遮光对杉木幼苗树干表面CO_2通量的影响
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摘要
树干表面CO_2通量是森林生态系统碳收支的重要组成部分,对全球碳平衡产生重要影响。近年来,全球变化导致植物光合产物供应发生改变,这将影响树干表面CO_2通量。然而,关于光合产物供应如何影响树干表面CO_2通量的机理仍不清楚。以盆栽杉木(Cunninghamia lanceolata)幼苗为研究对象,采用遮光方法减少光合产物供应,通过自制呼吸气室,使用Li-8100测定树干表面CO_2通量,并结合树干可溶性糖、淀粉和非结构性碳及树干温度等数据分析遮光对树干表面CO_2通量的影响。结果表明:遮光后树干可溶性糖、淀粉和非结构性碳含量分别显著下降了55.0%、78.9%和64.3%。遮光处理阶段树干表面CO_2通量平均下降39.9%,且下降幅度随着遮光时间的延长而增加;此外,遮光降低了树干表面CO_2通量的温度敏感性。恢复光照后,树干表面CO_2通量、树干可溶性糖、淀粉和非结构性碳含量以及树干温度均恢复至对照水平。可见,光合产物供应变化对树干表面CO_2通量具有调控作用,而且能够通过调控树干表面CO_2通量对温度变化的响应对全球碳循环产生重要影响。
Stem surface CO_2 effluxes are an important part of the carbon(C) budget of forest ecosystems, which play an important role in sustaining global C balance. In recent years, global changes have altered the supply of plant photosynthates, and this could affect plant stem surface CO_2 effluxes. However, the underlying mechanisms of how changes in plant photosynthates affect stem surface CO_2 effluxes are still unclear. In the present study, potted Chinese fir(Cunninghamia lanceolata) seedlings were used, and shading was applied to alter the supply of photosynthate. We measured stem surface CO_2 effluxes using a Li-8100 Automated Soil Gas Flux System, and measured the concentrations of stem soluble sugar, starch, non-structural C, and stem temperature to clarify the effects of shading on stem surface CO_2 effluxes. The results showed that the concentrations of stem soluble sugar, starch, and non-structural C were significantly reduced by 55.0%, 78.9% and 64.3%, respectively. The mean stem surface CO_2 effluxes significantly decreased by 39.9% during shading, and the decrease rate gradually increased as shading time increased. Moreover, shading decreased the temperature sensibility of stem surface CO_2 effluxes. The stem surface CO_2 effluxes and the concentrations of stem soluble sugar, starch, non-structural C, and stem temperature were recovered after regaining normal light conditions, as compared with the control. These results suggest that the changes in photosynthate supply could affect stem surface CO_2 effluxes and consequently global C cycling by regulating the response of stem surface CO_2 effluxes to temperature.
Stem surface CO_2 effluxes are an important part of the carbon(C) budget of forest ecosystems, which play an important role in sustaining global C balance. In recent years, global changes have altered the supply of plant photosynthates, and this could affect plant stem surface CO_2 effluxes. However, the underlying mechanisms of how changes in plant photosynthates affect stem surface CO_2 effluxes are still unclear. In the present study, potted Chinese fir(Cunninghamia lanceolata) seedlings were used, and shading was applied to alter the supply of photosynthate. We measured stem surface CO_2 effluxes using a Li-8100 Automated Soil Gas Flux System, and measured the concentrations of stem soluble sugar, starch, non-structural C, and stem temperature to clarify the effects of shading on stem surface CO_2 effluxes. The results showed that the concentrations of stem soluble sugar, starch, and non-structural C were significantly reduced by 55.0%, 78.9% and 64.3%, respectively. The mean stem surface CO_2 effluxes significantly decreased by 39.9% during shading, and the decrease rate gradually increased as shading time increased. Moreover, shading decreased the temperature sensibility of stem surface CO_2 effluxes. The stem surface CO_2 effluxes and the concentrations of stem soluble sugar, starch, non-structural C, and stem temperature were recovered after regaining normal light conditions, as compared with the control. These results suggest that the changes in photosynthate supply could affect stem surface CO_2 effluxes and consequently global C cycling by regulating the response of stem surface CO_2 effluxes to temperature.
引文
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[14] 刘兰兰,汪思龙,杨庆朋.修枝对杉木人工林树干表面和土壤CO2释放通量的短期影响.生态学杂志,2014,33(6):1474- 1479.
[15] Poorter H,Niklas K J,Reich P B,Oleksyn J,Poot P,Mommer L.Biomass allocation to leaves,stems and roots:meta-analyses of interspecific variation and environmental control.New Phytologist,2012,193(1):30- 50.
[16] 许中旗,黄选瑞,徐成立,许晴,纪晓林.光照条件对蒙古栎幼苗生长及形态特征的影响.生态学报,2009,29(3):1121- 1128.
[17] Kuzyakov Y,Cheng W.Photosynthesis controls of rhizosphere respiration and organic matter decomposition.Soil Biology and Biochemistry,2001,33(14):1915- 1925.
[18] Bahn M,Lattanzi F A,Hasibeder R,Wild B,Koranda M,Danese V,Brüggemann N,Schmitt M,Siegwolf R,Richter A.Responses of belowground carbon allocation dynamics to extended shading in mountain grassland.New Phytologist,2013,198(1):116- 126.
[19] Damesin C,Ceschia E,Le Goff N,Ottorini J M,Dufrêne E.Stem and branch respiration of beech:from tree measurements to estimations at the stand level.New Phytologist,2002,153(1):159- 172.
[20] Wieser G,Bahn M.Seasonal and spatial variation of woody tissue respiration in a Pinus cembra tree at the alpine timberline in the central Austrian Alps.Trees,2004,18(5):576- 580.
[21] Gruber A,Wieser G,Oberhuber W.Intra-annual dynamics of stem CO2 efflux in relation to cambial activity and xylem development in Pinus cembra.Tree Physiology,2009,29(5):641- 649.
[17] 严玉平,沙丽清,曹敏.西双版纳热带季节雨林优势树种树干呼吸特征.植物生态学报,2008,32(1):23- 30.
[23] Yang Q P,Liu L L,Zhang W D,Xu M,Wang S L.Different responses of stem and soil CO2 efflux to pruning in a Chinese fir (Cunninghamia lanceolata) plantation.Trees,2015,29(4):1207- 1218.
[24] 国家林业局.中国森林资源报告(2009- 2013).北京:中国林业出版社,2014.
[25] Chen D M,Zhang C L,Wu J P,Zhou L X,Lin Y B,Fu S L.Subtropical plantations are large carbon sinks:evidence from two monoculture plantations in South China.Agricultural and Forest Meteorology,2011,151(9):1214- 1225.
[26] Bergmeyer H U,Bergmeyer J,Grassl M.Methods of enzymatic analysis.Cambridge,UK:VCH Publishers (UK) Ltd,1988.
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[28] Xu X L,Kuzyakov Y,Wanek W,Richter A.Root-derived respiration and non-structural carbon of rice seedlings.European Journal of Soil Biology,2008,44(1):22- 29.
[29] He T X,Wang Q K,Wang S L,Zhang F Y.Nitrogen addition altered the effect of belowground C allocation on soil respiration in a subtropical forest.PLoS One,2016,11(5):e0155881.
[30] 王文杰,王慧梅,祖元刚,李雪莹,小池孝良.林木非同化器官与土壤呼吸的温度系数Q10值的特征分析.植物生态学报,2005,29(4):680- 691.
[31] Atkin O K,Tjoelker M G.Thermal acclimation and the dynamic response of plant respiration to temperature.Trends in Plant Science,2003,8(7):343- 351.
[32] Azcón-Bieto J,Lambers H,Day D A.Effect of photosynthesis and carbohydrate status on respiratory rates and the involvement of the alternative pathway in leaf respiration.Plant Physiology,1983,72(3):598- 603.
[33] Covey-Crump E M,Attwood R G,Atkin O K.Regulation of root respiration in two species of plantago that differ in relative growth rate:the effect of short- and long-term changes in temperature.Plant,Cell & Environment,2002,25(11):1501- 1513.
[2] Luyssaert S,Inglima I,Jung M,Richardson A D,Reichstein M,Papale D,Piao S L,Schulze E D,Wingate L,Matteucci G,Aragao L,Aubinet M,Beer C,Bernhofer C,Black K G,Bonal D,Bonnefond J M,Chambers J,Ciais P,Cook B,Davis K J,Dolman A J,Gielen B,Goulden M,Grace J,Granier A,Grelle A,Griffis T,Grünwald T,Guidolotti G,Hanson P J,Harding R,Hollinger D Y,Hutyra L R,Kolari P,Kruijt B,Kutsch W,Lagergren F,Laurila T,Law B E,Le Maire G,Lindroth A,Loustau D,Malhi Y,Mateus J,Migliavacca M,Misson L,Montagnani L,Moncrieff J,Moors E,Munger J W,Nikinmaa E,Ollinger S V,Pita G,Rebmann C,Roupsard O,Saigusa N,Sanz M J,Seufert G,Sierra C,Smith M L,Tang J,Valentini R,Vesala T,Janssens I A.CO2 balance of boreal,temperate,and tropical forests derived from a global database.Global Change Biology,2007,13(12):2509- 2537.
[3] Xu M,DeBiase T A,Qi Y,Goldstein A,Liu Z G.Ecosystem respiration in a young Ponderosa Pine plantation in the Sierra Nevada Mountains,California.Tree Physiology,2001,21(5):309- 318.
[4] Maier C A,Albaugh T J,Allen H L,Dougherty P M.Respiratory carbon use and carbon storage in mid-rotation loblolly pine (Pinus taeda L.) plantations:the effect of site resources on the stand carbon balance.Global Change Biology,2004,10(8):1335- 1350.
[5] Wang W J,Zu Y G,Wang H M,Li X Y,Hirano T,Koike T.Newly-formed photosynthates and the respiration rate of girdled stems of Korean pine (Pinus koraiensis Sieb.et Zucc.).Photosynthetica,2006,44(1):147- 150.
[6] Maier C A,Johnsen K H,Clinton B D,Ludovici K H.Relationships between stem CO2 efflux,substrate supply,and growth in young loblolly pine trees.New Phytologist,2010,185(2):502- 513.
[7] Yang Q P,Xu M,Chi Y G,Zheng Y P.Relationship between stem CO2 efflux and stem temperature at different measuring depths in Pinus massoniana trees.Acta Ecologica Sinica,2016,36(4):229- 235.
[8] 马玉娥,项文化,雷丕锋.林木树干呼吸变化及其影响因素研究进展.植物生态学报,2007,31(3):403- 412.
[9] 崔之益,徐大平,杨曾奖,张宁南,刘小金,洪舟.土壤含水量对旱季降香黄檀树干呼吸和非结构性碳水化合物的影响.生态学杂志,2018,37(2):374- 382.
[10] Edwards N T,Tschaplinski T J,Norby R J.Stem respiration increases in CO2-enriched sweetgum trees.New Phytologist,2002,155(2):239- 248.
[11] Wertin T M,Teskey R O.Close coupling of whole-plant respiration to net photosynthesis and carbohydrates.Tree Physiology,2008,28(12):1831- 1840.
[12] Clinton B D,Maier C A,Ford C R,Mitchell R J.Transient changes in transpiration,and stem and soil CO2 ef?ux in longleaf pine (Pinus palustris Mill.) following fire-induced leaf area reduction.Trees,2011,25(6):997- 1007.
[13] H?gberg P,Nordgren A,Buchmann N,Taylor A F S,Ekblad A,H?gberg M N,Nyberg G,Ottosson-L?fvenius M,Read D J.Large-scale forest girdling shows that current photosynthesis drives soil respiration.Nature,2001,411(6839):789- 792.
[14] 刘兰兰,汪思龙,杨庆朋.修枝对杉木人工林树干表面和土壤CO2释放通量的短期影响.生态学杂志,2014,33(6):1474- 1479.
[15] Poorter H,Niklas K J,Reich P B,Oleksyn J,Poot P,Mommer L.Biomass allocation to leaves,stems and roots:meta-analyses of interspecific variation and environmental control.New Phytologist,2012,193(1):30- 50.
[16] 许中旗,黄选瑞,徐成立,许晴,纪晓林.光照条件对蒙古栎幼苗生长及形态特征的影响.生态学报,2009,29(3):1121- 1128.
[17] Kuzyakov Y,Cheng W.Photosynthesis controls of rhizosphere respiration and organic matter decomposition.Soil Biology and Biochemistry,2001,33(14):1915- 1925.
[18] Bahn M,Lattanzi F A,Hasibeder R,Wild B,Koranda M,Danese V,Brüggemann N,Schmitt M,Siegwolf R,Richter A.Responses of belowground carbon allocation dynamics to extended shading in mountain grassland.New Phytologist,2013,198(1):116- 126.
[19] Damesin C,Ceschia E,Le Goff N,Ottorini J M,Dufrêne E.Stem and branch respiration of beech:from tree measurements to estimations at the stand level.New Phytologist,2002,153(1):159- 172.
[20] Wieser G,Bahn M.Seasonal and spatial variation of woody tissue respiration in a Pinus cembra tree at the alpine timberline in the central Austrian Alps.Trees,2004,18(5):576- 580.
[21] Gruber A,Wieser G,Oberhuber W.Intra-annual dynamics of stem CO2 efflux in relation to cambial activity and xylem development in Pinus cembra.Tree Physiology,2009,29(5):641- 649.
[17] 严玉平,沙丽清,曹敏.西双版纳热带季节雨林优势树种树干呼吸特征.植物生态学报,2008,32(1):23- 30.
[23] Yang Q P,Liu L L,Zhang W D,Xu M,Wang S L.Different responses of stem and soil CO2 efflux to pruning in a Chinese fir (Cunninghamia lanceolata) plantation.Trees,2015,29(4):1207- 1218.
[24] 国家林业局.中国森林资源报告(2009- 2013).北京:中国林业出版社,2014.
[25] Chen D M,Zhang C L,Wu J P,Zhou L X,Lin Y B,Fu S L.Subtropical plantations are large carbon sinks:evidence from two monoculture plantations in South China.Agricultural and Forest Meteorology,2011,151(9):1214- 1225.
[26] Bergmeyer H U,Bergmeyer J,Grassl M.Methods of enzymatic analysis.Cambridge,UK:VCH Publishers (UK) Ltd,1988.
[27] McCleary B V,Gibson T S,Mugford D C.Measurement of total starch in cereal products by amyloglucosidase—α-amylase method:collaborative study.Journal of AOAC International,1997,80(3):571- 579.
[28] Xu X L,Kuzyakov Y,Wanek W,Richter A.Root-derived respiration and non-structural carbon of rice seedlings.European Journal of Soil Biology,2008,44(1):22- 29.
[29] He T X,Wang Q K,Wang S L,Zhang F Y.Nitrogen addition altered the effect of belowground C allocation on soil respiration in a subtropical forest.PLoS One,2016,11(5):e0155881.
[30] 王文杰,王慧梅,祖元刚,李雪莹,小池孝良.林木非同化器官与土壤呼吸的温度系数Q10值的特征分析.植物生态学报,2005,29(4):680- 691.
[31] Atkin O K,Tjoelker M G.Thermal acclimation and the dynamic response of plant respiration to temperature.Trends in Plant Science,2003,8(7):343- 351.
[32] Azcón-Bieto J,Lambers H,Day D A.Effect of photosynthesis and carbohydrate status on respiratory rates and the involvement of the alternative pathway in leaf respiration.Plant Physiology,1983,72(3):598- 603.
[33] Covey-Crump E M,Attwood R G,Atkin O K.Regulation of root respiration in two species of plantago that differ in relative growth rate:the effect of short- and long-term changes in temperature.Plant,Cell & Environment,2002,25(11):1501- 1513.