòà ìû èçó÷èëè âëèÿíèå ïîâûøåííûõ òåìïåðàòóð âîçäóõà â ïåðèîä âåñåííå-ëåòíåé âåãåòàöèè íà ïîêàçàòåëè ôîòîñèíòåòè÷åñêîãî àïïàðàòà ó êîðîòêîñòåáåëüíûõ âûñîêîóðîæàéíûõ ñîðòîâ ìÿãêîé îçèìîé ïøåíèöû Ñìóãëÿíêà (ñðåäíåðàííèé) è Ïåðåÿñëàâêà (ñðåäíåñïåëûé) íà ðàçíûõ ôîíàõ ìèíåðàëüíîãî ïèòàíèÿ (áåç ïðèìåíåíèÿ óäîáðåíèé è ñ èõ âíåñåíèåì â äîçàõ N90P60K60S10 è N120P90K90S20 îñåíüþ è â ðàçíûå ôàçû âåñåííå-ëåòíåé âåãåòàöèè). Èññëåäîâàíèÿ ïðîâîäèëè â 2009 è 2011 ãîäàõ, êîòîðûå õàðàêòåðèçîâàëèñü ïîâûøåííûìè òåìïåðàòóðàìè â ïåðèîä âåãåòàöèè, â îïûòíîì ñåëüñêîõîçÿéñòâåííîì ïðîèçâîäñòâå Èíñòèòóòà ôèçèîëîãèè ðàñòåíèé è ãåíåòèêè ÍÀÍ Óêðàèíû (Êèåâñêàÿ îáë.). Ïîêàçàíî, ÷òî òàêèå ïîãîäíûå óñëîâèÿ ïðèâåëè ê ñíèæåíèþ ñîäåðæàíèÿ õëîðîôèëëà â ëèñòüÿõ, ðàçìåðîâ àññèìèëÿöèîííîé ïîâåðõíîñòè ïîñåâîâ è ïðîäîëæèòåëüíîñòè åå ôóíêöèîíèðîâàíèÿ, à òàêæå óðîæàéíîñòè. Âî âñåõ âàðèàíòàõ (è íà âûñîêîì ôîíå ìèíåðàëüíîãî ïèòàíèÿ, è áåç âíåñåíèÿ óäîáðåíèé) ïëîùàäü ëèñòîâîé ïîâåðõíîñòè â 2009 ãîäó áûëà âûøå, ÷åì â ñîîòâåòñòâóþùèõ âàðèàíòàõ â 2011 ãîäó âî âñå òðè èññëåäóåìûå ôàçû âåãåòàöèè. Âíåñåíèå ìèíåðàëüíûõ óäîáðåíèé óâåëè÷èâàëî ëèñòîâîé èíäåêñ â îáà ãîäà â ñðåäíåì â 1,5-2,0 ðàçà (ó ñîðòà Ïåðåÿñëàâêà â ôàçó ìîëî÷íî-âîñêîâîé ñïåëîñòè — ïî÷òè â 5 ðàç ïî ñðàâíåíèþ ñ êîíòðîëåì). Ñîäåðæàíèå õëîðîôèëëà ðàçëè÷àëîñü â çàâèñèìîñòè îò ãîäà: â ôàçó öâåòåíèÿ â âàðèàíòàõ ñ âíåñåíèåì ìèíåðàëüíûõ óäîáðåíèé îíî ñîñòàâèëî 5,5-6,0 è 3,5-4,5 ìã/äì2 ñîîòâåòñòâåííî â 2009 è â 2011 ãîäó, â êîíòðîëå — ñîîòâåòñòâåííî 3,2-4,3 è 2,8 ìã/äì2, è òàêîå ñîîòíîøåíèå ñîõðàíÿëîñü â ïîñëåäóþùèå ôàçû ðàçâèòèÿ.  áîëüøèíñòâå ñëó÷àåâ ìàêñèìàëüíûå çíà÷åíèÿ ñîäåðæàíèÿ ïèãìåíòîâ â ñðåäíåé ïðîáå çåëåíûõ ëèñòüåâ ïîáåãà îòìå÷àëèñü â ôàçó ìîëî÷íîé ñïåëîñòè, î÷åâèäíî, âñëåäñòâèå ïîëíîãî îòìèðàíèÿ ïëàñòèíêè ëèñòüåâ íèæíèõ ÿðóñîâ. Âíåñåíèå ìèíåðàëüíûõ óäîáðåíèé óâåëè÷èâàëî êîëè÷åñòâî õëîðîôèëëà â ëèñòüÿõ ó îáîèõ ñîðòîâ. Ïî ðåçóëüòàòàì äèñïåðñèîííîãî àíàëèçà (F-êðèòåðèé Ôèøåðà), íàèáîëüøåå âëèÿíèå íà èçìåíåíèÿ ïîêàçàòåëåé ìîùíîñòè ðàçâèòèÿ ôîòîñèíòåòè÷åñêîãî àïïàðàòà â íàøèõ ýêñïåðèìåíòàõ îêàçûâàëè óñëîâèÿ ãîäà, âòîðûì ïî çíà÷èìîñòè áûë óðîâåíü ìèíåðàëüíîãî ïèòàíèÿ. Âåëè÷èíà õëîðîôèëëüíîãî ôîòîñèíòåòè÷åñêîãî ïîòåíöèàëà âåñüìà ñóùåñòâåííî çàâèñåëà îò êîìáèíàöèè ôàêòîðîâ ñîðò ½ óñëîâèÿ ãîäà è ìåíüøå — îò âçàèìîäåéñòâèé óñëîâèÿ ãîäà ½ ôîí ìèíåðàëüíîãî ïèòàíèÿ è ñîðò ½ ôîí ìèíåðàëüíîãî ïèòàíèÿ. Âûñîêèå è áëèçêèå ïî âåëè÷èíå çíà÷åíèÿ êîýôôèöèåíòà êîððåëÿöèè (îò 0,93 äî 0,99) ìåæäó çåðíîâîé ïðîäóêòèâíîñòüþ ïîñåâà è õëîðîôèëëüíûì ôîòîñèíòåòè÷åñêèì ïîòåíöèàëîì îòìå÷àëèñü äëÿ âûáîðîê ïî êàæäîìó ãîäó, ñîðòó èëè âàðèàíòó ìèíåðàëüíîãî ïèòàíèÿ, ñèëüíóþ ïîëîæèòåëüíóþ çàâèñèìîñòü âûÿâèëè è äëÿ îáúåäèíåííîé âûáîðêè çà îáà ãîäà. Òàêèì îáðàçîì, óñòàíîâëåíî, ÷òî òåñíàÿ êîððåëÿöèÿ ìåæäó âåëè÷èíîé õëîðîôèëëüíîãî ôîòîñèíòåòè÷åñêîãî ïîòåíöèàëà ëèñòüåâ â ïîñåâå è óðîæàéíîñòüþ îçèìîé ïøåíèöû ñîõðàíÿåòñÿ è â óñëîâèÿõ ïîâûøåííûõ òåìïåðàòóð, à ñàìà çàâèñèìîñòü ìîæåò áûòü îïèñàíà óðàâíåíèåì ðåãðåññèè, îäèíàêîâûì äëÿ ëþáîãî ôàêòîðà, âûçûâàþùåãî âàðüèðîâàíèå ïîêàçàòåëåé. Êëþ÷åâûå ñëîâà: Triticum aestivum L., çåðíîâàÿ ïðîäóêòèâíîñòü, ëèñòîâîé èíäåêñ, õëîðîôèëë, õëîðîôèëëüíûé ôîòîñèíòåòè÷åñêèé ïîòåíöèàë ëèñòüåâ. Èçó÷åíèå âçàèìîñâÿçè ìåæäó ïîêàçàòåëÿìè ôîòîñèíòåòè÷åñêîãî àïïàðàòà ñåëüñêîõîçÿéñòâåííûõ êóëüòóð è óðîæàéíîñòüþ èìååò âàæíîå çíà÷åíèå êàê äëÿ ïîíèìàíèÿ çàêîíîìåðíîñòåé ïðîäóêöèîííîãî ïðîöåññà, òàê è äëÿ ðàçðàáîòêè êðèòåðèåâ ñåëåêöèè íà âûñîêóþ ïðîäóêòèâíîñòü, à 88 òàêæå ìîäåëåé ïðîãíîçèðîâàíèÿ óðîæàéíîñòè â àãðîöåíîçàõ. Ïî äàííûì ìíîãèõ àâòîðîâ, íàèáîëåå òåñíî ñ óðîæàéíîñòüþ êîððåëèðóåò õëîðîôèëëüíûé ôîòîñèíòåòè÷åñêèé ïîòåíöèàë (ÕëÔÏ), õàðàêòåðèçóþùèé ñóììàðíîå êîëè÷åñòâî õëîðîôèëëà â íàäçåìíîé ÷àñòè ðàñòåíèé (èëè ëèñòüÿõ) íà åäèíèöó ïëîùàäè ïîñåâà çà âåãåòàöèþ èëè åå îïðåäåëåííûé ïåðèîä (1-3). Ó ðàçíûõ ãåíîòèïîâ îçèìîé ïøåíèöû íàìè áûëà ïîêàçàíà òåñíàÿ ñâÿçü ìåæäó ÕëÔÏ ëèñòüåâ è ïðîäóêòèâíîñòüþ ïîñåâîâ è âîçìîæíîñòü èñïîëüçîâàòü òàêóþ çàâèñèìîñòü äëÿ ïðîãíîçèðîâàíèÿ óðîæàéíîñòè (4). Âûñîêàÿ ñòåïåíü êîððåëÿöèè ÕëÔÏ ñ óðîæàåì îáóñëîâëåíà òåì, ÷òî ýòîò ïîêàçàòåëü õîðîøî îòðàæàåò äèíàìèêó ôîðìèðîâàíèÿ áèîìàññû è ýôôåêòèâíîñòü ïîãëîùåíèÿ ôîòîñèíòåòè÷åñêè àêòèâíîé ðàäèàöèè (ÔÀÐ) ïîñåâîì â ïðîöåññå âåãåòàöèè (5-7). Âûñîêèå êîýôôèöèåíòû êîððåëÿöèè ìåæäó ÕëÔÏ çëàêîâûõ ðàñòåíèé è èõ ïðîäóêòèâíîñòüþ, êàê ïðàâèëî, îòìå÷àþò â áëàãîïðèÿòíûå ïî ïîãîäíûì óñëîâèÿì ãîäû (4, 8, 9). Ñ÷èòàåòñÿ, ÷òî ôîðìèðîâàíèå ïîñåâîâ ñ âûñîêèìè ëèñòîâûìè è õëîðîôèëëüíûìè èíäåêñàìè, îñîáåííî ïðè âíåñåíèè âûñîêèõ äîç àçîòíûõ óäîáðåíèé, ìîæåò îêàçàòüñÿ êîíòðïðîäóêòèâíûì â óñëîâèÿõ äåôèöèòà âëàãîîáåñïå÷åííîñòè è ïîâûøåííûõ òåìïåðàòóð âñëåäñòâèå óâåëè÷åíèÿ ïîòåðü ïî÷âåííîé âëàãè è çàòðàò àññèìèëèðîâàííîãî óãëåðîäà íà äûõàíèå (10). Ïðè ýòîì èç-çà ãëîáàëüíûõ èçìåíåíèé êëèìàòà ÷àñòîòà è ïðîäîëæèòåëüíîñòü ïåðèîäîâ ñ ïîâûøåííûìè òåìïåðàòóðàìè ñóùåñòâåííî óâåëè÷èëèñü (11, 12). Òàê, â Óêðàèíå ñ 2007 ïî 2012 ãîäû ðåêîðäíûå çíà÷åíèÿ ñðåäíåñóòî÷íîé òåìïåðàòóðû âîçäóõà â ïåðèîä âåñåííå-ëåòíåé âåãåòàöèè íå áûëè çàðåãèñòðèðîâàíû òîëüêî â 2008 ãîäó. Îçèìàÿ ìÿãêàÿ ïøåíèöà (Triticum aestivum L.) — ÷óâñòâèòåëüíàÿ ê âûñîêîé òåìïåðàòóðå êóëüòóðà, è îñîáåííî çíà÷èòåëüíûå ïîòåðè óðîæàéíîñòè âûçûâàþòñÿ ïîâûøåííûìè òåìïåðàòóðàìè âîçäóõà â ðåïðîäóêòèâíûé ïåðèîä ðàçâèòèÿ (13-15). Âûñîêîòåìïåðàòóðíûé ñòðåññ îêàçûâàåò èíãèáèðóþùåå äåéñòâèå íà ñèíòåç õëîðîôèëëà è àêòèâíîñòü ôîòîñèíòåòè÷åñêîãî àïïàðàòà, óñêîðÿåò ñòàðåíèå è óìåíüøàåò ïðîäîëæèòåëüíîñòü æèçíè ëèñòüåâ, óãíåòàåò è íàðóøàåò ïðîöåññû ôîðìèðîâàíèÿ ýëåìåíòîâ êîëîñà è ôåðòèëüíîñòü ïûëüöû, ïîäàâëÿåò îáðàçîâàíèå è íàëèâ çåðíîâêè, ÷òî â êîíå÷íîì èòîãå âûçûâàåò ñíèæåíèå ÷èñëà è ìàññû çåðåí â êîëîñå (16, 17). Õîòÿ îñòàåòñÿ íåÿñíûì, ÿâëÿþòñÿ ëè èçìåíåíèÿ â ðåïðîäóêòèâíîé ñôåðå ñàìîñòîÿòåëüíûìè ôàêòîðàìè ñíèæåíèÿ óðîæàéíîñòè èëè îíè ñâÿçàíû ñ íåäîñòàòî÷íûì îáåñïå÷åíèåì ôîòîàññìèëÿòàìè (14, 18), î÷åâèäíî, ÷òî ïîâûøåííûå òåìïåðàòóðû â ïåðèîä âåãåòàöèè ìîãóò ñóùåñòâåííî íàðóøàòü âçàèìîñâÿçü ôîòîñèíòåòè÷åñêèõ ïîêàçàòåëåé è çåðíîâîé ïðîäóêòèâíîñòè. Ïîýòîìó â ãîäû âûñîêîòåìïåðàòóðíûõ ýêñòðåìóìîâ èñïîëüçîâàíèå çàâèñèìîñòåé ìåæäó ïîêàçàòåëÿìè ðàçâèòèÿ ôîòîñèíòåòè÷åñêîãî àïïàðàòà è ïðîäóêòèâíîñòüþ äëÿ öåëåé ñåëåêöèè è ïðîãíîçèðîâàíèÿ óðîæàéíîñòè ìîæåò îêàçàòüñÿ ïðîáëåìàòè÷íûì. Öåëüþ íàøåé ðàáîòû áûëî èññëåäîâàíèå çàâèñèìîñòè ìåæäó ïîêàçàòåëÿìè ðàçâèòèÿ ôîòîñèíòåòè÷åñêîãî àïïàðàòà è çåðíîâîé ïðîäóêòèâíîñòüþ â öåíîçàõ îçèìîé ïøåíèöû íà ðàçíûõ ôîíàõ ìèíåðàëüíîãî ïèòàíèÿ â ãîäû, êîãäà îòìå÷àëèñü ïîâûøåííûå òåìïåðàòóðû âîçäóõà â òå÷åíèå âåãåòàöèè. Ìåòîäèêà. Èññëåäîâàíèÿ ïðîâîäèëè â 2009 è 2011 ãîäàõ, êîòîðûå õàðàêòåðèçîâàëèñü ïîâûøåííûìè òåìïåðàòóðàìè â âåñåííå-ëåòíèé ïåðèîä, íà äâóõ êîðîòêîñòåáåëüíûõ âûñîêîóðîæàéíûõ ñîðòàõ ìÿãêîé îçèìîé ïøåíèöû, âûðàùåííûõ íà ó÷àñòêàõ ñîðòîèñïûòàíèÿ îçèìîé ïøåíèöû â îïûòíîì ñåëüñêîõîçÿéñòâåííîì ïðîèçâîäñòâå Èíñòèòóòà ôèçèîëîãèè ðàñòåíèé è ãåíåòèêè ÍÀÍ Óêðàèíû (Êèåâñêàÿ îáë.). Ñîðò Ñìóãëÿíêà ñðåäíåðàííèé (âåãåòàöèîííûé ïåðèîä — 278-281 ñóò, óðîæàéíîñòü — îò 60 äî 89 115 ö/ãà), ñîðò Ïåðåÿñëàâêà — ñðåäíåñïåëûé (âåãåòàöèîííûé ïåðèîä — 280-287 ñóò, óðîæàéíîñòü — 60-103 ö/ãà) (19). Íîðìà âûñåâà ñåìÿí ñîñòàâëÿëà 5,0-5,5 ìëí/ãà. ×èñëî ïðîäóêòèâíûõ ïîáåãîâ ïðè óáîðêå óðîæàÿ êîëåáàëîñü îò 500 äî 750 øò./ì2 ïî÷âû. Ïëîùàäü ó÷àñòêà â êàæäîé èç ÷åòûðåõ ïîâòîðíîñòåé ðàâíÿëàñü 40 ì2, áèîëîãè÷åñêàÿ ïîâòîðíîñòü — 20 ðàñòåíèé. Îïûòû ïðîâîäèëè íà äåðíîâî-ñëàáî- è ñðåäíåïîäçîëèñòûõ îãëååííûõ ñóïåñ÷àíûõ ïî÷âàõ. Ñîäåðæàíèå ãóìóñà — 1,8 %, ðÍ (ÊCl) — 6,2. Ìèíåðàëüíûå óäîáðåíèÿ (N90P60K60S10 è N120P90K90S20) âíîñèëè îñåíüþ è â îòäåëüíûå ôàçû âåñåííå-ëåòíåé âåãåòàöèè. Êîíòðîëåì ñëóæèë âàðèàíò áåç ïðèìåíåíèÿ óäîáðåíèé. Îòáîð îáðàçöîâ äëÿ îïðåäåëåíèÿ ñîäåðæàíèÿ ôîòîñèíòåòè÷åñêèõ ïèãìåíòîâ è ïëîùàäè àññèìèëÿöèîííîé ïîâåðõíîñòè ïîñåâîâ îñóùåñòâëÿëè â òå÷åíèå ïåðèîäà öâåòåíèå—ìîëî÷íî-âîñêîâàÿ ñïåëîñòü çåðíà. Ñîäåðæàíèå ñóììû õëîðîôèëëîâ a è b â ñðåäíåé ïðîáå âñåõ ëèñòüåâ ïîáåãà îïðåäåëÿëè ñïåêòðîôîòîìåòðè÷åñêèì ìåòîäîì ñ ýêñòðàêöèåé â äèìåòèëñóëüôîêñèäå (ÄÌÑÎ) (20). Ïëîùàäü çåëåíûõ ëèñòüåâ îòäåëüíîãî ïîáåãà îçèìîé ïøåíèöû ðàññ÷èòûâàëè êàê ïðîèçâåäåíèå çíà÷åíèé èõ äëèíû è ìàêñèìàëüíîé øèðèíû ñ ïîïðàâî÷íûì êîýôôèöèåíòîì 0,76, ëèñòîâîé èíäåêñ (ËÈ) — óìíîæàÿ ïîëó÷åííóþ âåëè÷èíó íà ãóñòîòó ñòîÿíèÿ ïîáåãîâ. Õëîðîôèëëüíûé èíäåêñ (ÕëÈ) âû÷èñëÿëè êàê ïðîèçâåäåíèå ËÈ è ñîäåðæàíèÿ õëîðîôèëëà â ëèñòüÿõ. Õëîðîôèëëüíûé ôîòîñèíòåòè÷åñêèé ïîòåíöèàë îïðåäåëÿëè êàê ñóììó åæåñóòî÷íûõ çíà÷åíèé õëîðîôèëëüíîãî èíäåêñà çà îïðåäåëåííûé ïåðèîä ðàçâèòèÿ (21). Äëÿ åãî ðàñ÷åòà áûëè ïîñòðîåíû ãðàôèêè èçìåíåíèÿ õëîðîôèëëüíûõ èíäåêñîâ äëÿ çåëåíûõ ëèñòüåâ â òå÷åíèå ïåðèîäà öâåòåíèå—ìîëî÷íî-âîñêîâàÿ ñïåëîñòü. Ðåçóëüòàòû ýêñïåðèìåíòîâ îáðàáàòûâàëè ñòàòèñòè÷åñêè ñ ïîìîùüþ ïðîãðàììû Microsoft Excel. Ðåçóëüòàòû.  îáà ãîäà èññëåäîâàíèé ñðåäíåñóòî÷íàÿ òåìïåðàòóðà âîçäóõà â ïåðèîä ðåïðîäóêòèâíîãî ðàçâèòèÿ îçèìîé ïøåíèöû áûëà ñóùåñòâåííî âûøå ñðåäíèõ ìíîãîëåòíèõ çíà÷åíèé.  2009 ãîäó ïðåâûøåíèå ñðåäíåñóòî÷íûõ òåìïåðàòóð âîçäóõà íàä êëèìàòè÷åñêèìè ôèêñèðîâàëè â òå÷åíèå 2 íåä ñíà÷àëà íåáîëüøîå (íà 0,8 °Ñ) â ïåðèîä îò êîëîøåíèÿ äî öâåòåíèÿ, à ïîòîì áîëåå ñóùåñòâåííîå (íà 1,5 °Ñ) — îò ôàçû ìîëî÷íîé ñïåëîñòè (ÌÑ) è äî ìîëî÷íî-âîñêîâîé ñïåëîñòè (ÌÂÑ).  ñðåäíåì çà ïåðèîä öâåòåíèå—ÌÂÑ ðàçíèöà ðàâíÿëàñü 1,2 °Ñ.  2011 ãîäó ñðåäíÿÿ òåìïåðàòóðà áûëà çíà÷èòåëüíî (íà 5,1 °Ñ) âûøå íîðìû â òå÷åíèå âñåãî ïåðèîäà îò öâåòåíèÿ è äî ÌÂÑ.  îòäåëüíûå äíè ñðåäíåñóòî÷íàÿ òåìïåðàòóðà ïðåâîñõîäèëà íîðìó áîëåå ÷åì íà 8 °Ñ. Êðîìå òîãî, â ýòîì ãîäó ïðåâûøåíèå ñðåäíåñóòî÷íûõ òåìïåðàòóð íà÷àëîñü çà 1,5 íåä äî íà÷àëà ôàçû öâåòåíèÿ è ñîâïàëî ñ îòñóòñòâèåì îñàäêîâ. Ïîêàçàòåëü óâëàæíåííîñòè òåïëîãî ïåðèîäà ãîäà (ãèäðîòåðìè÷åñêèé êîýôôèöèåíò) â ìàå 2009 ãîäà ñîñòàâëÿë 0,8, 2011 ãîäà — 0,6, òîãäà êàê åãî ñðåäíåå ìíîãîëåòíåå çíà÷åíèå äëÿ Êèåâñêîé îáëàñòè â ýòîò ìåñÿö — 1,2. Òàêîé õàðàêòåð ïîãîäíûõ óñëîâèé ñóùåñòâåííî ïîâëèÿë íà ðàçâèòèå ðàñòåíèÿ, îñîáåííî íà ñòàäèè âåãåòàòèâíîãî ðîñòà. Òàê, â 2011 ãîäó ôàçà öâåòåíèÿ íàñòóïèëà íà 12 ñóò ðàíüøå, ÷åì â 2009 ãîäó. Äëèòåëüíîñòü ïåðèîäà öâåòåíèå—ÌÂÑ ðàçëè÷àëàñü ìåíüøå (â 2009 ãîäó îíà áûëà êîðî÷å ïðèìåðíî íà 4 ñóò). Ïîãîäíûå óñëîâèÿ ãîäà îêàçàëè ñóùåñòâåííîå âëèÿíèå íà ôîðìèðîâàíèå àññèìèëÿöèîííîãî àïïàðàòà ïîñåâîâ è íà èçìåíåíèå âåëè÷èíû ËÈ â òå÷åíèå âåãåòàöèè ïøåíèöû (ðèñ. 1). Âî âñåõ âàðèàíòàõ (è íà âûñîêîì ôîíå ìèíåðàëüíîãî ïèòàíèÿ, è áåç âíåñåíèÿ óäîáðåíèé) ïëîùàäü ëèñòîâîé ïîâåðõíîñòè â 2009 ãîäó áûëà âûøå, ÷åì â ñîîòâåòñòâóþùèõ âàðè90 àíòàõ â 2011 ãîäó âî âñå òðè èññëåäóåìûå ôàçû âåãåòàöèè. Âíåñåíèå ìèíåðàëüíûõ óäîáðåíèé òàêæå çíà÷èòåëüíî (â ñðåäíåì â 1,5-2,0 ðàçà) óâåëè÷èâàëî ËÈ â îáà ãîäà. Äëÿ ñîðòà Ïåðåÿñëàâêà âëèÿíèå ìèíåðàëüíûõ óäîáðåíèé íà ïëîùàäü ëèñòîâîé ïîâåðõíîñòè â ôàçó ÌÂÑ áûëî íàèáîëåå âûñîêèì — ïî÷òè â 5 ðàç ïî ñðàâíåíèþ ñ êîíòðîëåì. Ðèñ. 1. Äèíàìèêà ëèñòîâîãî èíäåêñà (ËÈ) ïîñåâîâ (à, á) è ñîäåðæàíèÿ õëîðîôèëëà (â, ã) â ãîäû ñ ïîâûøåííîé òåìïåðàòóðîé âîçäóõà â ïåðèîä âåãåòàöèè (à, â — 2009 ãîä, á, ã — 2011 ãîä) ó îçèìîé ïøåíèöû (ñïëîøíûå ëèíèè — ñîðò Ñìóãëÿíêà, ïóíêòèðíûå ëèíèè — ñîðò Ïåðåÿñëàâêà) çà âðåìÿ îò öâåòåíèÿ (I) äî ìîëî÷íî-âîñêîâîé ñïåëîñòè (III) íà ðàçíûõ ôîíàõ ìèíåðàëüíîãî ïèòàíèÿ: è — êîíòðîëü; è — N90P60K60S10; S è U — N120P90K90S20 (îïûòíîå ñåëüñêîõîçÿéñòâåííîå ïðîèçâîäñòâî Èíñòèòóòà ôèçèîëîãèè ðàñòåíèé è ãåíåòèêè ÍÀÍ Óêðàèíû, Êèåâñêàÿ îáë.). Cîäåðæàíèå õëîðîôèëëà â ëèñòüÿõ ó îáîèõ ñîðòîâ îçèìîé ïøåíèöû òàêæå ñóùåñòâåííî ðàçëè÷àëîñü â çàâèñèìîñòè îò ãîäà: â 2011 ãîäó çíà÷åíèÿ áûëè ìåíüøå ïî ñðàâíåíèþ ñ òàêîâûìè â ñîîòâåòñòâóþùèõ âàðèàíòàõ â 2009 ãîäó. Òàê, â ôàçó öâåòåíèÿ ýòîò ïîêàçàòåëü â 2009 è 2011 ãîäàõ ïðè âíåñåíèè óäîáðåíèé ñîñòàâëÿë ñîîòâåòñòâåííî 5,5-6,0 è 3,5-4,5 ìã/äì2, â êîíòðîëå — 3,2-4,3 è 2,8 ìã/äì2 (ñì. ðèñ. 1). Ïîõîæåå ñîîòíîøåíèå â öåëîì ñîõðàíÿëîñü è íà ïðîòÿæåíèè äâóõ ïîñëåäóþùèõ ôàç ðàçâèòèÿ. Âìåñòå ñ òåì ñëåäóåò îòìåòèòü áîëåå ðåçêîå ñíèæåíèå ñîäåðæàíèÿ õëîðîôèëëà â ëèñòüÿõ â 2011 ãîäó, ñèëüíåå âûðàæåííîå ó ñîðòà Ñìóãëÿíêà, îñîáåííî â óñëîâèÿõ äåôèöèòà ìèíåðàëüíîãî ïèòàíèÿ. Ïî-âèäèìîìó, ýòî áûëî ñâÿçàíî ñ óñêîðåíèåì ïðîöåññîâ ñòàðåíèÿ è ðåóòèëèçàöèè àçîòà èç ëèñòüåâ ó ñîðòà Ñìóãëÿíêà, èíäóöèðîâàííûì ïîâûøåííûìè òåìïåðàòóðàìè. Ó ñîðòà Ïåðåÿñëàâêà êîëè÷åñòâî õëîðîôèëëà ìåíåå çàâèñåëî îò ïîãîäíûõ óñëîâèé âåãåòàöèîííîãî ïåðèîäà, è â áîëåå æàðêîì 2011 ãîäó â öåëîì ïî âàðèàíòàì îíî ïðåâûøàëî ñîîòâåòñòâóþùèå ïîêàçàòåëè ó ñîðòà Ñìóãëÿíêà. Âíåñåíèå ìèíåðàëüíûõ óäîáðåíèé óâåëè÷èâàëî êîëè÷åñòâî õëîðîôèëëà â ëèñòüÿõ ó îáîèõ ñîðòîâ.  2009 ãîäó ñîäåðæàíèå çåëåíûõ ïèãìåíòîâ âîçðîñëî â 1,3-1,5 ðàçà ïî ñðàâíåíèþ ñ êîíòðîëåì, à â 2011 ãîäû ïîâûñèëîñü åùå çíà÷èòåëüíåå — â 1,7-2,2 ðàçà. Óâåëè÷åíèå äîç âíîñèìûõ óäîáðåíèé âûçûâàëî äîñòîâåðíîå âîçðàñòàíèå ñîäåðæàíèÿ ôîòîñèíòåòè÷å91 ñêèõ ïèãìåíòîâ â ëèñòüÿõ ó îáîèõ ñîðòîâ. Õàðàêòåðíî, ÷òî â áîëüøèíñòâå ñëó÷àåâ ìàêñèìàëüíîå ñîäåðæàíèå õëîðîôèëëà â ñðåäíåé ïðîáå ëèñòüåâ ïîáåãà îòìå÷àëè â ôàçó ìîëî÷íîé ñïåëîñòè (î÷åâèäíî, âñëåäñòâèå ïîëíîãî îòìèðàíèÿ ïëàñòèíêè ëèñòüåâ íèæíèõ ÿðóñîâ). Ïðè ýòîì îíòîãåíåòè÷åñêèå èçìåíåíèÿ èçó÷àåìîãî ïîêàçàòåëÿ ó ñîðòà Ïåðåÿñëàâêà áûëè ñõîäíûìè äëÿ âñåõ âàðèàíòîâ â îáà ãîäà, îáåñïå÷èâàÿ ïðèìåðíî îäèíàêîâûå îòíîñèòåëüíûå ðàçëè÷èÿ ìåæäó çíà÷åíèÿìè íà ïîçäíèõ ýòàïàõ îíòîãåíåçà êàê ïðè ïðèìåíåíèè, òàê è áåç èñïîëüçîÐèñ. 2. Õëîðîôèëëüíûé ôîòîñèíòåòè÷åñêèé ïîòåíöèàë âàíèÿ óäîáðåíèé. Ó ñîðòà (ÕëÔÏ) îçèìîé ïøåíèöû (1-3 — ñîðò Ñìóãëÿíêà, Ñìóãëÿíêà íàèáîëüøåå âëè4-6 — ñîðò Ïåðåÿñëàâêà) â ãîäû ñ ïîâûøåííîé òåìïåðàòóðîé âîçäóõà â ïåðèîä âåãåòàöèè çà âðåìÿ îò öâåòåíèÿ ÿíèå óäîáðåíèé íà ñèíòåç äî ìîëî÷íî-âîñêîâîé ñïåëîñòè íà ðàçíûõ ôîíàõ ìèíå- õëîðîôèëëà ðåãèñòðèðîâàëè â ðàëüíîãî ïèòàíèÿ: 1 è 4 — êîíòðîëü, 2 è 5 — ïåðèîä ÌÑ (îñîáåííî â 2011 N90P60K60S10, 3 è 6 — N120P90K90S20 (îïûòíîå ñåëüñêî- ãîäó), à â ôàçó ÌÂÑ ðàçëè÷èÿ õîçÿéñòâåííîå ïðîèçâîäñòâî Èíñòèòóòà ôèçèîëîãèè ìåæäó âàðèàíòàìè óìåíüøàðàñòåíèé è ãåíåòèêè ÍÀÍ Óêðàèíû, Êèåâñêàÿ îáë.). ëèñü âñëåäñòâèå áûñòðîãî ñíèæåíèÿ êîëè÷åñòâà ïèãìåíòà ïðè ïðèìåíåíèè óäîáðåíèé. Çíà÷åíèÿ õëîðîôèëëüíîãî ôîòîñèíòåòè÷åñêîãî ïîòåíöèàëà, ñôîðìèðîâàííîãî â ïîñåâàõ, êàê èíòåãðàëüíîãî ïîêàçàòåëÿ, îòðàæàþùåãî ïëîùàäü àññèìèëÿöèîííîé ïîâåðõíîñòè ðàñòåíèé, à òàêæå ñîäåðæàíèå õëîðîôèëëà è èçìåíåíèÿ ýòèõ ïîêàçàòåëåé â îíòîãåíåçå (ðèñ. 2), â 2011 ãîäó áûëè ñóùåñòâåííî íèæå, ÷åì â 2009 ãîäó. Âìåñòå ñ òåì âëèÿíèå ïîãîäíûõ óñëîâèé ðàçëè÷àëîñü â çàâèñèìîñòè îò ñîðòà è ôîíà ìèíåðàëüíîãî ïèòàíèÿ. Òàê, â 2009 ãîäó ïî ñðàâíåíèþ ñ 2011 ãîäîì ó ñîðòà Ñìóãëÿíêà âåëè÷èíà ÕëÔÏ â êîíòðîëå áûëà âûøå â 4,8 ðàçà, à â âàðèàíòàõ ñ N90P60K60S10 è N120P90K90S20 — ñîîòâåòñòâåííî òîëüêî â 3,0 è 2,0 ðàçà. Ó ñîðòà Ïåðåÿñëàâêà ðàçëè÷èÿ áûëè çíà÷èòåëüíî ìåíüøå — â 1,8 ðàçà (êîíòðîëü) è â 1,5 ðàçà (âàðèàíò ñ N120P90K90S20).  ñâîþ î÷åðåäü, âëèÿíèå óäîáðåíèé íà âåëè÷èíó ÕëÔÏ òàêæå çàâèñåëî îò ñîðòà è ãîäà ïðîâåäåíèÿ îïûòà.  2009 ãîäó îíî áûëî ìåíüøèì è ñõîäíûì (ðàçëè÷èÿ ñîîòâåòñòâåííî â 1,7 è 2,2 ðàçà ó ñîðòîâ Ñìóãëÿíêà è Ïåðåÿñëàâêà).  2011 ãîäó âíåñåíèå óäîáðåíèé â äîçå N120P90K90S20 â 4,0 ðàçà óâåëè÷èâàëî ÕëÔÏ ó ñîðòà Ñìóãëÿíêà è â 2,5 ðàçà ó ñîðòà Ïåðåÿñëàâêà. Êàê ïîêàçàëè ðåçóëüòàòû äèñïåðñèîííîãî àíàëèçà, íàèáîëüøåå âëèÿíèå íà ÕëÈ è ÕëÔÏ â íàøèõ ýêñïåðèìåíòàõ îêàçûâàëè óñëîâèÿ ãîäà (òàáë. 1). Ôîí ìèíåðàëüíîãî ïèòàíèÿ áûë âòîðûì ïî çíà÷èìîñòè ôàêòîðîì âàðüèðîâàíèÿ ýòèõ ïîêàçàòåëåé, è íàèìåíüøåå âëèÿíèå îêàçûâàë ñîðò.  òî æå âðåìÿ âåëè÷èíà ÕëÔÏ âåñüìà ñóùåñòâåííî çàâèñåëà îò êîìáèíàöèè ôàêòîðîâ ñîðò ½ óñëîâèÿ ãîäà è ìåíüøå — îò âçàèìîäåéñòâèÿ ôàêòîðîâ óñëîâèÿ ãîäà ½ ôîí ìèíåðàëüíîãî ïèòàíèÿ è ñîðò ½ ôîí ìèíåðàëüíîãî ïèòàíèÿ. Ó ðàñòåíèé èçó÷àåìûõ ñîðòîâ âìåñòå ñ ïîêàçàòåëÿìè ðàçâèòèÿ ôîòîñèíòåòè÷åñêîãî àïïàðàòà èçìåíÿëàñü çåðíîâàÿ ïðîäóêòèâíîñòü. Ïðè íå92 áëàãîïðèÿòíûõ óñëîâèÿõ 2011 ãîäà ïî ñðàâíåíèþ ñ 2009 ãîäîì ïðîèñõîäèëî ðåçêîå ñíèæåíèå óðîæàéíîñòè (òàáë. 2). Óðîæàé çåðíà ó ñîðòà Ïåðåÿñëàâêà â çàâèñèìîñòè îò ôîíà óäîáðåíèé óìåíüøèëñÿ â 1,8-2,2 ðàçà, ó ñîðòà Ñìóãëÿíêà — â 2,0-2,5 ðàçà. 1. Îöåíêà âëèÿíèÿ ôàêòîðîâ è èõ êîìáèíàöèé íà âàðüèðîâàíèå ïîêàçàòåëåé ðàçâèòèÿ ôîòîñèíòåòè÷åñêîãî àïïàðàòà ó ñîðòîâ îçèìîé ïøåíèöû Ñìóãëÿíêà è Ïåðåÿñëàâêà (F-êðèòåðèé Ôèøåðà; îïûòíîå ñåëüñêîõîçÿéñòâåííîå ïðîèçâîäñòâî Èíñòèòóòà ôèçèîëîãèè ðàñòåíèé è ãåíåòèêè ÍÀÍ Óêðàèíû, Êèåâñêàÿ îáë.) Ôàêòîð, ñî÷åòàíèå ôàêòîðîâ Fôàêò. õëîðîôèëëüíûé èíäåêñ ïî ôàçàì öâåòåíèå MC MBC ÕëÔÏ çà ïåðèîä Ãîä (a) 374 633 431 3913 Ñîðò (b) 2,30 1,08 6,84 54,30 Ôîí ìèíåðàëüíîãî ïèòàíèÿ (c) 123 103 30,1 1516 ab 0,40 28,40 7,30 264 ac 28,00 8,53 3,73 46,80 bc 2,57 2,47 3,61 33,10 abc 1,28 0,13 0,42 2,57 Ï ð è ì å ÷ à í è å. ÕëÔÏ — õëîðîôèëëüíûé ôîòîñèíòåòè÷åñêèé ïîòåíöèàë ëèñòüåâ, ÌÑ âåòñòâåííî ôàçû ìîëî÷íîé ñïåëîñòè è ìîëî÷íî-âîñêîâîé ñïåëîñòè. Fòåîð. 05 01 4,84 9,85 3,98 4,84 7,20 9,85 3,98 7,20 è ÌÂÑ — ñîîò- 2. Óðîæàéíîñòü çåðíà (ö/ãà) ó ñîðòîâ îçèìîé ïøåíèöû Ñìóãëÿíêà è Ïåðåÿñëàâêà â ãîäû ñ ïîâûøåííîé òåìïåðàòóðîé âîçäóõà â ïåðèîä âåãåòàöèè íà ðàçíûõ ôîíàõ ìèíåðàëüíîãî ïèòàíèÿ (X±x; îïûòíîå ñåëüñêîõîçÿéñòâåííîå ïðîèçâîäñòâî Èíñòèòóòà ôèçèîëîãèè ðàñòåíèé è ãåíåòèêè ÍÀÍ Óêðàèíû, Êèåâñêàÿ îáë.) Âàðèàíò Êîíòðîëü N90P60K60S10 N120P90K90S20 Ñîðò Ñìóãëÿíêà 2009 2011 60,0±3,1 102,3±2,9 105,8±2,6 27,7±1,9 39,6±4,5 45,3±5,6 Ñîðò Ïåðåÿñëàâêà 2009 2011 55,1±2,6 98,4±2,7 99,0±3,8 24,5±1,4 53,0±3,8 56,4±2,8 Ïðîâåäåííûé êîððåëÿöèîííûé àíàëèç ïîêàçàë òåñíóþ ïîëîæèòåëüíóþ çàâèñèìîñòü ìåæäó çåðíîâîé ïðîäóêòèâíîñòüþ ïîñåâà è âåëè÷èíîé ÕëÔÏ çà ïåðèîä öâåòåíèå—ÌÂÑ äëÿ îáúåäèíåííîé âûáîðêè âàðèàíòîâ çà îáà ãîäà íàáëþäåíèé (ðèñ. 3). Âàæíî îòìåòèòü, ÷òî âûñîêèå è áëèçêèå ïî âåëè÷èíå çíà÷åíèÿ r (îò 0,93 äî 0,99) îòìå÷àëèñü òàêæå äëÿ âûáîÐèñ. 3. Çàâèñèìîñòü ìåæäó âåëè÷èíîé õëîðîôèëëüíîãî ðîê ïî êàæäîìó îòäåëüíîìó ôîòîñèíòåòè÷åñêîãî ïîòåíöèàëà (ÕëÔÏ) è óðîæàéíîñòüþ ãîäó, îòäåëüíîìó ñîðòó èëè ó ñîðòîâ îçèìîé ïøåíèöû â ãîäû ñ ïîâûøåííîé òåìïåðàîòäåëüíîìó âàðèàíòó ìèíåòóðîé âîçäóõà â ïåðèîä âåãåòàöèè: ▲, ∆ — ñîðò Ñìóãðàëüíîãî ïèòàíèÿ. ëÿíêà, ñîîòâåòñòâåííî 2009 è 2011 ãîäû; ♦, ◊ — ñîðò Òàêèì îáðàçîì, ïîÏåðåÿñëàâêà, ñîîòâåòñòâåííî 2009 è 2011 ãîäû (îïûòíîå ñåëüñêîõîçÿéñòâåííîå ïðîèçâîäñòâî Èíñòèòó- ëó÷åííûå ðåçóëüòàòû ñâèäåòà ôèçèîëîãèè ðàñòåíèé è ãåíåòèêè ÍÀÍ Óêðàèíû, òåëüñòâóþò, ÷òî òåñíàÿ êîðÊèåâñêàÿ îáë.). ðåëÿöèÿ ìåæäó èíòåãðàëüíûì ïîêàçàòåëåì ìîùíîñòè ôîòîñèíòåòè÷åñêîãî àïïàðàòà — õëîðîôèëëüíûì ôîòîñèíòåòè÷åñêèì ïîòåíöèàëîì (ÕëÔÏ) ëèñòüåâ è óðîæàéíîñòüþ ñîõðàíÿåòñÿ ó îçèìîé ïøåíèöû è â óñëîâèÿõ ïîâûøåííûõ òåìïåðàòóð. Äëÿ ëþ93 áîãî ôàêòîðà, âûçûâàþùåãî âàðüèðîâàíèå ïîêàçàòåëåé (óñëîâèÿ ãîäà, ôîí ìèíåðàëüíîãî ïèòàíèÿ, ñîðò), íàáëþäàåìàÿ âçàèìîñâÿçü ìîæåò áûòü îïèñàíà îáùèì óðàâíåíèåì ðåãðåññèè. Ïîâûøåíèå òåìïåðàòóðû âîçäóõà â ïåðèîä âåãåòàòèâíîãî ðîñòà è ðåïðîäóêòèâíîãî ðàçâèòèÿ óìåíüøàëî ðàçìåðû àññèìèëÿöèîííîé ïîâåðõíîñòè, ñîäåðæàíèå õëîðîôèëëà è ïðîäîëæèòåëüíîñòü ôóíêöèîíèðîâàíèÿ ëèñòüåâ, ÷òî ïðèâîäèëî ê ñíèæåíèþ óðîæàéíîñòè. Óâåëè÷åíèå ïëîùàäè ëèñòüåâ è ñîäåðæàíèÿ õëîðîôèëëà ïðè âíåñåíèè áîëåå âûñîêèõ äîç óäîáðåíèé ñîïðîâîæäàëîñü ïðîïîðöèîíàëüíûì âîçðàñòàíèåì óðîæàéíîñòè êàê ïðè óìåðåííîì, òàê è ïðè ñóùåñòâåííîì ïîâûøåíèè òåìïåðàòóð â ïåðèîä âåãåòàöèè ó îáîèõ ñîðòîâ, õîòÿ ïîñëåäíèå äîâîëüíî çíà÷èòåëüíî ðàçëè÷àëèñü ïî ÷óâñòâèòåëüíîñòè ê óñëîâèÿì âûðàùèâàíèÿ. Áîëüøàÿ ïðîäóêòèâíîñòü öåíîçîâ îçèìîé ïøåíèöû â âàðèàíòå ñ âíåñåíèåì âûñîêèõ äîç óäîáðåíèé áûëà îáóñëîâëåíà, âî-ïåðâûõ, áîëüøèì êîëè÷åñòâîì õëîðîôèëëà â ëèñòüÿõ, âî-âòîðûõ, áîëåå ïðîäîëæèòåëüíûì ïåðèîäîì, â òå÷åíèå êîòîðîãî ëèñòîâîé èíäåêñ â ïîñåâàõ ó ñîðòîâ îïòèìàëåí. Àâòîðû áëàãîäàðÿò íà÷àëüíèêà Ãèäðîìåòöåíòðà Óêðàèíû Í.È. Êóëüáèäó è íà÷àëüíèêà îòäåëà ìåòåîïðîãíîçîâ Ë.È. Ñàâ÷åíêî çà ïðåäîñòàâëåííûå äàííûå ìåòåîíàáëþäåíèé ïî Êèåâó è Êèåâñêîé îáëàñòè. ËÈÒÅÐÀÒÓÐÀ 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 94 À í ä ð è à í î â à Þ.Å. Õëîðîôèëëüíûå èíäåêñû è õëîðîôèëëüíûå ôîòîñèíòåòè÷åñêèå ïîòåíöèàëû — êðèòåðèè îöåíêè ïîòåíöèàëüíîé ïðîäóêòèâíîñòè ñåëüñêîõîçÿéñòâåííûõ ðàñòåíèé. Àâòîðåô. äîêò. äèñ. Ì., 1998. Ø à ä ÷ è í à Ò.Ì., Ï ð ÿ ä ê і í à Ã.Î., Ì î ð ã ó í Â.Â. Çâ'ÿçîê ìіæ õàðàêòåðèñòèêàìè ôîòîñèíòåòè÷íîãî àïàðàòó òà çåðíîâîþ ïðîäóêòèâíіñòþ ó ðіçíèõ ñîðòіâ îçèìîї ïøåíèöі.  ñá.: Äîñÿãíåííÿ і ïðîáëåìè ãåíåòèêè, ñåëåêöії і áіîòåõíîëîãії. Ò. 2. Êèїâ, 2007: 410-415. K u t a s y E., C s a j b o k J., H u n y a d i B.E. Relations between yield and photosynthetic activity of winter wheat varieties. Cereal Res. Commun., 2005, 1: 173-176 (doi: 10.1556/CRC.33.2005.1.42). Ï ð ÿ ä ê і í à Ã.Î., Ñ ò à ñ è ê Î.Î. Ñïîñіá ïðîãíîçóâàííÿ âðîæàéíîñòі îçèìîї ïøåíèöі. Ïàòåíò íà êîðèñíó ìîäåëü 67232 (Óêðàїíà) ÌÏÊ (2012.01) À 01G 7/00 A01G 1/00. Èíñòèòóò ôèçèîëîãèè ðàñòåíèé è ãåíåòèêè ÍÀÍ Óêðàèíû (Óêðàèíà). ¹ u 2011 086464. Çàÿâë. 06.0.11. Îïóáë. 10.02.12. Áþë. ¹ 3. À í ä ð è à í î â à Þ.Å., Ò à ð ÷ å â ñ ê è é È.À. Õëîðîôèëë è ïðîäóêòèâíîñòü ðàñòåíèé. Ì., 2000. C a l d e r i n i D.F., D r e c c e r M.F., S l a f e r G.A. Consequences of plant breeding on biomass growth, radiation interception and radiation use efficiency in wheat. Field Crop Res., 1997, 3: 271-281. L i Q., C h e n Y., L i u M., Z h o u X., Y u S., D o n g B. Effect of irrigation and planting patterns on radiation use efficiency and yield of winter wheat in North China. Agricultural Water Management, 2008, 4: 469-476 (doi: 10.1016/j.agwat.2007.11.010). Ä ó ä å í ê î Í.Â., À í ä ð è à í î â à Þ.Å., Ì à ê ñ þ ò î â à Í.Í. Ôîðìèðîâàíèå õëîðîôèëëüíîãî ôîòîñèíòåòè÷åñêîãî ïîòåíöèàëà ïøåíèöû â ñóõîé è âëàæíûé ãîäû. Ôèçèîëîãèÿ ðàñòåíèé, 2002, 5: 684-687. Ä å ð å í ä î â ñ ê à ÿ À., Æ î ñ à í Ñ. Õëîðîôèëëüíûå ïîêàçàòåëè è èõ ñâÿçü ñ ïðîäóêòèâíîñòüþ ðàñòåíèé îçèìîãî ÿ÷ìåíÿ. Stiinta agricola, 2008, 1: 4-6. S m i t h A.M., S t i t t M. Coordination of carbon supply and plant growth. Plant Cell Environ., 2007, 9: 1126-1149 (doi: 10.1111/j.1365-3040.2007.01708.x). Ì î ð ã ó í Â.Â., Ê è ð è ç è é Ä.À., Ø à ä ÷ è í à Ò.Ì. Ýêîôèçèîëîãè÷åñêèå è ãåíåòè÷åñêèå àñïåêòû àäàïòàöèè êóëüòóðíûõ ðàñòåíèé ê ãëîáàëüíûì èçìåíåíèÿì êëèìàòà. Ôèçèîëîãèÿ è áèîõèìèÿ êóëüòóðíûõ ðàñòåíèé, 2010, 1: 3-22. B a t t i s t i D.S., N a y l o r R.L. Historical warnings of future food insecurity with unprecedented seasonal heat. Science, 2009, 5911: 240-244 (doi: 10.1126/science.1164363). B a r n a b a s B., J a g e r K., F e h e r A. The effect of drought and heat stress on reproductive processes in cereals. Plant Cell Environ., 2008, 1: 11-38. F a r o o q M., B r a m l e y H., P a l t a J.A., S i d d i q u e K.H.M. Heat stress in wheat during reproductive and grain-filling phases. Critical Reviews in Plant Sciences, 2011, 6: 491-507. A s h r a f M., H a r r i s P.J.C. Photosynthesis under stressful environment: An overview. Photosynthetica, 2013, 51(2): 163-190 (doi: 10.1007/s11099-013-0021-6). B o j o v i ć B . , S t o j a n o v i ć J . Chlorophyll and carotenoid content in cultivars as function of mineral nutrition. Arch. Biol. Sci., 205, 4: 283-290. L o n g S.P., O r t D.R. More than taking the heat: crops and global change. Current Opin. Plant Biol., 2010, 3: 241-248 (doi: 10.1016/j.pbi.2010.04.008). 18. A i n s w o r t h E.A., O r t D.R. How do we improve crop production in a warming word? Plant Physiol., 2010, 2: 526-530. 19. Ì î ð ã ó í Â.Â., Ñ à í і í Є.Â., Ø â à ð ò à ó Â.Â., À ð ò å ì ÷ ó ê І.Ï., Ñ å ì å ð ó í ü Ò.Á., Î ì å ë ÿ í å í ê î Î.À. Êëóá 100 öåíòíåðіâ. Ñîðòè îçèìîї ïøåíèöі Іíñòèòóòó ôіçіîëîãії ðîñëèí і ãåíåòèêè ÍÀÍ Óêðàїíè òà ñèñòåìà çàõèñòó êîìïàíії Ñèíãåíòà. Êèїâ, 2008. 20. H i s c o x J. D., I s r a e l s t a m R.J. The method for the extraction of chlorophyll from leaf tissue without maceration. Can. J. Bot., 1979, 12: 1332-1334. 21. Ò à ð ÷ å â ñ ê è é È.À., À í ä ð è à í î â à Þ.Å. Ñîäåðæàíèå ïèãìåíòîâ êàê ïîêàçàòåëü ìîùíîñòè ðàçâèòèÿ ôîòîñèíòåòè÷åñêîãî àïïàðàòà ó ïøåíèöû. Ôèçèîëîãèÿ ðàñòåíèé, 1980, 2: 341-347. Èíñòèòóò ôèçèîëîãèè ðàñòåíèé è ãåíåòèêè ÍÀÍ Óêðàèíû, Ïîñòóïèëà â ðåäàêöèþ 19 ìàðòà 2014 ãîäà 03022 Óêðàèíà, ã. Êèåâ, óë. Âàñèëüêîâñêàÿ, 31/17, e-mail: pryadk@yandex.ru A RELATIONSHIP BETWEEN CHLOROPHYLL PHOTOSYNTHETIC POTENTIAL AND YIELD IN WINTER WHEAT (Triticum aestivum L.) AT ELEVATED TEMPERATURES G.À. Priadkina, O.O. Stasik, L.N. Mikhalskaya, V.V. Shvartau Institute of Plant Physiology and Genetics, National Academy of Sciences of Ukraine, 31/17, Vasilkovskaya St., Kiev, 03022 Ukraine, e-mail pryadk@yandex.ru Received March 19, 2014 doi: 10.15389/agrobiology.2014.5.88eng Abstract Development of the indices for selection of high-yielding crops and the models, forecasting agrocenosis yields, necessitate the analysis of the relationship between photosynthetic traits and productivity. Chlorophyll photosynthetic potential, characterizing the total amount of chlorophyll in the above-ground parts of plants (or in leaves) per unit of ground surface area during the growing season or a certain period, under optimal weather conditions correlated most closely with the magnitude of yield (T.M. Shadchina et al., 2007; E. Kutasy et al., 2005). Modern global climate change may primarily affect the heat-sensitive crops in particular winter wheat causing disruption in the relationship between photosynthetic traits and productivity. In field experiment, we examined the effects of increased air temperatures during the spring—summer growing season on performance of photosynthetic apparatus in the high-yielding varieties of common winter wheat Smuhlyanka and Pereyaslavka on the different levels of mineral nutrition (without fertilizers and fertilizing in the fall and at different phases of the spring—summer vegetation in doses N90P60K60S10 and N120P90K90S20). Investigations were carried out in 2009 and 2011 with elevated air temperatures during the growing season at the experimental agricultural station of the Institute of Plant Physiology and Genetics, National Academy of Sciences of Ukraine (Kyiv region.). It was shown that such weather conditions led to a decrease in chlorophyll content in the leaves, the size of crop surface and the duration of its operation, as well as grain yield. In all cases (both at high level of mineral nutrition and without fertilizers) leaf surface area in 2009 was higher than under the corresponding treatments in 2011 at all studied vegetation phases. Mineral fertilizers increased the leaf area index in both years by average 1.5-2.0 times (in Pereyaslavka at milk-wax ripeness phase it increased almost 5 times if compared to the control). Chlorophyll content in leaves of both varieties depended on year: it was 5.5-6.0 and 3.5-4.5 mg/dm2, respectively, in 2009 and 2011 if fertilizers were used, and 3.2-4.3 and 2.8 mg/dm2, respectively, in the control. In most cases, the maximum values of the pigments in the averaged sample of all green leaves from plant were observed in the phase of milk ripeness, probably, due to the complete withering away of leaves of lower layers. Mineral fertilizers increased the amount of chlorophyll in the leaves of both varieties. According to the results of dispersion analysis (F-Fisher test), the greatest influence on changes in the indices of the photosynthetic capacity in our experiments have the conditions of the year, the second most important factor was the level of mineral nutrition. The value of chlorophyll photosynthetic potential was more dependent on a combination of factors variety ½ year conditions and less on the interaction of the factors year conditions ½ mineral nutrition level and variety ½ mineral nutrition level. High and similar correlation coefficient values (0.93 to 0.99) between grain yield and chlorophyll photosynthetic potential were observed for data sets within single year, variety or mineral nutrition treatment, a strong positive dependence revealed for the combined data sets for both years. The data have shown that the close correlation between the leaf chlorophyll photosynthetic potential and yield of winter wheat is retained in the conditions of air high temperature and this dependence can be described by the same regression equation for any varying factors. Keywords: Triticum aestivum L., grain yield, leaf area index, chlorophyll, chlorophyll photosynthetic potential of leaves. 95
