Îáùåñòâî Ôèçèîëîãîâ ðàñòåíèé Ðîññèè Óðàëüñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò èì. À.Ì. Ãîðüêîãî Èíñòèòóò ôèçèîëîãèè ðàñòåíèé èì. Ê.À. Òèìèðÿçåâà ÐÀÍ Íàó÷íûé ñîâåò ïî ôèçèîëîãèè ðàñòåíèé è ôîòîñèíòåçó ÐÀÍ
ÃÎÄÈ×ÍÎÅ ÑÎÁÐÀÍÈÅ
ÎÁÙÅÑÒÂÀ ÔÈÇÈÎËÎÃÎÂ ÐÀÑÒÅÍÈÉ
ÐÎÑÑÈÈ
Ìåæäóíàðîäíàÿ êîíôåðåíöèÿ
«Ôèçèêî-õèìè÷åñêèå îñíîâû
ñòðóêòóðíî-ôóíêöèîíàëüíîé îðãàíèçàöèè ðàñòåíèé»
06 îêòÿáðÿ – 10 îêòÿáðÿ 2008 ã. Åêàòåðèíáóðã, ÐîññèÿÒÅÇÈÑÛ ÊÎÍÔÅÐÅÍÖÈÈ
Åêàòåðèíáóðã 2008Ïðåäñòàâëåíû ìàòåðèàëû òåçèñîâ Ìåæäóíàðîäíîé êîíôåðåíöèè «Ôèçèêî-õèìè÷åñêèå îñíîâû ñòðóêòóðíî-ôóíêöèîíàëüíîé îðãàíèçàöèè ðàñòåíèé», ïðîâåäåííîé â ðàìêàõ Ãîäè÷íîãî ñîáðàíèÿ Îáùåñòâà ôèçèîëîãîâ ðàñòåíèé Ðîññèè 6-11 îêòÿáðÿ 2008 ãîäà. Îðãàíèçàòîðû êîíôåðåíöèè - Îáùåñòâî ôèçèîëîãîâ ðàñòåíèé Ðîññèè, Íàó÷íûé ñîâåò ïî ôèçèîëîãèè ðàñòåíèé è ôîòîñèíòåçó ÐÀÍ, Óðàëüñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, Èíñòèòóò ôèçèîëîãèè ðàñòåíèé èì. Ê. À. Òèìèðÿçåâà ÐÀÍ. Íà êîíôåðåíöèè áûëè îáñóæäåíû ñîâðåìåííûå äîñòèæåíèÿ â îáëàñòè ôèçèêî-õèìè÷åñêèõ îñíîâ ôóíêöèîíèðîâàíèÿ ðàñòåíèé. Îñîáîå âíèìàíèå ó÷àñòíèêîâ óäåëåíî âîïðîñàì ìåõàíèçìîâ ðåãóëÿöèè è èíòåãðàöèè êëåòî÷íîãî ìåòàáîëèçìà, ñòðóêòóðíî-ôóíêöèîíàëüíîé îðãàíèçàöèè äîíîðíî-àêöåïòîðíûõ ñèñòåì â ðàñòåíèè, à òàêæå ðàçëè÷íûì àñïåêòàì ýêîëîãî-ôèçèîëîãè÷åñêèõ ìåõàíèçìîâ óñòîé÷èâîñòè è ïðîäóêöèîííîãî ïðîöåññà ðàñòåíèé.  ðàìêàõ êîíôåðåíöèè ïðîâåäåí êðóãëûé ñòîë, ïîñâÿùåííûé 80-ëåòèþ àêàäåìèêà À.Ò. Ìîêðîíîñîâà; äàí àíàëèç íàñëåäèÿ àêàäåìèêà À.Ò. Ìîêðîíîñîâà è ñîâðåìåííîãî ðàçâèòèÿ åãî èäåé.  êîíôåðåíöèè ïðèíÿëè ó÷àñòèå ó÷åíûå èç Ðîññèè, Áåëîðóññèè, Óêðàèíû, Òàäæèêèñòàíà, Ïîëüøè, Èíäèè, Ãåðìàíèè. Ìàòåðèàëû ïðåäíàçíà÷åíû äëÿ ôèçèîëîãîâ, áèîõèìèêîâ è ìîëåêóëÿðíûõ áèîëîãîâ ðàñòåíèé, áîòàíèêîâ è ýêîëîãîâ. Ïðîâåäåíèå Ìåæäóíàðîäíîé êîíôåðåíöèè ïîääåðæàíî Ðîññèéñêèì ôîíäîì ôóíäàìåíòàëüíûõ èññëåäîâàíèé; ãðàíò ÐÔÔÈ ¹ 08-04-06109-ã. Ðåäàêöèîííàÿ êîëëåãèÿ: È.Ñ. Êèñåëåâà (îòâ. ðåäàêòîð), Âë.Â. Êóçíåöîâ, Ã.À. Ðîìàíîâ, Ë.Ä. Êèñëîâ, Ñ.Í. ×ìîðà (ñåêðåòàðü) © Óðàëüñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò èì. À.Ì. Ãîðüêîãî, 2008 © Èíñòèòóò ôèçèîëîãèè ðàñòåíèé èì. Ê.À. Òèìèðÿçåâà ÐÀÍ, 2008
ÎÐÃÀÍÈÇÀÖÈÎÍÍÛÉ ÊÎÌÈÒÅÒ Ñîïðåäñåäàòåëè: Êóçíåöîâ Âë. Â. ÷ë.-êîðð. ÐÀÍ, Ìîñêâà Áóãðîâ Ä.Â. ê.è.í., ðåêòîð ÓðÃÓ, Åêàòåðèíáóðã Êèñåëåâà È.Ñ. ê.á.í., Åêàòåðèíáóðã ×ëåíû îðãêîìèòåòà: Áîðèñîâà Ã.Ã. ïðîô., Åêàòåðèíáóðã Âåñåëîâ À.Ï. ïðîô., Íèæíèé Íîâãîðîä Âîðîíèí Ï.Þ. ä.á.í., Ìîñêâà Ãàìàëåé Þ.Â. ÷ë.-êîðð. ÐÀÍ, Ñàíêò-Ïåòåðáóðã Ãîëîâêî Ò.Ê. ïðîô., Ñûêòûâêàð Ãîí÷àðîâà Ý.À ïðîô., Ñàíêò-Ïåòåðáóðã Åðìàêîâ Å. È. àêàä. ÐÀÑÕÍ, Ñàíêò-Ïåòåðáóðã Æèðîâ Â.Â. ÷ë.-êîðð. ÐÀÍ, Àïàòèòû Æóðàâëåâ Þ.Í. àêàä. ÐÀÍ, Âëàäèâîñòîê Èâàíîâ À.Î. ä.ô.-ì.í., ïðîðåêòîð ÓðÃÓ, Åêàòåðèíáóðã Êóçíåöîâ Â.Â. ä.á.í., Ìîñêâà Ìåäâåäåâ Ñ.Ñ. ïðîô., Ñàíêò-Ïåòåðáóðã Ðîìàíîâ Ã.À. ä.á.í., Ìîñêâà Ñàëÿåâ Ð.Ê. ÷ë.-êîðð. ÐÀÍ, Èðêóòñê Òèòîâ À.Ô. ÷ë.-êîðð. ÐÀÍ, Ïåðòîçàâîäñê Òðóíîâà Ò.È. ïðîô., Ìîñêâà Õîëîäîâà Â.Ï. ê.á.í., Ìîñêâà Õðÿíèí Â.Í. ïðîô., Ïåíçà ×èêîâ Â.È. ïðîô., Êàçàíü ×ìîðà Ñ.Í. ê.á.í., Ìîñêâà (ó÷åíûé ñåêðåòàðü) Øàâíèí Ñ.À. ïðîô., Åêàòåðèíáóðã Ëîêàëüíûé îðãêîìèòåò: Áîðçåíêîâà Ð.À. ê.á.í., Åêàòåðèíáóðã Äçþáåíêî Î.À. Åêàòåðèíáóðã Ìàëåâà Ì.Ã. ê.á.í., Åêàòåðèíáóðã Íåêðàñîâà Ã.Ô. ê.á.í., Åêàòåðèíáóðã Ôåäîñååâà Ã.Ï. ê.á.í., Åêàòåðèíáóðã Ôèðñîâ Í.Í. ïðîô., Åêàòåðèíáóðã ×óêèíà Í.Â. Åêàòåðèíáóðã
ÒÅÇÈÑÛ
ÊËÈÌÀÒÈ×ÅÑÊÈÅ ÔÀÊÒÎÐÛ ÔÎÐÌÈÐÎÂÀÍÈß NPP ÐÀÑÒÈ-ÒÅËÜÍÎÃÎ ÏÎÊÐÎÂÀ ÑÅÂÅÐÍÎÉ ÅÂÐÀÇÈÈ Âîðîíèí Ï.Þ. Èíñòèòóò ôèçèîëîãèè ðàñòåíèé èì. Ê.À. Òèìèðÿçåâà Ðîññèéñêîé àêàäåìèè íàóê, ã. Ìîñêâà, Ðîññèÿ, [email protected] Íà îñíîâàíèè ðåçóëüòàòîâ êîëëåêòèâà ó÷àñòíèêîâ ìíîãîëåòíåé ïðîãðàììû «Èçìåíåíèÿ îêðóæàþùåé ñðåäû è êëèìàòà: ïðèðîäíûå êàòàñòðîôû» ïðåäñòàâëåíû òåíäåíöèè èçìåíåíèÿ áèîëîãè÷åñêîé ïðîäóêòèâíîñòè ðàñòèòåëüíîãî ïîêðîâà Ñåâåðíîé Åâðàçèè ïîä âëèÿíèåì àðèäèçàöèè êëèìàòà. Òàê, èññëåäîâàíî äëèòåëüíîå âîçäåéñòâèå ïî÷âåííîé çàñóõè, ïåðåóâëàæíåíèÿ, òåìïåðàòóðû âîçäóõà è óäâîåííîé êîíöåíòðàöèè ÑÎ2 â àòìîñôåðå íà ôîòîñèíòåòè÷åñêèé àïïàðàò äðåâåñíûõ ðàñòåíèé. Ïðîàíàëèçèðîâàíû ìíîãîëåòíèå äàííûå ïî ïðîäóêòèâíîñòè ñåâåðíîé òàéãè â çàâèñèìîñòè îò òåìïåðàòóðíî-âëàæíîñòíîãî ðåæèìà. Âûÿâëåíû ôàêòîðû ëèìèòèðóþùèå ïðîäóêòèâíîñòü äðåâîñòîåâ íà óðîâíå ãàçî-îáìåíà ëèñòà è ôèòîöåíîçà â öåëîì.  ðàìêàõ ðåçåðâóàðíî-ïîòîêîâîé ìîäåëè ïðîäóêöèîííîãî ïðîöåññà íà òåððèòîðèè Ñåâåðíîé Åâðàçèè ïîëó÷åíû óòî÷íåííûå çíà÷åíèÿ ôîòîñèíòåòè÷åñêîãî ñòîêà óãëåðîäà (4.7 Ãò Ñ/ãîä) è NPP (4.4 Ãò Ñ/ãîä). Ïðåäñòàâëåíû äàííûå ïîëåâûõ è ýêñïåðèìåíòàëüíûõ èññëåäîâàíèé ïðîöåññà ìèêîãåííîãî ðàçëîæåíèÿ äðåâåñèíû, îïðåäåëÿþùèå âîçâðàòíûé ïîòîê óãëåðîäà èç ðàñòèòåëüíîãî ïîêðîâà â àòìîñôåðó.  ðåçóëüòàòå ïîëó÷åíî ïåðâîå ïðèáëèæåíèå óãëåðîäíîãî áàëàíñà ðàñòèòåëüíîãî ïîêðîâà Ñåâåðíîé Åâðàçèè. Ðàáîòà âûïîëíåíà ïðè ïîääåðæêå ïðîãðàììû ôóíäàìåíòàëüíûõ èññëåäîâàíèé ÐÀÍ «Èçìåíåíèÿ îêðóæàþùåé ñðåäû è êëèìàòà: ïðè-ðîäíûå êàòàñòðîôû», ÐÔÔÈ (ãðàíò No 06-04-48383), ãðàíòîâ ÍØ No 5551.2006.4, à òàêæå ÓðÎ ÐÀÍ (èíòåãðàöèîííûé ïðîåêò ñ ÄÂÎ ÐÀÍ).
SOME CLIMATE FACTORS HAVE BEEN EFFECTING A NORTHERN
EURASIA PLANT COVER NET PRIMARY PRODUCTION Voronin P.Yu.
K.A. Timiriazev Plant Physiology Institute of Rus. Acad. Sci., Moscow, Russia, [email protected]
General tendency of modern Northern Eurasia plant cover NPP changes ever done with climate aridization have been presented on the basis of data collected by participants of a many years’ fundamental research RAS program “Global Environmental and Climate Changes: Natural Disasters”. Effects of longterm soil water deprivation and fludity, temperature and dry air stress, double CO2 air content on photosynthesis apparatus had been seached. Northern taiga bioproduction dependence of temperature-humid regimen have been analysed. Climate factors ever limit forest stand’s bioproduction have been considered on different system levels: from leaf gas-exchange to phytocenosis. Northern Eurasia plant cover NPP is estimated as much as 4.4 Gton C/year and every year photosynthetic carbon incorporation to phytomass – 4.7 Gton C/year with the aid of an original organic carbon reservoir-flow model. Large scale Northern Eurasia carbon emission through fungi debris wood decomposition has also been estimated. The first step estimation for an organic carbon balance at the Northern Eurasia territory has been presented as a final result of the work.
This research has been being supported with the fundamental esearch program of RAS “Global Environmental and Climate Changes: Natural Disasters”, Russian Fund for Fundamental Research (grant # 06-04-48383), grant for Research School Support (# 5551.2006.4) and Ural Branch of RAS grant for fundamental research (integration project with Far East Branch of RAS).
ÔÎÒÎÑÈÍÒÅÇ Â ÑÈÑÒÅÌÅ ÄÎÍÎÐÍÎ-ÀÊÖÅÏÒÎÐÍÛÕ ÑÂßÇÅÉ Â ÐÀÑÒÅÍÈÈ Êèñåëåâà È.Ñ. Óðàëüñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò èì. À.Ì. Ãîðüêîãî, Åêàòåðèíáóðã, Ðîññèéñêàÿ Ôåäåðàöèÿ, [email protected] Öåëîñòíîñòü ðàñòåíèÿ êàê æèâîé ñèñòåìû îáåñïå÷èâàåòñÿ ñóùåñò-âîâàíèåì ìíîãîîáðàçíûõ ñâÿçåé è âçàèìîäåéñòâèé ìåæäó åãî îðãàíàìè, òêàíÿìè, êëåòêàìè.  îíòîãåíåçå ïðîèñõîäèò óñëîæíåíèå åãî ñòðóêòóðíîé è ôóíêöèîíàëüíîé îðãàíèçàöèè, ÷òî ïðèâîäèò ê èçìåíåíèþ òàêèõ ñèñòåìíûõ ñâîéñòâ êàê ôîòîñèíòåç, äûõàíèå, ðîñò, ðàçâèòèå è äð. Ïðîÿâëåíèåì ýòîãî ÿâëÿþòñÿ èçìåíåíèÿ äîíîðíî-àêöåïòîðíûõ ñèñòåì è ñâÿçåé ìåæäó íèìè. Åäèíñòâåííûì ïîñòàâùèêîì ïëàñòè÷åñêèõ âåùåñòâ â çåëåíûõ ðàñòåíèÿõ ÿâëÿåòñÿ ôîòîñèíòåç, òîãäà êàê ïîòðåáèòåëè äîñòàòî÷íî ðàçíîîáðàçíû: ïðîöåññû ðîñòà è ðàçâèòèÿ, çàïàñàíèå âåùåñòâ, äûõàíèå, ìåõàíèçìû îáåñïå÷åíèÿ óñòîé÷èâîñòè è ðåïàðàöèè è äðóãèå. Âçàèìîñâÿçü ôîòîñèíòåçà è ðîñòà, ðåàëèçóåìàÿ ÷åðåç òðàíñïîðò ôîòîàññèìèëÿòîâ è èõ óòèëèçàöèþ â àòòðàãèðóþùèõ òêàíÿõ è îðãàíàõ ðàñòåíèÿ, ñîñòàâëÿåò îñíîâó äîíîðíî-àêöåïòîðíûõ ñâÿçåé. Ñîãëàñíî êîíöåïöèè À.Ò. Ìîêðîíîñîâà âåäóùàÿ ðîëü â ýòèõ âçàèìîîòíîøåíèÿõ ïðèíàäëåæèò ðîñòó, îäíàêî ðîëü ôîòîñèíòåçà íå ñâîäèòñÿ òîëüêî ê òðîôè÷åñêîìó îáåñïå÷åíèþ ðàñòåíèÿ. Ýíäîãåííàÿ ðåãóëÿöèÿ è êîîð-äèíàöèÿ ôóíêöèé ôîòîñèíòåçà è ðîñòà âêëþ÷àåò, ïðåæäå âñåãî, ãåíåòè÷åñêèé è ãîðìîíàëüíûé êîíòðîëü. Îòäåëüíûå ñòîðîíû ýòîé ðåãóëÿöèè õîðîøî èçó÷åíû. Îäíàêî ìåæäó ìîëåêóëÿðíî-ãåíåòè÷åñêèì óðîâíåì ðåãóëÿöèè è ãîðìîíàëüíîé è òðîôè÷åñêîé ðåãóëÿöèåé â öåëîì ðàñòåíèè ëåæèò îãðîìíîå ÷èñëî ïðîìåæóòî÷íûõ ìåòàáîëè÷åñêèõ çâåíüåâ, íà÷àëî êîòîðûõ – â ïðîöåññàõ ïåðâè÷íîãî óñâîåíèÿ ÑÎ2. Îöåíêà ðîëè ôîòîñèíòåçà â äîíîðíî-àêöåïòîðíîé ñèñòåìå ðàñòåíèÿ ïðåäïîëàãàåò êîìïëåêñíîå èññëåäîâàíèå ñòðóêòóðíî-ôóíêöèîíàëüíîé îðãàíèçàöèè ôîòîñèíòåòè÷åñêîãî àïïàðàòà ðàñòåíèÿ, òðàíñïîðòà, ïóòåé óòèëèçàöèè è ðåóòèëèçàöèè ôîòîàññèìèëÿòîâ â äîíîðíûõ è àêöåïòîðíûõ îðãàíàõ. Èññëåäîâàíèå ðîñòà è ðàçâèòèÿ èíäèâèäóàëüíîé ôîòîàâòîòðîôíîé êëåòêè (ìîäåëü – ðàçíîâîçðàñòíûå çîíû ïåðâîãî ëèñòà ÿ÷ìåíÿ), îòäåëüíîãî ëèñòà, öåëîãî ðàñòåíèÿ ïîêàçàë, ÷òî â äîíîðíûõ îðãàíàõ ñòàíîâëåíèå è ðàçâèòèå ôîòîñèíòåòè÷åñêîé ôóíêöèè è èìïîðòíî-ýêñïîðòíîé ñïîñîáíîñòè ïðîèñõîäèò íà ôîíå îñëàáëåíèÿ ãåòåðîòðîôíîé ôèêñàöèè ÑÎ2, èçìåíå-íèÿ ïàðàìåòðîâ ìåçîñòðóêòóðû ôîòîòðîôíûõ òêàíåé (îïòèìèçàöèÿ óñëîâèé äèôôóçèè ÑÎ2), ôîòîñèíòåòè÷åñêîãî ìåòàáîëèçìà è ïîñòôî-òîñèíòåòè÷åñêîé ìîäèôèêàöèè ôîòîñèíòàòîâ, à òàêæå ãîðìîíàëüíîãî
ñòàòóñà. Þâåíèëüíûé ëèñò, êàê ïîòðåáèòåëü ñîáñòâåííûõ àññèìèëÿòîâ, èñïîëüçóåò èõ, ïðåæäå âñåãî, íà ñèíòåç ñòðóêòóðíûõ ñîåäèíåíèé – öåë-ëþëîçû, ãåìèöåëöåë-ëþëîçû, áåëêîâ, à çðåëûé ëèñò-äîíîð – íà îáðàçîâàíèå òðàíñïîðòíûõ ôîðì óãëåâîäîâ – ñàõàðîçû è îëèãîñàõàðîâ. Ñ âîçðàñòîì ïðîèñõîäèò ðàçâèòèå ýêñïîðòíîé àêòèâíîñòè ëèñòà, óìåíüøåíèå ñêîðîñòè ñèíòåçà ñòðóêòóðíûõ óãëåâîäîâ, óâåëè÷åíèå äîëè ñïèðòîâîäîðàñòâîðèìûõ âåùåñòâ, ÷òî êîððåëèðóåò ñ óìåíüøåíèåì ñîäåðæàíèÿ ÀÁÊ è ÈÓÊ è óâåëè÷åíèåì èõ ñîîòíîøåíèÿ. Âåðîÿòíî, ÀÁÊ ìîæåò âûñòóïàòü êàê ôàêòîð ðåãóëÿöèè ïóòåé óòèëèçàöèè ôîòîñèíòàòîâ â ëèñòå: 1) â íàïðàâëåíèè ñèíòåçà ñòðóêòóðíûõ âûñîêîìîëåêóëÿðíûõ ñîåäèíåíèé (öåëëþëîçà, ãåìèöåëëþëîçà, áåëêè), íåîáõîäèìûõ äëÿ îáåñïå÷åíèÿ ðîñòà êëåòîê ëèñòà èëè 2) â íàïðàâëåíèè ñèíòåçà ðàñòâîðèìûõ óãëåâîäîâ – òðàíñïîðòíûõ ôîðì àññèìèëÿòîâ. Ïðè âûñîêîé êîíöåíòðàöèè ýòîãî ãîðìîíà â áîëüøåé ñòåïåíè ðåàëèçóåòñÿ ïåðâîå íàïðàâëåíèå, à ïðè íèçêîé – âòîðîå. ÈÓÊ è öèòîêèíèíû òàêæå îïðåäåëÿþò íàïðàâëåíèå ìåòà-áîëèçàöèè ôîòîñèíòàòîâ â ëèñòå. Ïîäîáíûå ãîðìîíàëüíûå è ñòðóêòóðíî-ôóíêöèîíàëüíûå ïåðåñòðîéêè ïðîèñõîäÿò è â àêöåïòîðíûõ îðãàíàõ. Òàê, â êîëîñå ñèíòåçèðîâàííûå èëè òðàíñïîðòèðîâàííûå óãëåâîäû èñïîëüçóþòñÿ ïåðâîíà÷àëüíî íà ðîñò åãî ýëåìåíòîâ (ìåòàáîëèçàöèÿ êàê ñîáñòâåííûõ ôîòîñèíòàòîâ, òàê è ïîñòóïèâøèõ àññèìèëÿòîâ â íàïðàâ-ëåíèè ñòðóêòóðíûõ óãëåâîäîâ è áåëêîâ), à ïîçäíåå – íà çàïàñàíèå (ïðåèìóùåñòâåííûé ñèíòåç êðàõìàëà â çåðíîâêàõ èç ïðîäóêòîâ ôîòî-ñèíòåçà ëèñòüåâ è/èëè óãëåâîäîâ, ïîñòóïàþùèõ èç âòîðè÷íûõ äîíîðîâ). Èçìåíåíèå äîíîðíî-àêöåïòîðíûõ ñâÿçåé â îíòîãåíåçå îáóñëîâëåíî ñìåíîé è ïîñòàâùèêîâ, è ïîòðåáèòåëåé àññèìèëÿòîâ. Ó çëàêîâ â ïðåäåëàõ ãëàâíîãî ïîáåãà â ðàçíûå ïåðèîäû îíòîãåíåçà ïåðâè÷íûìè äîíîðàìè ïëàñòè÷åñêèõ âåùåñòâ ÿâëÿþòñÿ ëèñòüÿ, çåëåíûé êîëîñ (åãî îñòè è ÷åøóè), âåðõíåå ìåæäîóçëèå ñîëîìèíû, à àêöåïòîðàìè – ãåòåðîòðîôíûå àêòèâíî ðàñòóùèå îðãàíû, ãëàâíûì îáðàçîì, êîëîñ è ñîëîìèíà. Îäíàêî îíè ìîãóò âûñòóïàòü â êà÷åñòâå âòîðè÷íûõ äîíîðîâ è ïîñòàâëÿòü ðàñòóùèì çåðíîâêàì ïðîäóêòû ìåòàáîëèçàöèè ðàíåå äåïîíèðîâàííûõ âåùåñòâà, ãëàâíûì îáðàçîì, ôðóêòîçàíîâ. Òàêèì îáðàçîì, ðîëü ôîòîñèíòåçà â ñèñòåìå äîíîðíî-àêöåïòîðíûõ ñâÿçåé â ðàñòåíèè èñêëþ÷èòåëüíî âàæíà íå òîëüêî êàê èñòî÷íèêà îðãà-íè÷åñêîãî âåùåñòâà, íî è êàê ïðîöåññà, îíòîãåíåòè÷åñêèå îñîáåííîñòè êîòîðîãî îïðåäåëÿþò õàðàêòåð ìåòàáîëèçìà êàê ñàìèõ ôîòîàâòîòðîôíûõ ýëåìåíòîâ ðàñòåíèÿ, òàê è ïîòðåáëÿþùèõ ãåòåðîòðîôíûõ êëåòîê, òêàíåé, îðãàíîâ è â çíà÷èòåëüíîé ñòåïåíè îïðåäåëÿþò èõ ôîðìèðîâàíèå.
ÊÓËÜÒÈÂÈÐÓÅÌÛÅ in vitro ÊÎÐÍÈ ËÅÊÀÐÑÒÂÅÍÍÛÕ ÐÀÑÒÅÍÈÉ ÊÀÊ ÌÎÄÅËÜÍÀß ÑÈÑÒÅÌÀ ÈÇÓ×ÅÍÈß ÏÐÎÑÒÐÀÍÑÒÂÅÍÍÎÉ ÎÐÃÀÍÈÇÀÖÈÈ ÂÒÎÐÈ×ÍÎÃÎ ÌÅÒÀÁÎËÈÇÌÀ Êóçîâêèíà È.Í. Èíñòèòóò ôèçèîëîãèè ðàñòåíèé èì. Ê.À.Òèìèðÿçåâà ÐÀÍ, Ìîñêâà, Ðîññèÿ, [email protected] Êîðíè ëåêàðñòâåííûõ ðàñòåíèé ÿâëÿþòñÿ ìåñòîì ñèíòåçà ðàçíîîá-ðàçíûõ âòîðè÷íûõ ìåòàáîëèòîâ, êîòîðûå íàõîäÿò øèðîêîå ïðèìåíåíèå â ìåäèöèíñêîé è ïèùåâîé ïðîìûøëåííîñòè. Îäíîâðåìåííî ñ ýòèì íèçêîìîëåêóëÿðíûå ñîåäèíåíèÿ, îáðàçóþùèåñÿ â ìîëîäûõ è ôóíêöèî-íàëüíî àêòèâíûõ çîíàõ êîðíåé, ìîãóò âûïîëíÿòü ðîëü ñèãíàëüíûõ âåùåñòâ ïðè êîíòàêòàõ ñ ïî÷âåííûìè ìèêðîîðãàíèçìàìè è ïðè àëëåëîïàòè÷åñêèõ âçàèìîîòíîøåíèÿõ ðàñòåíèé â áèîöåíîçàõ. Âîïðîñ î òîì, íàñêîëüêî ñîñòàâ âòîðè÷íûõ âåùåñòâ ôèçèîëîãè÷åñêè àêòèâíûõ çîí êîðíÿ èäåíòè÷åí ñîñòàâó êîìïîíåíòîâ åãî çðåëîé îáëàñòè, îñòàâàëñÿ äîëãîå âðåìÿ îòêðûòûì, ÷òî îáúÿñíÿëîñü ñëîæíîñòüþ ïîëó÷åíèÿ äîñòàòî÷íîãî äëÿ õèìè÷åñêîãî àíàëèçà ðàñòèòåëüíîãî ìàòåðèàëà ïðè ðàáîòå ñ öåëûì ðàñòåíèåì.  íàñòîÿùåå âðåìÿ îòâåò íà ýòîò âîïðîñ ìîæíî íàéòè, èñïîëüçóÿ â êà÷åñòâå îáúåêòà, ãåíåòè÷åñêè òðàíñôîðìèðîâàííûå êîðíè («hairy roots»), ïîëó÷åííûå ïóòåì èíîêóëÿöèè ñòåðèëüíûõ ðàñòåíèé äèêèìè øòàììàìè Agrobacterium rhizogenes. Hairy roots, èìåþùèå ñòðóêòóðó ïåðâè÷íûõ êîðíåé, áûñòðî ðàñòóò íà áåçãîðìîíàëüíûõ ïèòàòåëüíûõ ñðåäàõ â óñëîâèÿõ in vitro è â ðåçóëüòàòå ìîùíîãî âåòâëåíèÿ îáðàçóþò áîëüøîå êîëè÷åñòâî àïåêñîâ, ÷òî ïîçâîëÿåò ïîëó÷àòü îäíîðîäíûé ðàñòèòåëüíûé ìàòåðèàë äëÿ õèìè÷åñêèõ àíàëèçîâ è ìèêðîñêîïèðîâàíèÿ. Óäà÷íûì îáúåêòîì äëÿ ïðîâåäåíèÿ äàííîé ðàáîòû ïîñëóæèëè hairy roots ðóòû äóøèñòîé (Ruta graveolens L.), êîòîðàÿ ñèíòåçèðóåò áîëåå 120 âòîðè÷íûõ âåùåñòâ. Îñíîâíàÿ ÷àñòü ìåòàáîëèòîâ ðóòû èìååò èíòåí-ñèâíóþ ôëóîðåñöåíöèþ, ÷òî îáëåã÷àåò èõ äåòåêöèþ â òêàíÿõ ðàñòåíèÿ ñ ïîìîùüþ ñîâðåìåííîé ìèêðîñêîïè÷åñêîé òåõíèêè. Ïðè èññëåäîâàíèè ðàñïðåäåëåíèÿ âòîðè÷íûõ ìåòàáîëèòîâ â òðåõ çîíàõ ðàñòóùèõ êîí÷èêîâ êîðíåé ðóòû - ìåðèñòåìàòè÷åñêîé, ðàñòÿæåíèÿ è â çîíå äèôôåðåíöèàöèè áûëà èñïîëüçîâàíà àíàëèòè÷åñêàÿ è ïðåïàðàòèâíàÿ ÂÝÆÕ ýêñòðàêòîâ â ñî÷åòàíèè ñ 13C, 1H ßÌÐ-ñïåêòðîñêîïèåé è ìàññ-ñïåêòðîìåòðèåé, à òàêæå êîíôîêàëüíàÿ ëàçåðíàÿ ñêàíèðóþùàÿ ìèêðîñêîïèÿ (CLSM), ôëóîðåñ-öåíòíàÿ ìèêðîñêîïèÿ è ìèêðîñïåêòðôëóîðîìåòðèÿ. Êîìïëåêñíûé ïîäõîä îáåñïå÷èâàë ïîëó÷åíèå ïîëíîé êàðòèíû ëîêàëèçàöèè âòîðè÷íûõ âåùåñòâ â êîðíåâûõ êëåòêàõ è äîïîëíÿë ðåçóëüòàòû ÂÝÆÕ. Óñòàíîâëåíà ÷åòêàÿ çàêîíîìåðíîñòü ðàñïðåäåëåíèÿ âòîðè÷íûõ ìåòàáîëèòîâ â
îñíîâíûõ çîíàõ ðîñòà êîðíåé ðóòû. Ìåðèñòåìàòè÷åñêàÿ çîíà, ïðåäñòàâ-ëåííàÿ ìåðèñòåìîé è ïðèëåãàþùèì ê íåé êîðíåâûì ÷åõëèêîì, ñîäåðæèò òîëüêî ãëèêîçèäû äâóõ àêðèäîíîâûõ àëêàëîèäîâ – ãðàâàêðèäîíäèîëà è òðèîëà, ñ êîòîðûìè ñâÿçàíà èíòåíñèâíàÿ ôëóîðåñöåíöèÿ êîðíåâûõ àïåêñîâ ðóòû. Ðåçóëüòàòû CLSM ïîêàçàëè, ÷òî ãëèêîçèäû àêðèäîíîâûõ àëêàëîèäîâ ëîêàëèçîâàíû òîëüêî â âàêóîëÿõ êëåòîê êîðíåâîãî ÷åõëèêà è îòñóòñòâóþò â êëåòêàõ ìåðèñòåìû.  çîíå ðàñòÿæåíèÿ êîðíåé ðóòû îáíàðóæåíû êóìàðèíû è õèíîëèíîâûå àëêàëîèäû, êîòîðûå ñîäåðæàòñÿ â âèäå ãðàíóëèðîâàííûõ âêëþ÷åíèé â ïàðåíõèìàòè÷åñêèõ èäèîáëàñòàõ. Çîíà äèôôåðåíöèàöèè, ãóñòî ïîêðûòàÿ êîðíåâûìè âîëîñêàìè, ñîäåðæèò ïîëíûé íàáîð õàðàêòåðíûõ äëÿ êîðíåé ðóòû àêðèäîíîâûõ àëêàëîèäîâ, ñðåäè êîòîðûõ äîìèíèðóþò ëèïîôèëüíûå àêðèäîíû. Îíè ëîêàëèçîâàíû â ëèïîôèëüíûõ êàïëÿõ êîðíåâûõ âîëîñêîâ, ãðóïïèðóþùèõñÿ â ñðåäíåé ÷àñòè è â êîí÷èêàõ êîðíåâûõ âîëîñêîâ, îòêóäà âûäåëÿþòñÿ â âèäå ýêññóäàòîâ. Îñîáûé èíòåðåñ ïðåäñòàâëÿþò ïîãðàíè÷íûå êëåòêè («border cells») êîðíåé ðóòû, êîòîðûå ÿâëÿþòñÿ êëåòêàìè êàëèïòðû, îòäåëÿþ-ùèìèñÿ îò êîí÷èêà êîðíÿ ïî ìåðå åãî ðîñòà. Ìíîãî÷èñëåííûå border cells (ÂÑ) ïîñëå óòðàòû êîíòàêòà êàëèïòðû ñ ìåðèñòåìîé ïðîäîëæàþò ñâîå ñóùåñòâîâàíèå â óñëîâèÿõ in vitro â æèäêîé ïèòàòåëüíîé ñðåäå â òå÷åíèå íåñêîëüêèõ íåäåëü. Ðåçóëüòàòû ÂÝÆÕ è CLSM ïîêàçàëè, ÷òî ñîñòàâ âòî-ðè÷íûõ ìåòàáîëèòîâ ÂÑ ñóùåñòâåííî îòëè÷àåòñÿ îò ñîñòàâà êîìïîíåíòîâ êëåòîê êàëèïòðû, è â íèõ îáíàðóæèâàåòñÿ ïîëíûé íàáîð õàðàêòåðíûõ äëÿ êîðíåé ðóòû ëèïîôèëüíûõ âòîðè÷íûõ âåùåñòâ. Ïðè ìèêðîñêîïè÷åñêîì èçó÷åíèè ÂÑ óñòàíîâëåíî, ÷òî êàæäàÿ èç íèõ ñîäåðæèò ëèïîôèëüíîå âêëþ÷åíèå, ÿâëÿþùååñÿ ìåñòîì àêêóìóëÿöèè õàðàêòåðíûõ äëÿ êîðíåé ðóòû ãèäðîôîáíûõ ñîåäèíåíèé. Ýòî îçíà÷àåò, ÷òî ÂÑ ñïîñîáíû ê ñàìîñòîÿòåëüíîìó ñèíòåçó è ìåòàáîëèçìó âòîðè÷íûõ âåùåñòâ, òèïè÷íûõ äëÿ êîðíåé öåëîãî ðàñòåíèÿ ðóòû.  ïî÷âåííûõ óñëîâèÿõ ÂÑ áûñòðî ïîãèáàþò, è ñîäåðæàùèåñÿ â íèõ ìåòàáîëèòû, â êîòîðûõ òàêæå, êàê è â ýêññóäàòàõ êîðíåâûõ âîëîñêîâ, äîìèíèðóþò àêðèäîíîâûå àëêàëîèäû, ïîïàäàþò â ñóáñòðàò, ãäå ñîçäàþò çîíû, îáîãàùåííûå ñîåäèíåíèÿìè, èìåþùèìè ýêñïåðèìåíòàëüíî äîêàçàííóþ àíòèáàêòåðèàëüíóþ àêòèâ-íîñòü. Ðåçóëüòàòû èçó÷åíèÿ ïðîñòðàíñòâåííîé îðãàíèçàöèè âòîðè÷íîãî ìåòàáîëèçìà ãåíåòè÷åñêè òðàíñôîðìèðîâàííûõ êîðíåé ðóòû îáúÿñíÿþò ïðè÷èíó èõ íåñïîñîáíîñòè ê óñòàíîâëåíèþ êîíòàêòîâ ñ ìèêîðèçíûìè ãðèáàìè. Âòîðè÷íûå ìåòàáîëèòû, ïîñòóïàþùèå â ïî÷âó ïðè ãèáåëè ïîãðàíè÷íûõ êëåòîê êîðíåâîãî ÷åõëèêà è âûäåëÿåìûå êîðíåâûìè âîëîñêàìè ðàñòóùèõ êîðíåé, ìîãóò òàêæå ïðèíèìàòü àêòèâíîå ó÷àñòèå â àëëåëîïàòè÷åñêèõ âçàèìîîòíîøåíèÿõ ðàñòåíèé â áèîöåíîçàõ. Ðàáîòà âûïîëíåíà ïðè ïîääåðæêå Ôîíäà Àëåêñàíäðà ôîí Ãóìáîëüäòà.
ÁÈÎÐÀÇÍÎÎÁÐÀÇÈÅ ÐÀÑÒÈÒÅËÜÍÎÃÎ ÌÈÐÀ ÓÐÀËÀ Êóëèêîâ Ï.Â. Áîòàíè÷åñêèé ñàä ÓðÎ ÐÀÍ, Åêàòåðèíáóðã, Ðîññèÿ, [email protected] Ïðîòÿæåííîñòü Óðàëüñêîé ãîðíîé ñòðàíû ñ ñåâåðà íà þã ñîñòàâëÿåò îêîëî 2000 êì – îò òóíäðîâîé äî þãà ñòåïíîé çîíû. Âî ôëîðå Óðàëà íàñ÷èòûâàåòñÿ îêîëî 3000 âèäîâ ñîñóäèñòûõ ðàñòåíèé, â òîì ÷èñëå îêîëî 2700 àáîðèãåííûõ è îêîëî 300 àäâåíòèâíûõ. Ýíäåìèçì, îïðåäåëÿþùèé ñâîåîáðàçèå è ñàìîáûòíîñòü ôëîðû Óðàëà, ñîñòàâëÿåò îêîëî 5% åå âèäîâîãî ñîñòàâà. Ýíäåìèêàìè Óðàëüñêîé ãîðíîé ñòðàíû ÿâëÿþòñÿ 90 àìôèìèêòè÷åñêèõ âèäîâ è ïîäâèäîâ ñîñóäèñòûõ ðàñòåíèé, à òàêæå íå ìåíåå 60 àïîìèêòè÷åñêèõ âèäîâ èç ðîäîâ Alchemilla (33), Hieracium (îêîëî 25), Ranunculus (3). Áîëüøèíñòâî ýíäåìèêîâ ñâÿçàíî ñ î÷åíü õàðàêòåðíûìè äëÿ Óðàëà ïåòðîôèòíûìè ôèòîöåíîçàìè ñ ðàçðåæåííûì ðàñòèòåëüíûì ïîêðîâîì è îñëàáëåííîé ìåæâèäîâîé êîíêóðåíöèåé – êàìåíèñòûìè ãîðíûìè òóíäðàìè, ñêàëüíûìè îáíàæåíèÿìè ïî áåðåãàì ðåê ëåñíîé çîíû è êàìåíèñòûìè ñòåïÿìè. Ëèøü î÷åíü íåìíîãèå èç óðàëüñêèõ ýíäåìèêîâ ðàñïðîñòðàíåíû îò Ïîëÿðíîãî äî Þæíîãî Óðàëà (Gypsophila uralensis, Thymus
paucifolius, Gagea samojedorum, Salix uralicola), òîãäà êàê áîëüøèíñòâî èõ
ñâîéñòâåííî îïðåäåëåííûì øèðîòíûì ñåêòîðàì Óðàëà.  òî æå âðåìÿ íà âñåì ïðîòÿæåíèè îò þæíîé ÷àñòè Ïðèïîëÿðíîãî Óðàëà äî öåíòðàëüíîé ÷àñòè Þæíîãî Óðàëà íàáëþäàåòñÿ îïðåäåëåííàÿ îáùíîñòü ñîñòàâà ôëîðû, äëÿ êîòîðîé õàðàêòåðíû ñïåöèôè÷åñêèå ýíäåìè÷íûå è äîâîëüíî ìíîãî÷èñëåííûå ðåëèêòîâûå ýëåìåíòû, ñâÿçûâàþùèå åå ñ ôëîðîé ãîð Þæíîé Ñèáèðè, îñîáåííî ñ Àëòàåì. Äîâîëüíî ñëîæíàÿ èñòîðèÿ ôîðìè-ðîâàíèÿ ôëîðû Óðàëà, â õîäå êîòîðîé ïðîèñõîäèëè ìíîãî÷èñëåííûå ìèãðàöèè àðêòè÷åñêèõ è àðêòîàëüïèéñêèõ ýëåìåíòîâ íà þã âäîëü ãîðíûõ õðåáòîâ, à ãîðíî-ñòåïíûõ âèäîâ – íà ñåâåð ïî ñêëîíàì ðå÷íûõ äîëèí, îáóñëîâèëà ñëîæíîñòü ñîâðåìåííîãî ïðîñòðàíñòâåííîãî ðàñïðåäåëåíèÿ ýëåìåíòîâ ôëîðû, íà êîòîðóþ íàêëàäûâàþòñÿ ÿâëåíèÿ, ñâÿçàííûå ñ âûñîòíî-ïîÿñíîé äèôôåðåíöèàöèåé ðàñòèòåëüíîãî ïîêðîâà è âëèÿíèåì ýäàôè÷åñêèõ ôàêòîðîâ (ñîñòàâà ãîðíûõ ïîðîä). Èçó÷åíèåì ôëîðû Óðàëà çàíèìàëèñü â XVIII â. È.Ã. Ãìåëèí, Ï.Ñ. Ïàëëàñ, È.Ï. Ôàëüê, È.Ã. Ãåîðãè, â XIX â. – Õ. Ô. Ëåññèíã, À. À. Ëåìàí, Ê.Ô. Ìåéíñãàóçåí, À.Ã. Øðåíê, Þ.Ê. Øåëëü, Ï.Í. Êðûëîâ, Ñ.È. Êîðæèíñêèé, À.ß. Ãîðäÿãèí, Î.Å. Êëåð, Ï.Â. Ñþçåâ, Í.È. Êóçíåöîâ, Ä.È. Ëèòâèíîâ, â XX â. – È.Ì. Êðàøåíèííèêîâ, Ð.Ð. Ïîëå, Â.Í. Ñóêà÷åâ, Á.Í. Ãîðîäêîâ, Â.Á. Ñî÷àâà, Ì.Ì. Èëüèí, Á.À. Ôåä÷åíêî, Â.Ñ. Ãîâîðóõèí, Ê.Í. Èãîøèíà, Ñ.Þ. Ëèïøèö, À.Ê. Íîñêîâ, Ë.Í. Òþëèíà, Ñ.Å. Êó÷åðîâñêàÿ, Ë.À. Óòêèí, À.Í. Ïîíîìàðåâ, Â.Á. Êóâàåâ,
Ï. Ë. Ãîð÷àêîâñêèé è äð. Ïåðâûå îáîáùàþùèå ðàáîòû ïî ôëîðå Óðàëà áûëè îïóáëèêîâàíû â êîíöå XIX – íà÷àëå XIX â. (Êðûëîâ, 1878–1885; Ôåä÷åíêî, Ôåä÷åíêî, 1893; Korshinsky, 1898; Ñþçåâ, 1912). Ïåðâîé ðàáîòîé, â êîòîðîé áûëè ïðåäñòàâëåíû ñâåäåíèÿ î ôëîðå âñåãî Óðàëà îò ïîáåðåæüÿ Ñåâåðíîãî Ëåäîâèòîãî îêåàíà äî þæíîé ãðàíèöû Ñðåäíåãî Óðàëà, ñòàëà «Ôëîðà Óðàëà» Â. Ñ. Ãîâîðóõèíà (1937).  1966 ã. áûë èçäàí ïåðâûé ðåãèîíàëüíûé îïðåäåëèòåëü ðàñòåíèé íà Óðàëå – «Îïðåäåëèòåëü ðàñòåíèé Áàøêèðñêîé ÀÑÑл ïîä ðåäàêöèåé Á. Ê. Øèøêèíà è Â. È. Ãðóáîâà, íàïèñàííûé ëåíèíãðàäñêèìè è óêðàèíñêèìè áîòàíèêàìè. Îáîáùàþùèå ðàáîòû ïî ôëîðå âûñîêîãîðèé Óðàëà áûëè îïóáëèêîâàíû Ê. Í. Èãîøèíîé (1966) è Ï. Ë. Ãîð÷àêîâñêèì (1966, 1975). Ôëîðà ñåâåðíîé ÷àñòè Óðàëà (â ïðåäåëàõ Ðåñïóáëèêè Êîìè, Íåíåöêîãî è ßìàëî-Íåíåöêîãî ÀÎ) íàøëà îòðàæåíèå âî «Ôëîðå ñåâåðî-âîñòîêà åâðîïåéñêîé ÷àñòè ÑÑÑл ïîä ðåäàêöèåé À. È. Òîëìà÷åâà (1974–1977), «Àðêòè÷åñêîé ôëîðå ÑÑÑл (1960– 1987) è ñâîäêå «Ðàñòèòåëüíûé ïîêðîâ è ðàñòèòåëüíûå ðåñóðñû Ïîëÿðíîãî Óðàëà» (2006).  òå÷åíèå ïîñëåäíèõ 20 ëåò äëÿ îáëàñòåé è ðåñïóáëèê Óðàëà áûë îïóáëèêîâàí ðÿä ðåãèîíàëüíûõ ôëîðèñòè÷åñêèõ ñâîäîê: «Îïðåäå-ëèòåëü âûñøèõ ðàñòåíèé Áàøêèðñêîé ÀÑÑл (1988, 1989), «Îïðåäå«Îïðåäå-ëèòåëü ñîñóäèñòûõ ðàñòåíèé Ñðåäíåãî Óðàëà» (1994) è êîíñïåêòû ôëîðû Ïåðìñêîé (Îâåñíîâ, 1997), Îðåíáóðãñêîé (Ðÿáèíèíà, 1998) è ×åëÿáèíñêîé îáëàñòåé (Êóëèêîâ, 2005; Ðÿçàíîâà, 2006). Íåñìîòðÿ íà äîâîëüíî âûñîêóþ èíòåíñèâíîñòü ôëîðèñòè÷åñêèõ èññëåäîâàíèé íà Óðàëå â ïîñëåäíèå ãîäû è íàëè÷èå êðóïíûõ ðåãèîíàëü-íûõ êîëëåêöèé (ãåðáàðèè Èíñòèòóòà ýêîëîãèè ðàñòåíèé è æèâîòðåãèîíàëü-íûõ ÓðÎ ÐÀÍ è Èíñòèòóòà áèîëîãèè Êîìè ÍÖ ÓðÎ ÐÀÍ – ïî 180 òûñ. ýêç., ãåðáàðèè Èíñòèòóòà áèîëîãèè Óôèìñêîãî ÍÖ ÐÀÍ è Ïåðìñêîãî ãîñóäàðñòâåííîãî óíèâåðñèòåòà – ïî 90 òûñ. ýêç.), ôëîðà Óðàëà äî ñèõ ïîð èññëåäîâàíà íåïîëíî è íåðàâíîìåðíî äàæå â îòíîøåíèè òàêñîíîìè÷åñêîãî ñîñòàâà. Îá ýòîì ñâèäåòåëüñòâóåò òîò ôàêò, ÷òî çà ïîñëåäíèå 10 ëåò ñîòðóäíèêàìè Áîòàíè÷åñêîãî ñàäà ÓðÎ ÐÀÍ (Ì. Ñ. Êíÿçåâûì è àâòîðîì) îïèñàíû 25 íîâûõ äëÿ íàóêè âèäîâ, 3 ïîäâèäà è 3 ìåæâèäîâûõ ãèáðèäà (íîòîâèäà), áîëüøèíñòâî êîòîðûõ ÿâëÿåòñÿ ýíäåìèêàìè Óðàëà. Îõðàíà ðàñòèòåëüíîãî ìèðà Óðàëà îñóùåñòâëÿåòñÿ ñèñòåìîé îñîáî îõðàíÿåìûõ ïðèðîäíûõ òåððèòîðèé, êëþ÷åâûìè ýëåìåíòàìè êîòîðîé ÿâëÿþòñÿ 10 ãîñóäàðñòâåííûõ çàïîâåäíèêîâ è 5 íàöèîíàëüíûõ ïàðêîâ.  8 çàïîâåäíèêàõ (Ïå÷îðî-Èëû÷ñêîì, Âèøåðñêîì, Äåíåæêèí Êàìåíü, Áàñåãè, Âèñèìñêîì, Èëüìåíñêîì, Áàøêèðñêîì, Îðåíáóðãñêîì ñòåïíîì) è 3 íàöèîíàëüíûõ ïàðêàõ (Þãûä-Âà, Ïðèïûøìèíñêèå áîðû, Çþðàòêóëü) èíâåíòàðèçàöèÿ ôëîðû ñîñóäèñòûõ ðàñòåíèé â îñíîâíîì çàâåðøåíà, è ïî åå ðåçóëüòàòàì îïóáëèêîâàíû ôëîðèñòè÷åñêèå ñïèñêè. Âî âñåõ îáëàñòÿõ è ðåñïóáëèêàõ Óðàëà â íàñòîÿùåå âðåìÿ èçäàíû ðåãèîíàëüíûå Êðàñíûå êíèãè.
ÃÎÐÌÎÍÀËÜÍÀß ÐÅÃÓËßÖÈß ÄÎÍÎÐÍÎ-ÀÊÖÅÏÒÎÐÍÛÕ ÑÂßÇÅÉ Ó ÐÀÑÒÅÍÈÉ Ðîíüæèíà Å.Ñ. ÔÃÎÓ ÂÏÎ «Êàëèíèíãðàäñêèé ãîñóäàðñòâåííûé òåõíè÷åñêèé óíèâåðñèòåò», ã. Êàëèíèíãðàä, Ðîññèÿ, [email protected]  ðàçâèòèå ðàçðàáîòàííîé À.Ò. Ìîêðîíîñîâûì êîíöåïöèè äîíîðíî-àêöåïòîðíûõ îòíîøåíèé â äîêëàäå ðàññìîòðåí âîïðîñ î ïðèðîäå ïðîöåññîâ, êîíòðîëèðóþùèõ òðàíñïîðò è ðàñïðåäåëåíèå àññèìèëÿòîâ â ðàñòåíèÿõ è èíòåãðèðóþùèõ ôîòîñèíòåçèðóþùèå, ïðîâîäÿùèå è ïîòðåáëÿþùèå òêàíè è îðãàíû â åäèíóþ äîíîðíî-àêöåïòîðíóþ ñèñòåìó. Îñîáîå âíèìàíèå óäåëåíî âëèÿíèþ ôèòîãîðìîíîâ, â ïåðâóþ î÷åðåäü, öèòîêèíèíîâ, íà âñå ýëåìåíòû äîíîðíî-àêöåïòîðíîé ñèñòåìû: ôîðìèðî-âàíèå è ôóíêöèîíèðîôîðìèðî-âàíèå ôîòîñèíòåòè÷åñêîãî àïïàðàòà, îòòîê àññèìèëÿòîâ èç ëèñòà, èõ ðàñïðåäåëåíèå ìåæäó ðàçëè÷íûìè îðãàíàìè è òêàíÿìè, îòëîæåíèå â çàïàñ è èñïîëüçîâàíèå íà ïðîöåññû æèçíåäåÿ-òåëüíîñòè, ðîñòà è ðàçâèòèÿ ðàñòåíèé. Ïðîàíàëèçèðîâàíû âîçìîæíûå ìåõàíèçìû ôîðìèðîâàíèÿ àêöåï-òîðíîé ñïîñîáíîñòè ïîòðåáëÿþùèõ è çàïàñàþùèõ çîí ðàñòåíèÿ, êîòîðàÿ ïðîÿâëÿåòñÿ â: 1). Ðåãóëÿöèè èíòåíñèâíîñòè è êà÷åñòâåííîé íàïðàâ-ëåííîñòè ôîòîñèíòåçà. 2). Àòòðàãèðóþùåì äåéñòâèè, ò.å. îðèåíòàöèè òðàíñïîðòà ôîòîàññèìèëÿòîâ â ðàñòåíèè â ñâîþ ñòîðîíó. Îáñóæäåí âîïðîñ îá ó÷àñòèè ôèòîãîðìîíîâ â ðåãóëÿöèè àêöåïòîðíîé àêòèâíîñòè îðãàíîâ è òêàíåé. Ðàññìîòðåíû òðè ãðóïïû ôàêòîâ: 1). Ïîëîæèòåëüíàÿ êîððåëÿöèÿ ìåæäó èíòåíñèâíîñòüþ ïðèòîêà àññèìèëÿòîâ ê îðãàíàì è òêàíÿì è ñîäåðæàíèåì ýíäîãåííûõ ôèòîãîð-ìîíîâ â íèõ. 2). Èçìåíåíèå ïîä äåéñòâèåì ýêçîãåííûõ ôèòîãîðôèòîãîð-ìîíîâ íàïðàâëåíèÿ òðàíñïîðòà è ðàñïðåäåëåíèÿ àññèìèëÿòîâ, óñèëåíèå ïðèòîêà ïîäâèæíûõ ìåòàáîëèòîâ ê îáðàáîòàííûì çîíàì ðàñòåíèÿ. 3). Ñâÿçü ìåæäó èçìåíåíèåì ðàñïðåäåëåíèÿ ïðîäóêòîâ ôîòîñèíòåçà ïî ðàñòåíèþ è ñîäåðæàíèåì ôèòîãîðìîíîâ â îðãàíàõ â îòâåò íà èçìåíåíèå ñëîæèâøèõñÿ â îíòîãåíåçå äîíîðíî-àêöåïòîðíûõ îòíîøåíèé (íàïðèìåð, ïðè óäàëåíèè ÷àñòè äîíîðíûõ èëè àêöåïòîðíûõ îðãàíîâ, èçìåíåíèè óñëîâèé îêðóæàþ-ùåé ñðåäû – ñâåòà, òåìïåðàòóðû, ôîòîïåðèîäà, êîíöåíòðàöèè ÑÎ2 è ò.ä.). Èçëîæåíà ñèñòåìà âçãëÿäîâ, â êîòîðîé îáðàçîâàíèå, òðàíñïîðò, ðàñïðåäåëåíèå, ïîòðåáëåíèå è çàïàñàíèå âåùåñòâ èíòåãðèðîâàíû â åäèíóþ ñèñòåìó âçàèìîñâÿçàííûõ ïðîöåññîâ. Âûñêàçàíî ïðåäïîëîæåíèå, ÷òî âçàèìîñâÿçü ìåæäó íèìè îñóùåñòâëÿåòñÿ áëàãîäàðÿ ôîðìèðîâàíèþ ãðàäèåíòà îñìîòè÷åñêîãî ïîòåíöèàëà è äàâëåíèÿ ìåæäó äîíîðíûì è àêöåïòîðíûì êîíöàìè òðàíñïîðòíîãî ïóòè. Âåëè÷èíà îñìîòè÷åñêîãî
ïîòåíöèàëà íà äîíîðíîì êîíöå ôëîýìû îïðåäåëÿåòñÿ íàëè÷èåì òðàíñïîðòíîãî ïóëà àññèìèëÿòîâ, ðàçìåð êîòîðîãî çàâèñèò îò óãëåâîäíîé èëè íåóãëåâîäíîé íàïðàâëåííîñòè ïîñòôîòîñèíòåòè÷åñêîãî ìåòàáîëèçìà, ïîòðåáëåíèÿ óãëåðîäà â ñàìîì ëèñòå, îòëîæåíèÿ â êëåòêàõ ìåçîôèëëà â âèäå çàïàñíûõ ñîåäèíåíèé èëè âêëþ÷åíèåì â òðàíñïîðòíûå ôîðìû àññèìèëÿòîâ. Âûñêàçàíî ïðåäïîëîæåíèå î òîì, ÷òî âåëè÷èíà îñìîòè-÷åñêîãî ïîòåíöèàëà â êîíöå òðàíñïîðòíîãî ïóòè ïîâûøàåòñÿ áëàãîäàðÿ ðîñòó ðàñòÿæåíèåì è íàêîïëåíèþ ïîëèìåðíûõ è/èëè ãèäðîôîáíûõ ñîåäèíåíèé, êîòîðûå îïðåäåëÿþò àêöåïòîðíûå ñâîéñòâà îðãàíîâ è òêàíåé ðàñòåíèÿ. Ñäåëàí âûâîä î òîì, ÷òî öèòîêèíèíû àêòèâèðóþò ñïåöèôè-÷åñêóþ, ãåíåòè÷åñêè äåòåðìèíèðîâàííóþ ôóíêöèîíàëüíóþ àêòèâíîñòü äîíîðíûõ è àêöåïòîðíûõ òêàíåé è îðãàíîâ ðàñòåíèÿ, óâåëè÷èâàÿ ãðàäèåíò îñìîòè÷åñêîãî ïîòåíöèàëà ìåæäó íèìè è ãðàäèåíò äàâëåíèÿ âäîëü òðàíñïîðòíîãî ïóòè, âëèÿÿ, òåì ñàìûì, íà èíòåíñèâíîñòü è íàïðàâ-ëåííîñòü ôëîýìíîãî òðàíñïîðòà âåùåñòâ.
ÝÊÎËÎÃÈ×ÅÑÊÈÅ ÏÐÎÁËÅÌÛ ÓÐÀËÜÑÊÎÃÎ ÐÅÃÈÎÍÀ Øàâíèí Ñ.À., Âëàñåíêî Â.Ý. Áîòàíè÷åñêèé ñàä ÓðÎ ÐÀÍ, ã. Åêàòåðèíáóðã, Ðîññèÿ, [email protected] Ïðèðîäíûå óñëîâèÿ Óðàëà õàðàêòåðèçóþòñÿ áîëüøèì ðàçíîîáðàçèåì ëàíäøàôòîâ, íà åãî òåððèòîðèè ðàñïîëàãàþòñÿ áîëüøèå ìàññèâû ëåñîâ (áîëåå 50% ïëîùàäè, èëè 31,8 ìëí. ãà ëåñîïîêðûòîé ïëîùàäè), ñòåïåé, åñòü òóíäðà, ëåñîòóíäðà, âîäîåìû, ãîðíûå òóíäðû. Çíà÷èòåëüíàÿ ÷àñòü òåððèòîðèè ïîäâåðæåíà ñèëüíîìó òåõíîãåííîìó è àíòðîïîãåííîìó âëèÿíèþ. Ñîòíè òûñÿ÷ ãåêòàð ëåñà ñòðàäàþò îò ãàçîâûõ âûáðîñîâ ïðåäïðèÿòèé. Âîäîåìû, îñîáåííî íà Ñðåäíåì è Þæíîì Óðàëå, îòëè÷àþòñÿ âûñîêèì çàãðÿçíåíèåì. Íà Óðàëå áîëåå 150 âèäîâ ðàñòåíèé îòíîñÿòñÿ ê ÷èñëó èñ÷åçàþùèõ. Ïî ýòîé ïðè÷èíå ïðîáëåìà ñîõðàíåíèÿ ïðèðîäíûõ îáúåêòîâ ÿâëÿåòñÿ èñêëþ÷èòåëüíî îñòðîé. Îíà ðåøàåòñÿ ïóòåì ñîçäàíèÿ ñèñòåìû îõðàíÿåìûõ òåððèòîðèé (ÎÎÏÒ). Îñîáîå ìåñòî çàíèìàåò Ñâåðäëîâñêàÿ îáëàñòü. Äàííûé ðåãèîí ïðåäñòàâëÿåò ñîáîé áîãàòóþ è ñàìîáûòíóþ âî ôëîðèñòè÷åñêîì îòíîøåíèè òåððèòîðèþ – çäåñü ïðîèçðàñòàåò îêîëî 1600 âèäîâ ñîñóäèñòûõ ðàñòåíèé.  Ñâåðäëîâñêîé îáëàñòè ðàñïîëàãàþòñÿ äâå áîòàíèêî-ãåîãðàôè÷åñêèå çîíû – òàåæíàÿ ñ òðåìÿ ïîäçîíàìè, ëåñîñòåïíàÿ è ïåðåõîäíàÿ ìåæäó íèìè ïîäçîíà ïðåäëåñîñòåïíûõ ñîñíîâî-áåðåçîâûõ ëåñîâ. Ñâåðäëîâñêàÿ îáëàñòü ÿâëÿåòñÿ îäíèì èç êðóïíåéøèõ ïðîìûøëåííûõ ðåãèîíîâ Ðîññèéñêîé Ôåäåðàöèè. Íà åå òåððèòîðèè ñóùåñòâóåò 2225 ïðåäïðèÿòèé è îðãàíèçàöèé ïî ñëåäóþùèì îòðàñëÿì: òîïëèâíî-ýíåðãå-òè÷åñêàÿ; ìåòàëëóðãè÷åñêàÿ; õèìè÷åñêàÿ; ëåñíàÿ; ìàøèíîñòðîèòåëüíàÿ; òðàíñïîðòíàÿ; ñåëüñêîõîçÿéñòâåííàÿ; æèëèùíî-êîììóíàëüíàÿ è äð. Âêëàä îñíîâíûõ îòðàñëåé ýêîíîìèêè â çàãðÿçíåíèå ïî÷â, à òàêæå âîçäóøíîãî è âîäíîãî áàññåéíîâ ñîñòàâëÿåò: 44,3% ñóììàðíîãî âûáðîñà – îò ïðåäïðèÿòèé ìåòàëëóðãè÷åñêîãî êîìïëåêñà, 32,8% – îò ïðåäïðèÿòèé òîïëèâíî-ýíåðãåòè÷åñêîãî êîìïëåêñà. Íàèáîëüøåå êîëè÷åñòâî äèîêñèäîâ ñåðû ïîñòóïàåò â àòìîñôåðó îò ïðåäïðèÿòèé ìåòàëëóðãèè (50,8%), òîïëèâíîé ýíåðãåòèêè (42,7%). Ïðåäïðèÿòèÿ ìåòàëëóðãèè äàþò íàèáîëüøèé âêëàä òàêæå â âûáðîñû îêñèäà óãëåðîäà (71,6%). Çàãðÿçíåíèå àòìîñôåðíîãî âîçäóõà îêñèäàìè àçîòà îïðåäåëÿåòñÿ, ãëàâíûì îáðàçîì, âûáðîñàìè ïðåäïðèÿòèé òîïëèâíî-ýíåðãåòè÷åñêîãî êîìïëåêñà (63,9%). Îñîáóþ ðîëü çàíèìàþò ëåñà è çåëåíûå íàñàæäåíèÿ ãîðîäîâ Ñâåðäëîâ-ñêîé îáëàñòè, êàê ñðåäîçàùèòíûå è ñðåäîôîðìèðóþùèå êîìïëåêñû. Îáùàÿ ïëîùàäü ëåñíîãî ôîíäà è ëåñîâ, íå âõîäÿùèõ â ëåñíîé ôîíä íàøåãî ðåãèîíà ñîñòàâëÿåò 16063,0 òûñ. ãà, ïðè ëåñèñòîñòè òåððèòîðèè îáëàñòè 67,5%. Èõ ðîëü òðóäíî ïåðåîöåíèòü. Âî-ïåðâûõ, îíè ÿâëÿþòñÿ
÷àñòüþ ïðèðîäíîãî êîìïëåêñà, ðåãóëèðóþùåãî îñíîâíûå ôèçè÷åñêèå ïðîöåññû, ïðîèñõîäÿùèå â àòìîñôåðå è â ïî÷âå, âîññîçäàþùèå åñòåñòâåííûå óñëîâèÿ ñðåäû è îïðåäåëÿþùèå ðàäèàöèîííûé, ãèäðîëî-ãè÷åñêèé ìèêðîêëèìàòè÷åñêèé, ãàçîâûé è ìèêðîáèîëîãèäðîëî-ãè÷åñêèé ðåæèìû ñðåäû. Äëÿ ðåøåíèÿ ýêîëîãè÷åñêèõ ïðîáëåì íåîáõîäèìî ñîçäàíèå äîëãîñðî÷íûõ è ñðåäíåñðî÷íûõ ïðîåêòîâ ïî îðãàíèçàöèè ýêîëîãè÷åñêîãî ìîíèòîðèíãà ñ öåëüþ ïðîãíîçà ñîñòîÿíèÿ è óñòîé÷èâîñòè ëåñíûõ è äðóãèõ ýêîñèñòåì. Òåððèòîðèÿ Ñâåðäëîâñêîé îáëàñòè ïðèíàäëåæèò áàññåéíàì 7-ìè îñíîâíûõ ðåê, ê êîòîðûì îòíîñÿòñÿ ðåêè Òàâäà, Òóðà, Ïûøìà, Èñåòü, ×óñîâàÿ, Óôà, Ñûëâà.  öåëîì ãèäðîëîãè÷åñêàÿ ñåòü îáëàñòè ïðåäñòàâëåíà 18414 ðåêàìè îáùåé ïðîòÿæåííîñòüþ áîëåå 68 òûñ. êì. Ïî äàííûì èíâåíòàðèçàöèè, â Ñâåðäëîâñêîé îáëàñòè ýêñïëóàòèðóåòñÿ 129 âîäîõðà-íèëèù îáúåìîì áîëåå 1 ìëí. ì3 âñå ýòè âîäíûå îáúåêòû èñïûòûâàþò âûñîêóþ àíòðîïîãåííóþ íàãðóçêó. Çà ïðîøåäøèé ãîä áûëî ñáðîøåíî 385 òûñ. ò çàãðÿçíÿþùèõ âåùåñòâ. Îñíîâíûå èíãðåäèåíòû â ýòèõ ñáðîñàõ ïðåäñòàâëåíû: íåôòåïðîäóêòàìè, ñóëüôàòàìè, íèòðàòàìè, æåëåçîì, êàëèåì, êàëüöèåì, ëèãíèíîì, ìàãíèåì, ìàðãàíöåì, ìûøüÿêîì, íàòðèåì, öèíêîì, ôåíîëîì è äð. Íàèáîëüøåå êîëè÷åñòâî çàãðÿçíåííûõ ñòî÷íûõ âîä ïîñòóïàåò â ïîâåðõíîñòíûå âîäíûå îáúåêòû îò: ïðåäïðèÿòèé æèëèùíî-êîììóíàëüíîãî õîçÿéñòâà – 54,3%; ÷åðíîé ìåòàëëóðãèè – 10,5%; õèìè÷åñêîé ïðîìûøëåííîñòè – 10,2%; öâåòíîé ìåòàëëóðãèè – 6,7%; ìàøèíîñòðîåíèÿ – 5,8%. Âñå ýòî îïðåäåëÿåò àêòóàëüíîñòü èçó÷åíèÿ è ðåøåíèÿ ïðîáëåì âîäíîé ðàñòèòåëüíîñòè. Çíà÷èòåëüíàÿ ÷èñëåííîñòü íàñåëåíèÿ Ñâåðäëîâñêîé îáëàñòè, âûñîêàÿ êîíöåíòðàöèÿ ïðåäïðèÿòèé ÷åðíîé è öâåòíîé ìåòàëëóðãèè, ýëåêòðîýíåð-ãåòèêè îïðåäåëÿþò íàëè÷èå áîëüøîãî êîëè÷åñòâà îòõîäîâ ïðîèçâîäñòâà è ïîòðåáëåíèÿ.  ñâÿçè ñ ýòèì áîëüøóþ âàæíîñòü èìååò ðàçðàáîòêà ìåòîäîâ ðåêóëüòèâàöèè è áëàãîóñòðîéñòâà òåððèòîðèé, íàðóøåííûõ äåéñòâèåì ïðîìûøëåííûõ ïðåäïðèÿòèé.  äàííîì íàïðàâëåíèè àêòèâíî ðàáîòàþò ëàáîðàòîðèè Áîòàíè÷åñêîãî ñàäà ÓðÎ ÐÀÍ è ÓðÃÓ. Èìè íàêîïëåí çíà÷èòåëüíûé íàó÷íûé îïûò â îáîñíîâàíèè íîâûõ ïîäõîäîâ ê ðåêóëüòèâàöèè çåìåëü. Íåìàëîâàæíîé ïðîáëåìîé äëÿ Ñâåðäëîâñêîé îáëàñòè ÿâëÿåòñÿ ðàöèîíàëüíîå èñïîëüçîâàíèå ëåñíûõ ðåñóðñîâ.  ñâÿçè ñ ââåäåíèåì â äåéñòâèå íîâîãî Ëåñíîãî êîäåêñà âîçíèêëà îñòðàÿ ïîòðåáíîñòü â ðàçðàáîòêå îáëàñòíîé ïðîãðàììû, íàïðàâëåííîé íà ðåøåíèå ðåãèî-íàëüíûõ çàäà÷. Íàðÿäó ñ ïåðå÷èñëåííûì, îñîáîå ìåñòî çàíèìàåò ïðîáëåìà èçó÷åíèÿ, îõðàíû è èñïîëüçîâàíèÿ íåäðåâåñíîé ðàñòèòåëüíîñòè., âêëþ÷àÿ ëåêàðñòâåííûå, ðåäêèå è èñ÷åçàþùèå âèäû.
ÒÅÇÈÑÛ
POTATO TRANSFORMATION WITH A YEAST-DERIVED INVERTASE GENE MODIFIES SSR AND CHILLING TOLERANCE
Deryabin A.N., Klimov S.V., Sinkevich M.S., Trunova T.I.
Timirayzev Institute of Plant Physiology, Russian Academy of Sciences, Moscow, Russia; [email protected]
In contrast to frost-resistant plants, the role of source-sink relations (SSR) and carbohydrate metabolism in the formation of resistance in chilling-tolerance plants is much less investigated. It is quite expedient to study this issue in a typical chilling-tolerance potato plant. The potato (Solanum tuberosum L., cv. Desiree) transformed with vector carrying yeast invertase gene under the control of tuber-specific patatin promoter B33 class I, fused with proteinase II inhibitor leader peptide to provide enzyme location in apoplast is an organism with modified SSR. The boosted expression of yeast invertase gene caused a retardation of sucrose efflux from the cells because in the apoplast it was transformed into nontranslocated hexoses. As a result, the efflux of assimilates from photosynthesizing tissues is suppressed, and they accumulate in the leaves. Common potato plants of the same cultivar (wild type) served as control plants. The plants were taken from the collection of clones produced as a result of cooperation between the researchers of Max Planck Institute of Molecular Plant Physiology (Golm, Germany) and Chailakhyan Laboratory of Growth and Development, Timiryazev Institute of Plant Physiology, Russian Academy of Sciences. The plants were micropropagated in vitro and grown in a
controlled-climate chamber at 22oC and 16-h illumination from the luminescent lamps
producing white light (illuminance of 4 klx) for 5 weeks in test-tube culture on MS-medium containing 2% sucrose.
At a temperature of 22oC optimal for growth, the transformed plants differed
from the plants of wild type in retarded growth and a lower rate of photosynthesis as calculated per plant. On a leaf dry weight basis, photosynthesis of transformed
plants was higher than in control plants. Under hypothermia (5oC), dark
respiration and especially photosynthesis of transformed plants turned out to be more intense than in control plants. These changes modify the balance between photosynthesis and retarded efflux of assimilates, causing an increase in the intracellular level of sugars and a rise in the resistance to chilling. The exposure of plants to light under prolonged hypothermia (5oC, 6 days) activated all the
forms of invertase (predominantly, of acid invertase) and induced accumulation of sugars. In the leaves of potato expressing the gene for yeast invertase, these processes were more intense. Under chilling, the rate of lipid peroxidation and superoxide dismutase activity in the leaves of investigated genotypes of potato depended on the level of accumulated intracellular sugars. It was concluded that
changed SSR under hypothermia stimulate of sugar accumulation in leaves. Those sugars play an important role as stabilizers of cellular membranes upon the development of oxidative stress induced by hypothermia.
This work was supported by the Russian Foundation for Basic Research, project no. ¹ 07-04-00601.
MOLECULAR ASPECTS OF REGULATION OF THE PHOTOSYNTHETIC ANTENNA FUNCTION
Gruszecki W.I.1, Grudzinski W.1, Janik E.2, Gospodarek M.3
1Department of Biophysics, Institute of Physics, Maria Curie-Sklodowska
University, Lublin, Poland, [email protected]
2Department of Plant Physiology, Institute of Biology, Maria Curie-Sklodowska
University, Lublin, Poland
3Institute of Physics, Technical University, Lublin, Poland
Optimization of the photosynthetic activity of plants requires efficient regulation of both the antenna function and photoprotection against photoinhibition. Absorption of light quanta by accessory pigments bound to the antenna pigment-protein complexes creates singlet excitations utilized to drive photochemical reactions but can be also converted to triplet excitations which may initiate oxidative damage of the photosynthetic apparatus. Therefore, regulation of the photosynthetic antenna function seems to be an important aspect of overall regulatory processes operating in the photosynthetic apparatus. Our research was focused on regulatory processes operating at the molecular level, in the pigment-protein antenna complex of Photosystem II. Chlorophyll
a fluorescence excitation spectroscopy show that illumination of the complex is
associated with energetic uncoupling of a certain pool of accessory xanthophyll pigments from chlorophylls. Analysis of the resonance Raman scattering spectra of LHCII-bound xanthophyll pigments reveals that the energetic uncoupling observed is accompanied by liberation of a small fraction of xanthophyll pigments (identified as violaxanthin) from the protein bed. Such a transition can be discussed in terms of down-regulation of the photosynthetic antenna function under overexcitation conditions. On the other hand, this process can be also discussed in terms of the process of making violaxanthin available for enzymatic de-epoxidation within the xanthophyll cycle. Light-dependent reorganization of the pool of violaxanthin has also pronounced effect on organization of LHCII, as deduced from the analysis of FTIR spectra.
The model of regulation of the photosynthetic antenna function at the level of pigment-protein complexes, based on the results of recent studies, will be discussed.
INTERACTIVE FUNCTIONS OF Ni, Zn, Cu, Cd, Mn ON PHOTOSYN-THETIC PIGMENTS AND CHLOROPHYLL FLUORESCENCE IN Elodea
canadensis LEAVES
Maleva M.G.1, Malec P.2, Prasad M.N.V.3, Strzalka K.2
1A.M. Gorky Ural State University, Ekaterinburg, Russia, [email protected]
2Department of Plant Physiology and Biochemistry, Faculty of Biochemistry
Biophysics and Biotechnology, Jagiellonian University, Krakow, Poland; [email protected], [email protected]
3Department of Plant Sciences, University of Hyderabad, Hyderabad, India,
The pulse-amplitude modulation (PAM) technique has been applied to study effects of selected heavy metals on photosynthetic apparatus of Elodea sp. The
Elodea canadensis Michx. shoots were incubated for four days under the
standarized conditions (at temperature about 23°C and photon flux density of 60
ìM m-2s-1) in 5% modified Hoagland medium. Ni, Zn, Cu, Cd, Mn (as sulfate
salts) and their combinations were added in concentrations of 0.1 mg/l (μM values corresponding to their Maximum Acceptable Concentrations; MAC). The content of photosynthetic pigments (chl a, chl b, total carotenoids) in Elodea leaves was measured spectrophotometrically according to Lichtenthaler (1987) and normalized per gram of dry weight of material. Chlorophyll fluorescence parameters were measured by PAM 210 fluorometer (Walz, Effeltrich, Gemany). All heavy metals at used concentrations induced changes on the chlorophyll and carotenoid contents in E. canadensis leaves. The relative content of chl a decreased in leaves treated with Cu and Mn (12-16%), and also in leaves treated with compositions of Ni+Cu (13%) and Mn+Cu (24%). The chl b content showed no significant changes. Copper and a composition of Cu+Mn significantly reduced the accumulation of carotenoids (34%). On the contrary, cadmium stimulated the accumulation of chlorophylls and carotenoids in
E. canadensis leaves (hormesis). Also, the tendency to stimulate the pigment
accumulation was observed for simultaneous treatments: Ni+Zn and Cd+Mn. The photosynthetic efficiency of PS II (Fv/Fm), as measured in dark-adapted leaves treated with heavy metals, expressed no significant changes in comparison to the control material (0.787±0.01) with except of leaves treated with Cu (0.624±0.07). Interestingly, the presence of other ions (Ni or Mn) reduced the inhibitory effect of Cu on Fv/Fm value.
Electron transport rate (ETR) in the control material showed a maximum (23.8±2.6) at photon flux density of 314 ìÌ m-2s-1 ETR was strongly enhanced
by Ni and Zn (27.6±0.6 and 25.4±2.7 respectively). This effect was accompanied with a shift of the maximum of photon flux density to 444 ìÌ m-2s-1. Contrary,
in leaves treated with copper and cadmium the maximal values of ETR were observed at 214 ìÌ m-2s-1 (11.4±1.3 and 18.3±1.7 respectively). The presence
of Ni or Mn in the medium abolished the effect of Cu and Cd on ETR in leaves. Additionally, Cd, Mn and especially Cu, significantly reduced both photochemical quenching (qP) and nonphotochemical quenching (qN) of chlorophyll fluorescence (the reduction level of 35-74% at photon flux density
214-444 ìÌ m-2s-1). This effect was not observed if leaves were treated
simultaneously with more than one ion.
In conclusion, our results indicate that heavy metals as Cd, Cu, and Mn, applied at concentrations corresponding to their MAC could induce a remarkable inhibitory effect on the functioning of the photosynthetic apparatus (mainly photosystem II) in Elodea canadensis leaves. The observed hormetic effect of Cd on pigment accumulation was not accompanied with the stimulation of photosynthetic electron transport around PS II. Also, the presence of additional ions in the medium significantly reduced toxic effects of single metal ions on pigment accumulation and/or chlorophyll fluorescence parameters in
METAL-BIOMOLECULES COMPLEXES IN PLANTS: A CRUCIAL COMPONENT OF GREEN TECHNOLOGY TO CONTAIN AND CLEAN UP HEAVY METALS IN THE ENVIRONMENT
M.N.V. Prasad
Department of Plant Sciences. University of Hyderabad, Hyderabad 500046, India, [email protected], [email protected]
Potentially toxic metals are available to plants through: mining activities – smelting, river dredging, mine spoils and tailings, metal industries etc.; industries – plastics, textiles, microelectronics, wood preservatives, refineries etc.); atmospheric deposition – urban refuse disposal, pyrometallurgical industries, automobile exhausts, fossil fuel combustion etc.; excessive use of agrochemicals – fertilizers and pesticides and waste disposal -sewage sludge.
Self-cleaning of soils does not takes place or rather extremely slow. The toxic metals in top soil, thus get accumulated in plants. Plants can remediate metal pollutants in many ways as detailed below:
1. Phytoremediation: Use of plants to remediate contaminated soil or water 2. Phytoaccumulation: The uptake and concentration of contaminants (metals or organics) within the roots or aboveground portion of plants.
3. Phytoextraction: the use of plants at waste sites to accumulate metals into the harvestable, above ground portion of the plant and, thus, to decontaminate soils
4. Phytomining: Use of plants to extract inorganic substances from mine ore 5. Phytostabilisation: plants tolerant to the element in question are used to reduce the mobility of elements, thus, they are stabilised in the substrate or roots 6. Phytovolatilization: The uptake and transpiration of a contaminant by a plant, with release of the contaminant or a modified form of the contaminant to theatmosphere from the plant.
7. Rhizofiltration/Phytofiltration: Roots or whole plants of element accumulating plants absorb the element from polluted effluents and are later harvested to diminish the metals in the effluents.
Fields of economic and social interests of metal biomolecules and in the broad area of phytotechnology are:
a) Ecotoxicology and ecophysiology: Nutrient availability efficiency and deficiency for optimization of crop yield
b) Environmental chemistry waste management and mine reclamation: Bioremediation and restoration of metal contaminated and polluted ecosystems
c) Agricultural and nutritional sciences, food industry: Plant and Soil serve as vital links for supplementing nutrients on sustainable basis
d) Clinical biochemistry: Usage radionuclides and their disposal
e) Medicine and pharmacology: Usage of plants as a source of nutrients for certain disorders
The involved principal metal binding ligands, enzymes, proteins and genes and their application in green technologies for containment and cleanup of heavy metals in the environment are presented in this lecture.
THE INVESTIGATION OF PHYTOREMEDIATION POTENTIAL OF
Calamagrostis epigeios L. AND Brassica napus L. ARE PROMISING PLANT
SPECIES FOR THE OF Ni-CONTAMINATED SOILS IN RUSSIA N.I. Shevyakova, Yu.G. Madzhugina, E.N. Iljina, Vl.V. Kuznetsov
Timiryazev Institute of Plant Physiology, Russian Academy of Sciences, Botanicheskaya str. 35, 127276 Moscow, e-mail: nshevyakova@yandex. ru
The goal of these studies was to identify plant species with strong ability to take up Ni from contaminated soils in humidity zone of Russia. These investigations was based on assumption that such plant species may be found among which are ruderal plants growing naturally in contaminated areas or of cultured species belong to the family of Brassicaceae a majority of wild plants accumulating Ni. As well known, hyperaccumulators are defined as plants that can accumulate 1000 ìg/g Ni and up. We selected two plant species for detailed studies: ruderal cereal Calamagrostis epigeos and cultured species Brassica napus which were capable of high leaf biomass produced under Ni- or other heavy metals-contaminated soils. These plants show the tolerance to Ni during seed germination. Leaf of C. epigeos can accumulate Ni close to 700 ìg/g dry weight when they grown on heavy metal-contaminated soil-ground and their shoots were tolerant to cut that was of importance in Ni removal in process of soil cleaning. Leaf of 5-week age plants B. nutans grown in water culture in the
presence of NiCl2 (500 ìM) accumulated Ni (640 ìg/g dry mass). After the
treatment of leaf surface of plants grown in presence NiCl2 by exogenous
polyamine (putrescine, 1mM) as helator Ni content in leaf was increased 2-3 times greater.
This work was partially supported by the Russian Foundation for Basic Research (pr. no. 07-04-00241 and 07-04-00995).
EXOGENOUS CADAVERINE INDUCES OXIDATIVE BURST AND REDUCES CADAVERINE CONJUGATE CONTENT IN THE COMMON ICE PLANT
N. I. Shevyakova, L. A. Stetsenko, Vl.V. Kuznetsov
Moscow, Timiryazev Institute of Plant Physiology, Russian Academy of Sciences, Botanicheskaya str. 35, [email protected]
The effect of free cadaverine (Cad) on its conjugates formation was analyzed in roots of the common ice plants (Mesembryantemum crystallinum L.). It was found for the first time that Cad could induce oxidative burst in the roots of adult plants, as was evident from the sharp decrease in content of Cad soluble or insoluble congugates. This unusual effect was associated with the increased
oxidative degradation of exogenous Cad (1 mM, 1.5 h) and intense H2O2
production in the root cells of adult plants. Root treatment of both juvenile and
adult plants with H2O2 (1 mM, 1.5 h) reduced the content of soluble Cad
conjugates and increased the content of their components, free Cad and phenols. We also found that one of possible reasons of negative effect of exogenous diamine on the formation of conjugated forms in adult roots was alkalization of the root
apoplast at Cad addition to nutrient medium and the unusual O2•- synthase
function as a pH-dependent guaiacol peroxidase in the presence of a high content of H2O2. This was confirmed by the data on the accumulation of O2•- and
enhanced superoxide dismutase activity in adult roots under treatment with Cad.
It is possible, that the accumulation of O2•- together with H
2O2 was also
responsible for oxidative burst, which induced a decrease in the content of Cad conjugates in adult roots of the common ice plants.
This work was partially supported by the Russian Foundation for Basic Research (project no. 07-04-00241) and by the Program of the Presidium of RAS (Molecular and Cell Biology).
CARBON 13C ISOTOPE DISCRIMINATION WITH THE C
3 AND C4
PHOTOSYNTHESIS UNDER SOIL SALINITY STRESS
Shuyskaya E.V.1, Matsuo N.2, Toderich K.N.3, Sunada K.4, Gismatullina L3,
Radjabov T.3, Ivanova L.A.5, Ronjina D.A.5, Ivanov L.A.5, Voronin P.Yu.1, Black C.C.6
1K.A. Timiriazev Plant Physiology Institute, Rus. Acad. Sci., Moscow, Russia,
2Mie University, Mie, Japan
3Samarkand Branch of Acad. of Sci. of Uzbekistan, Samarkand, Republic of
Uzbekistan
4Interdisciplinary Graduate School of Medicine and Engineering University
of Yamanashi, Takeda Kofu, Japan
5Botany Garden, Ural Branch of Rus. Acad. Sci., Ekaterinburg, Russia
6The University of Georgia, Athens, USA
It is well known the carbon 13C isotope discrimination (CID) reflects a
photosynthetic productivity and a plant water use efficiency (WUE) (Farquhar et al., 1989). The C4 plant CID has ever revealed δ13C range valued between -10
and -17‰ is twice as low compare with the C3 ones (-25 —: -35‰) (Edwards et al., 1986). This persuades into a more effective water and CO2 gas exchange of the C4 plants compare with the C3 ones under non-light limiting conditions of Turan deserts at a hot and dry summer season at Middle Asia plain. Thus one would have cast a proposal of the strongest C4 plants’ salt tolerance in native
habitats of the Turan deserts. We had compared δ13C of some C
4 (Haloxylon
aphillum, Salsola paulsenii) and C3 (Tamarix hispida, Alhagi pseudalhagi) plants. Plant material had been gathered at places with gradual soil salinity measured with natural Na+, Mg2+, Ca2+, HCO
3-, SO42-, Cl- soil contents during two
vegetation seasons had been contrast on rainfall precipitation (0 and 26 mm). Carbon 13C isotope’s fractionation measured as δ13C had been determined with
mass-spectrometer Delta-S (Thermo Finnigan, USA). One can evaluate carbon
13C isotope discrimination taking in mind the δ13C increase really does mean the
CID decrease. The C3 plant probing from higher soil salinity places had revealed the stronger suppression of the CID (1‰) relative to proper C4 variants (0.1‰). From our data received the overall range of δ13C for C
4 was twice as high as C3
one according with current notion about. The CID C3 species specificity was
available. For the first, δ13C mean value of A. pseudalhagi was 1.5‰ lower than
one of T. hispida and δ13C increase (3.3‰) of A. pseudalhagi under the strongest
salinity was twice as strong as δ13C increase (1.6‰) under proper salt stress. For
the second, the C3 species specificity had been determined toward a Ca2+
WUE lower then T. hispida one. The higher soil salinity makes a water osmotic potential difference between soil and leaves decrease which in turn suppresses a transpiration. Under this reason T. hispida soil salinity stress tolerance is higher then one of A. pseudalhagi.
For the aridity season dependence of interest, the CID effect was opposed to salt stress one. The C3 plant (T. hispida) decreased its δ13C value from August
(-25.39‰) to November (-26.76‰). Meanwhile for the C4 plant (H. aphillum)
its δ13C November value (-11.75‰) excelled the August (-12.29‰) one.
Higher soil salinity didn’t change the mesophyll cell sizes of the C4 plant (H. aphillum), but it significantly did enlarge the cuticula depth. The salt stress increase made C3 plant (A. pseudalhagi) leaves change their leaf tissue complexity with a differ manner: cuticula didn’t changed, but mesophyll cells became much
small in size. These data argue the salt stress dependence of the C3 plant
transpiration is stronger then C4 one.
The results considered characterize the C3 and C4 photosynthetic gas exchange environment stress response specificity of the CID. Some authors (Tiezen & Boutton, 1989; Wang et al., 2006) demonstrated a light-limiting dependence of C4 plants at certain vegetation periods. Meanwhile a transpiration has been being depended of the C3 plant water supply limits the C3 photosynthetic production
at a hot and dry summer season. Under such a condition the highest C3 plant
production had been pointed out at the seasons with the highest annual rainfall precipitation (Ehleringer et al., 1988; Farquhar et al., 1989; Wang et al., 2003). At a case of the consideration, November was followed by the winter rainfall which would be the reason for the partial dilution of the soil salinity. This could have to diminish the salt stress for transpiration-limited C3 plants in extent much
higher compared with light-limited C4 plants allowing surpass of the C3
productivity v. C4 one.
One could conclude the soil salinity makes the CID reveal an explicit quantitative attribute for proper suppression of the C3 and C4 productivity at native arid plain Middle Asia Turan deserts.
