干旱林线区不同树种非结构性碳水化合物的季节格局及其主导因子
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摘要
为探究干旱林线区不同树种非结构性碳水化合物(NSC)的季节格局以及生物因素(树种)和非生物因素(气候)在其中的作用,选取内蒙古高原东南缘4种典型乔木(樟子松、白桦、小叶杨和华北落叶松)作为研究对象,对其生长季叶片、树干中的可溶性糖、淀粉及总非结构性碳水化合物的浓度进行比较。结果表明:不同功能型树种NSC各组分浓度具有相似的季节格局(生长季初期短暂上升,生长季旺期持续下降,生长季末期由于生长减缓而逐渐回升),说明气候因素在NSC季节动态中起主导作用。不同树种NSC季节均值总体上具有显著差异(p<0.05),白桦NSC各组分含量均高于樟子松,体现生物因素的影响。更加干旱的局地气候条件使得樟子松NSC含量高于小叶杨及华北落叶松,说明干旱林线NSC含量影响因子的复杂性。叶片可溶性糖及TNC含量高于树干,而淀粉含量低于树干,这与植物不同器官的功能差异有关。
To examine the seasonal patterns of non-structural carbohydrates(NSC) in different tree species and the roles of biotic(tree species) and abiotic(climate) factors on these patterns at xeric timberline, the authors monitored seasonal patterns of non-structural carbohydrates in four different tree species(Pinus sylvestris var.mongolica, Betula platyphylla, Populus simonii and Larix principis-rupprechtii) in southeastern Inner Mongolian Plateau of China. The results show that tree species with different foliar habit types have broadly similar seasonal non-structural carbohydrates(NSC) patterns(NSC concentration increased at the beginning of the growing season,and reduced in summer because of high growth rate, then increased slowly in the end), which suggests a climatic determination. Significant differences(p<0.05) in NSC concentrations are observed between four tree species.NSC concentration in birch is significant higher than that in pine, reproducing the pattern that deciduous species have higher NSC pool than ever green ones, which suggests a biotic determination. Due to the relatively drier climates, NSC concentration in larch and poplar turn to be lower than that in pine, implying a combined effect of both biotic and abiotic factors. The sugar and TNC(total non-structural carbohydrates) concentrations in leaf are significantly higher than those in stem, but the starch concentration in leaf is lower, which is consistent with the different roles of leaf and stem.
To examine the seasonal patterns of non-structural carbohydrates(NSC) in different tree species and the roles of biotic(tree species) and abiotic(climate) factors on these patterns at xeric timberline, the authors monitored seasonal patterns of non-structural carbohydrates in four different tree species(Pinus sylvestris var.mongolica, Betula platyphylla, Populus simonii and Larix principis-rupprechtii) in southeastern Inner Mongolian Plateau of China. The results show that tree species with different foliar habit types have broadly similar seasonal non-structural carbohydrates(NSC) patterns(NSC concentration increased at the beginning of the growing season,and reduced in summer because of high growth rate, then increased slowly in the end), which suggests a climatic determination. Significant differences(p<0.05) in NSC concentrations are observed between four tree species.NSC concentration in birch is significant higher than that in pine, reproducing the pattern that deciduous species have higher NSC pool than ever green ones, which suggests a biotic determination. Due to the relatively drier climates, NSC concentration in larch and poplar turn to be lower than that in pine, implying a combined effect of both biotic and abiotic factors. The sugar and TNC(total non-structural carbohydrates) concentrations in leaf are significantly higher than those in stem, but the starch concentration in leaf is lower, which is consistent with the different roles of leaf and stem.
引文
[1]Kozlowski T T.Carbohydrate sources and sinks in woody-plants.Botanical Review,1992,58(2):107-222
[2]Barbaroux C,Breda N,Dufrene E.Distribution of above-ground and below-ground carbohydrate reserves in adult trees of two contrasting broad-leaved species(Quercus petraea and Fagus sylvatica).New Phytologist,2003,157(3):605-615
[3]Bansal S,Germino M J.Carbon balance of conifer seedlings at timberline:relative changes in uptake,storage,and utilization.Oecologia,2008,158(2):217-227
[4]Martinez-Vilalta J,Sala A,Asensio D,et al.Dynamics of non-structural carbohydrates in terrestrial plants:a global synthesis.Ecological Monographs,2016,86(4):495-516
[5]Allen C D,Macalady A K,Chenchouni H,et al.Aglobal overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests.Forest Ecology and Management,2010,259(4):660-684
[6]Hartmann H.Will a 385 million year-struggle for light become a struggle for water and for carbon?-How trees may cope with more frequent climate change-type drought events.Global Change Biology,2011,17(1):642-655
[7]Mcdowell N,Pockman W T,Allen C D,et al.Mechanisms of plant survival and mortality during drought:why do some plants survive while others succumb to drought?.New Phytologist,2008,178(4):719-739
[8]Hoch G,Richter A,Korner C.Non-structural carbon compounds in temperate forest trees.Plant Cell and Environment,2003,26(7):1067-1081
[9]Chapin F S,Schulze E D,Mooney H A.The ecology and economics of storage in plants.Annual Review of Ecology and Systematics,1990,21:423-447
[10]Hoch G.Cell wall hemicelluloses as mobile carbon stores in non-reproductive plant tissues.Functional Ecology,2007,21(5):823-834
[11]Bonicel A,Haddad G,Gagnaire J.Seasonal-variations of starch and major soluble sugars in the different organs of young poplars.Plant Physiology and Biochemistry,1987,25(4):451-459
[12]Sauter J J,Wellenkamp S.Seasonal changes in content of starch,protein and sugars in the twig wood of Salix caprea L.Holzforschung,1998,52(3):255-262
[13]Piper F I,Fajardo A.Carbon dynamics of Acer pseudoplatanus seedlings under drought and complete darkness.Tree Physiology,2016,36(11):1400-1408
[14]Dietze M C,Sala A,Carbone M S,et al.Nonstructural Carbon in Woody Plants.Annual Review of Plant Biology,2014,65:667-687
[15]Liu H,Park Williams A,Allen C D,et al.Rapid warming accelerates tree growth decline in semi-arid forests of Inner Asia.Global Change Biology,2013,19(8):2500-2510
[16]印婧婧,郭大立,何思源,等.内蒙古半干旱区树木非结构性碳、氮、磷的分配格局.北京大学学报(自然科学版),2009,45(3):519-527
[17]Li H,Hoch G,K?rner C.Source/sink removal affects mobile carbohydrates in Pinus cembra at the Swiss treeline.Trees,2002,16(4):331-337
[18]Buysse J,Merckx R.An improved colorimetric method to quantify sugar content of plant-tissue.Journal of Experimental Botany,1993,44(10):1627-1629
[19]Barbaroux C,Breda N.Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuseporous beech trees.Tree Physiology,2002,22(17):1201-1210
[20]Richardson A D,Carbone M S,Keenan T F,et al.Seasonal dynamics and age of stemwood nonstructural carbohydrates in temperate forest trees.New Phytol,2013,197(3):850-861
[21]Dickson R E.Carbon and nitrogen allocation in trees.Annales Des Sciences Forestieres,1989,46:S631-S647
[22]Landhausser S M,Lieffers V J.Seasonal changes in carbohydrate reserves in mature northern Populus tremuloides clones.Trees-Structure and Function,2003,17(6):471-476
[23]Breda N,Huc R,Granier A,et al.Temperate forest trees and stands under severe drought:a review of ecophysiological responses,adaptation processes and long-term consequences.Annals of Forest Science,2006,63(6):625-644
[24]Hartmann H,Ziegler W,Kolle O,et al.Thirst beats hunger-declining hydration during drought prevents carbon starvation in Norway spruce saplings.New Phytologist,2013,200(2):340-349
[25]Dickman L T,Mcdowell N G,Sevanto S,et al.Carbohydrate dynamics and mortality in a pinonjuniper woodland under three future precipitation scenarios.Plant Cell And Environment,2015,38(4):729-739
[26]潘庆民,韩兴国,白永飞,等.植物非结构性贮藏碳水化合物的生理生态学研究进展.植物学通报,2002,19(1):30-38
[27]Gaudet D A,Laroche A,Yoshida M.Low temperature-wheat-fungal interactions:a carbohydrate connection.Physiologia Plantarum,1999,106(4):437-444
[28]Kozlowski T T,Pallardy S G.Acclimation and adaptive responses of woody plants to environmental stresses.Botanical Review,2002,68(2):270-334
[29]Fischer C,Holl W.Food reserves of scots pine(Pinus-sylvestris L.).1.seasonal changes in the carbohydrate and fat reserves of pine needles.TreesStructure and Function,1991,5(4):187-195
[30]Piispanen R,Saranp??P.Variation of non-structural carbohydrates in silver birch(Betula pendula Roth)wood.Trees,2001,15(7):444-451
[31]Fajardo A,Piper F I,Pfund L,et al.Variation of mobile carbon reserves in trees at the alpine treeline ecotone is under environmental control.New Phytologist,2012,195(4):794-802
[32]Sala A,Woodruff D R,Meinzer F C.Carbon dynamics in trees:feast or famine?.Tree Physiol,2012,32(6):764-775
[33]Fajardo A,Piper F I,Hoch G.Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients.Annuals of Botany,2013,112(3):623-631
[34]Sala A,Mencuccini M.Ecosystem science:plump trees win under drought.Nature Climate Change,2014,4(8):666-667
[2]Barbaroux C,Breda N,Dufrene E.Distribution of above-ground and below-ground carbohydrate reserves in adult trees of two contrasting broad-leaved species(Quercus petraea and Fagus sylvatica).New Phytologist,2003,157(3):605-615
[3]Bansal S,Germino M J.Carbon balance of conifer seedlings at timberline:relative changes in uptake,storage,and utilization.Oecologia,2008,158(2):217-227
[4]Martinez-Vilalta J,Sala A,Asensio D,et al.Dynamics of non-structural carbohydrates in terrestrial plants:a global synthesis.Ecological Monographs,2016,86(4):495-516
[5]Allen C D,Macalady A K,Chenchouni H,et al.Aglobal overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests.Forest Ecology and Management,2010,259(4):660-684
[6]Hartmann H.Will a 385 million year-struggle for light become a struggle for water and for carbon?-How trees may cope with more frequent climate change-type drought events.Global Change Biology,2011,17(1):642-655
[7]Mcdowell N,Pockman W T,Allen C D,et al.Mechanisms of plant survival and mortality during drought:why do some plants survive while others succumb to drought?.New Phytologist,2008,178(4):719-739
[8]Hoch G,Richter A,Korner C.Non-structural carbon compounds in temperate forest trees.Plant Cell and Environment,2003,26(7):1067-1081
[9]Chapin F S,Schulze E D,Mooney H A.The ecology and economics of storage in plants.Annual Review of Ecology and Systematics,1990,21:423-447
[10]Hoch G.Cell wall hemicelluloses as mobile carbon stores in non-reproductive plant tissues.Functional Ecology,2007,21(5):823-834
[11]Bonicel A,Haddad G,Gagnaire J.Seasonal-variations of starch and major soluble sugars in the different organs of young poplars.Plant Physiology and Biochemistry,1987,25(4):451-459
[12]Sauter J J,Wellenkamp S.Seasonal changes in content of starch,protein and sugars in the twig wood of Salix caprea L.Holzforschung,1998,52(3):255-262
[13]Piper F I,Fajardo A.Carbon dynamics of Acer pseudoplatanus seedlings under drought and complete darkness.Tree Physiology,2016,36(11):1400-1408
[14]Dietze M C,Sala A,Carbone M S,et al.Nonstructural Carbon in Woody Plants.Annual Review of Plant Biology,2014,65:667-687
[15]Liu H,Park Williams A,Allen C D,et al.Rapid warming accelerates tree growth decline in semi-arid forests of Inner Asia.Global Change Biology,2013,19(8):2500-2510
[16]印婧婧,郭大立,何思源,等.内蒙古半干旱区树木非结构性碳、氮、磷的分配格局.北京大学学报(自然科学版),2009,45(3):519-527
[17]Li H,Hoch G,K?rner C.Source/sink removal affects mobile carbohydrates in Pinus cembra at the Swiss treeline.Trees,2002,16(4):331-337
[18]Buysse J,Merckx R.An improved colorimetric method to quantify sugar content of plant-tissue.Journal of Experimental Botany,1993,44(10):1627-1629
[19]Barbaroux C,Breda N.Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuseporous beech trees.Tree Physiology,2002,22(17):1201-1210
[20]Richardson A D,Carbone M S,Keenan T F,et al.Seasonal dynamics and age of stemwood nonstructural carbohydrates in temperate forest trees.New Phytol,2013,197(3):850-861
[21]Dickson R E.Carbon and nitrogen allocation in trees.Annales Des Sciences Forestieres,1989,46:S631-S647
[22]Landhausser S M,Lieffers V J.Seasonal changes in carbohydrate reserves in mature northern Populus tremuloides clones.Trees-Structure and Function,2003,17(6):471-476
[23]Breda N,Huc R,Granier A,et al.Temperate forest trees and stands under severe drought:a review of ecophysiological responses,adaptation processes and long-term consequences.Annals of Forest Science,2006,63(6):625-644
[24]Hartmann H,Ziegler W,Kolle O,et al.Thirst beats hunger-declining hydration during drought prevents carbon starvation in Norway spruce saplings.New Phytologist,2013,200(2):340-349
[25]Dickman L T,Mcdowell N G,Sevanto S,et al.Carbohydrate dynamics and mortality in a pinonjuniper woodland under three future precipitation scenarios.Plant Cell And Environment,2015,38(4):729-739
[26]潘庆民,韩兴国,白永飞,等.植物非结构性贮藏碳水化合物的生理生态学研究进展.植物学通报,2002,19(1):30-38
[27]Gaudet D A,Laroche A,Yoshida M.Low temperature-wheat-fungal interactions:a carbohydrate connection.Physiologia Plantarum,1999,106(4):437-444
[28]Kozlowski T T,Pallardy S G.Acclimation and adaptive responses of woody plants to environmental stresses.Botanical Review,2002,68(2):270-334
[29]Fischer C,Holl W.Food reserves of scots pine(Pinus-sylvestris L.).1.seasonal changes in the carbohydrate and fat reserves of pine needles.TreesStructure and Function,1991,5(4):187-195
[30]Piispanen R,Saranp??P.Variation of non-structural carbohydrates in silver birch(Betula pendula Roth)wood.Trees,2001,15(7):444-451
[31]Fajardo A,Piper F I,Pfund L,et al.Variation of mobile carbon reserves in trees at the alpine treeline ecotone is under environmental control.New Phytologist,2012,195(4):794-802
[32]Sala A,Woodruff D R,Meinzer F C.Carbon dynamics in trees:feast or famine?.Tree Physiol,2012,32(6):764-775
[33]Fajardo A,Piper F I,Hoch G.Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients.Annuals of Botany,2013,112(3):623-631
[34]Sala A,Mencuccini M.Ecosystem science:plump trees win under drought.Nature Climate Change,2014,4(8):666-667