长白山阔叶红松林演替序列植物-凋落物-土壤碳同位素特征
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摘要
稳定碳同位素组成能精确指示生态系统碳循环过程,可以为深入研究森林演替进程对碳循环过程和固碳潜力的影响提供关键信息.利用稳定碳同位素技术对长白山阔叶红松林演替序列3种林分——中龄杨桦次生林、成熟杨桦次生林、阔叶红松林的叶片、树干、根系、凋落物和土壤δ~(13)C值及碳、氮元素含量进行测定.结果表明:各演替序列优势树种叶片δ~(13)C从冠上到冠下均呈降低趋势;树干δ~(13)C表现为树皮小于木质部;根系δ~(13)C表现为细根小于粗根.阔叶红松林未分解凋落物δ~(13)C小于半分解及全分解凋落物,次生林相反;土壤δ~(13)C沿深度逐渐增加.总体上,δ~(13)C值叶片<凋落物<根系<树干<土壤,说明植物各器官之间有明显的碳同位素分馏效应,且相同器官不同部位之间也存在差异;植物δ~(13)C沿演替方向先减小后增加,土壤δ~(13)C沿演替方向不断增加,且变化规律可以通过氮元素含量与碳同位素分馏效应的关系解释,说明长白山阔叶红松林演替过程优势树种和碳周转速率的变化影响了碳同位素分馏.
Stable carbon isotope composition can accurately indicate ecosystem carbon cycling and provide key information for the study of the influence of forest succession on the carbon cycling and carbon sequestration potential. We measured the δ~(13)C values and carbon and nitrogen contents of leaf, trunk, root, litter, and soil along a forest successional gradient in Changbai Mountain, which included a middle-aged poplar-birch secondary forest, a mature poplar-birch secondary forest, and an old-growth broad-leaved Korean pine forest. The results showed that leaf δ~(13)C reduced with their position from the upper canopy to lower canopy, bark δ~(13)C was less than xylem, fine root δ~(13)C was less than course root. In contrast to the secondary forests, δ~(13)C of the undecomposed litter layer was less than that of the semi-decomposed layer and decomposed litter layer in the broad-leaved Korean pine forest. Soil δ~(13)C increased with depth. The ascending order of mean δ~(13)C was leaf, litter, root, trunk, and soil, indicating that there is obvious fractionation among different organs of plants and among different parts of a specific organ. In addition, plant δ~(13)C first decreased and then increased with the succession process, but soil δ~(13)C increased with the succession processes. The different patterns of the changes of plant and soil δ~(13)C along forest succession could be explained by the relationship between nitrogen content and carbon isotope fractionation effect, indicating that carbon isotope fractionation was affected by the change of dominant tree species and the variation of carbon turnover rate.
Stable carbon isotope composition can accurately indicate ecosystem carbon cycling and provide key information for the study of the influence of forest succession on the carbon cycling and carbon sequestration potential. We measured the δ~(13)C values and carbon and nitrogen contents of leaf, trunk, root, litter, and soil along a forest successional gradient in Changbai Mountain, which included a middle-aged poplar-birch secondary forest, a mature poplar-birch secondary forest, and an old-growth broad-leaved Korean pine forest. The results showed that leaf δ~(13)C reduced with their position from the upper canopy to lower canopy, bark δ~(13)C was less than xylem, fine root δ~(13)C was less than course root. In contrast to the secondary forests, δ~(13)C of the undecomposed litter layer was less than that of the semi-decomposed layer and decomposed litter layer in the broad-leaved Korean pine forest. Soil δ~(13)C increased with depth. The ascending order of mean δ~(13)C was leaf, litter, root, trunk, and soil, indicating that there is obvious fractionation among different organs of plants and among different parts of a specific organ. In addition, plant δ~(13)C first decreased and then increased with the succession process, but soil δ~(13)C increased with the succession processes. The different patterns of the changes of plant and soil δ~(13)C along forest succession could be explained by the relationship between nitrogen content and carbon isotope fractionation effect, indicating that carbon isotope fractionation was affected by the change of dominant tree species and the variation of carbon turnover rate.
引文
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[22] Leavitt SW,Long A.Stable-carbon isotope variability in tree foliage and wood.Ecology,1986,67:1002-1010
[23] Benner R,Fogel ML,Sprague EK,et al.Depletion of 13C in lignin and its implications for stable carbon isotope studies.Nature,1987,329:708
[24] H?gberg P,Nordgren A,?gren GI.Carbon allocation between tree root growth and root respiration in boreal pine forest.Oecologia,2002,132:579-581
[25] Xue J-H (薛建辉),Wang Z (王智),Lyu X-S (吕祥生).Progress on the interaction between tree roots and soil environment.Journal of Nanjing Forestry Unversity (Natural Science) (南京林业大学学报:自然科学版),2002,26(3):79-84 (in Chinese)
[26] Si G-Y (司高月),Li X-Y (李晓玉),Cheng S-L (程淑兰),et al.Organic carbon dynamics of the leaf-litter-soil continuum in the typical forests of the Changbai Mountain transect:An analysis of stable carbon isotope technology.Acta Ecologica Sinica (生态学报),2017,37(16):5285-5293 (in Chinese)
[27] Xiong X (熊鑫),Zhang H-L (张慧玲),Wu J-P (吴建平),et al.13C and 15N isotopic signatures of plant-soil continuum along a successional gradient in Dinghushan Biosphere Reserve.Chinese Journal of Plant Ecology (植物生态学报),2016,40(6):533-542 (in Chinese)
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[29] Natelhoffer KJ,Fry B.Controls on natural nitrogen-15 and carbon-13 abundances in forest soil organic matter.Soil Science Society of America Journal,1988,52:1633-1640
[30] Becker-Heidmann P,Scharpenseel HW.Thin layer δ13C and D14C monitoring of “lessive” soil profiles.Radiocarbon,1986,28:383-390
[31] Sparling G.Ratio of microbial biomass carbon to soil organic carbon as a sensitive indicator of changes in soil organic matter.Soil Research,1992,30:195-207
[32] Vaughn BH,Evans CU,White JWC,et al.Global network measurements of atmospheric trace gas isotopes// West JB,Bowen GJ,Dawson TE,eds.Isoscapes:Understanding Movement,Pattern,and Process on Earth Through Isotope Mapping.Dordrecht:Springer,2010:3-31
[33] Li M-C (李明财),Luo T-X (罗天祥),Liu X-S (刘新圣),et al.Distribution characteristics of δ13C values in different organs of Abies georgei growing at alpine timberline.Chinese Journal of Applied Ecology (应用生态学报),2007,18(12):2654-2660 (in Chinese)
[34] Hobbie EA,Werner RA.Intramolecular,compound-specific,and bulk carbon isotope patterns in C3 and C4 plants:A review and synthesis.New Phytologist,2004,161:371-385
[35] Chai H (柴华),Zhong S-Z (钟尚志),Cui H-Y (崔海莹),et al.Variation in the carbon isotope composition of CO2 derived from plant autotrophic respiration.Acta Ecologica Sinica (生态学报),2018,38(8):2616-2624 (in Chinese)
[36] Marshall JD,Zhang J.Carbon isotope discrimination and water-use efficiency in native plants of the north-central Rockies.Ecology,1994,75:1887-1895
[37] K?rner C,Farquhar GD,Wong SC.Carbon isotope discrimination by plants follows latitudinal and altitudinal trends.Oecologia,1991,88:30-40
[38] Piao H-C (朴河春),Zhu J-M (朱建明),Zhu S-F (朱书法),et al.Altitudinal variations of nutrient concentrations and carbon isotope compositionsin a C3 plant and the effects of nutrient interactions on carbon isotope discrimination in limestone areas of southwest China.Advance in Earth Sciences (地球科学进展),2004,19(suppl.1):412-418 (in Chinese)
[39] Cernusak LA,Ubierna N,Winter K,et al.Environmental and physiological determinants of carbon isotope discrimination in terrestrial plants.New Phytologist,2013,200:950-965
[40] Vitousek PM,Field CB,Matson PA.Variation in foliar δ13C in Hawaiian Metrosideros polymorpha:A case of internal resistance?Oecologia,1990,84:362-370
[41] Wang C,Wei H,Liu D,et al.Depth profiles of soil carbon isotopes along a semi-arid grassland transect in northern China.Plant and Soil,2017,417:43-52
[2] Fang Y-T (方运霆),Mo J-M (莫江明),Peng S-L (彭少麟),et al.Role of forest succession on carbon sequestration of forest ecosystems in lower subtropical China.Acta Ecologica Sinica (生态学报),2003,23(9):1685-1694 (in Chinese)
[3] De Kovel CGF,Van Mierlo AEM,Wilms YJO,et al.Carbon and nitrogen in soil and vegetation at sites differing in successional age.Plant Ecology,2000,149:43-50
[4] Thuille A,Schulze ED.Carbon dynamics in successional and afforested spruce stands in Thuringia and the Alps.Global Change Biology,2006,12:325-342
[5] Bowling DR,Pataki DE,Randerson JT.Carbon isotopes in terrestrial ecosystem pools and CO2 fluxes.New Phytologist,2008,178:24-40
[6] Farquhar GD,Ehleringer JR,Hubick KT.Carbon isotope discrimination and photosynthesis.Annual Review of Plant Physiology and Plant Molecular Biology,1989,40:503-537
[7] Farquhar GD,O’Leary MH,Berry JA.On the relationship between carbon isotope discrimination and the intercellular carbon dioxide concentration in leaves.Functional Plant Biology,1982,9:121-137
[8] Bernoux M,Cerri CC,Neill C,et al.The use of stable carbon isotopes for estimating soil organic matter turnover rates.Geoderma,1998,82:43-58
[9] Yang L-W (杨丽韫),Luo T-X (罗天祥),Wu S-T (吴松涛).Root biomass and underground C and N storage of primitive Korean pine and broad-leaved climax forest in Changbai Mountains at its different succession stages.Chinese Journal of Applied Ecology (应用生态学报),2005,16(7):1195-1199 (in Chinese)
[10] Zhang X (张雪),Han S-J (韩士杰),Wang S-Q (王树起),et al.Change of soil organic carbon fractions at different successional stages of Betula platyphylla forest in Changbai Mountains.Chinese Journal of Ecology (生态学杂志),2016,35(2):282-289 (in Chinese)
[11] Zhang C-Y (张春雨),Zhao X-H (赵秀海),Zhao Y-Z (赵亚洲).Community structure in different successional stages in north temperate forests of Changbai Mountains,China.Chinese Journal of Plant Ecology (植物生态学报),2009,33(6):1090-1100 (in Chinese)
[12] Hu Y-S (胡耀升),Me X-Y (么旭阳),Liu Y-H (刘艳红).N and P stoichiometric traits of plant and soil in different forest succession stages in Changbai Mountains.Chinese Journal of Applied Ecology (应用生态学报),2014,25(3):632-638 (in Chinese)
[13] Dai LM,Shao GF,Chen G,et al.Forest cutting and regeneration methodology on Changbai Mountain.Journal of Forestry Research,2003,14:56-60
[14] Schleser GH.δ13C pattern in a forest tree as an indicator of carbon transfer in trees.Ecology,1992,73:1922-1925
[15] Buchmann N,Guehl JM,Barigah TS,et al.Interseasonal comparison of CO2 concentrations,isotopic composition,and carbon dynamics in an Amazonian rainforest (French Guiana).Oecologia,1997,110:120-131
[16] Wang Y-N (王云霓),Xiong W (熊伟),Wang Y-H (王彦辉),et al.Spatial-temporal pattern of stable carbon isotope composition of dominant tree species in Liupanshan Mountain.Research of Soil and Water Conservation (水土保持研究),2012,19(3):42-47 (in Chinese)
[17] Martinelli LA,Almeida S,Brown IF,et al.Stable carbon isotope ratio of tree leaves,boles and fine litter in a tropical forest in Rond?nia,Brazil.Oecologia,1998,114:170-179
[18] Da Silveira L,Sternberg L,Mulkey SS,et al.Ecological interpretation of leaf carbon isotope ratios:Influence of respired carbon dioxide.Ecology,1989,70:1317-1324
[19] Buchmann N,Kao WY,Ehleringer J.Influence of stand structure on carbon-13 of vegetation,soils,and canopy air within deciduous and evergreen forests in Utah,Uni-ted States.Oecologia,1997,110:109-119
[20] Chen T (陈拓),Qin D-H (秦大河),He Y-Q (何元庆),et al.The pattern of stable carbon isotope ratios in Sabina przewalskii.Journal of Glacio-lgy and Geocryology (冰川冻土),2002,24(5):571-573 (in Chinese)
[21] Han X-G (韩兴国),Yan C-R (严昌荣),Chen L-Z (陈灵芝),et al.Stable carbon isotope characteristics of some woody plants in warm temperate zone.Chinese Journal of Applied Ecology (应用生态学报),2000,11(4):497-500 (in Chinese)
[22] Leavitt SW,Long A.Stable-carbon isotope variability in tree foliage and wood.Ecology,1986,67:1002-1010
[23] Benner R,Fogel ML,Sprague EK,et al.Depletion of 13C in lignin and its implications for stable carbon isotope studies.Nature,1987,329:708
[24] H?gberg P,Nordgren A,?gren GI.Carbon allocation between tree root growth and root respiration in boreal pine forest.Oecologia,2002,132:579-581
[25] Xue J-H (薛建辉),Wang Z (王智),Lyu X-S (吕祥生).Progress on the interaction between tree roots and soil environment.Journal of Nanjing Forestry Unversity (Natural Science) (南京林业大学学报:自然科学版),2002,26(3):79-84 (in Chinese)
[26] Si G-Y (司高月),Li X-Y (李晓玉),Cheng S-L (程淑兰),et al.Organic carbon dynamics of the leaf-litter-soil continuum in the typical forests of the Changbai Mountain transect:An analysis of stable carbon isotope technology.Acta Ecologica Sinica (生态学报),2017,37(16):5285-5293 (in Chinese)
[27] Xiong X (熊鑫),Zhang H-L (张慧玲),Wu J-P (吴建平),et al.13C and 15N isotopic signatures of plant-soil continuum along a successional gradient in Dinghushan Biosphere Reserve.Chinese Journal of Plant Ecology (植物生态学报),2016,40(6):533-542 (in Chinese)
[28] Ehleringer JR,Buchmann N,Flanagan LB.Carbon isotope ratios in belowground carbon cycle processes.Ecological Applications,2000,10:412-422
[29] Natelhoffer KJ,Fry B.Controls on natural nitrogen-15 and carbon-13 abundances in forest soil organic matter.Soil Science Society of America Journal,1988,52:1633-1640
[30] Becker-Heidmann P,Scharpenseel HW.Thin layer δ13C and D14C monitoring of “lessive” soil profiles.Radiocarbon,1986,28:383-390
[31] Sparling G.Ratio of microbial biomass carbon to soil organic carbon as a sensitive indicator of changes in soil organic matter.Soil Research,1992,30:195-207
[32] Vaughn BH,Evans CU,White JWC,et al.Global network measurements of atmospheric trace gas isotopes// West JB,Bowen GJ,Dawson TE,eds.Isoscapes:Understanding Movement,Pattern,and Process on Earth Through Isotope Mapping.Dordrecht:Springer,2010:3-31
[33] Li M-C (李明财),Luo T-X (罗天祥),Liu X-S (刘新圣),et al.Distribution characteristics of δ13C values in different organs of Abies georgei growing at alpine timberline.Chinese Journal of Applied Ecology (应用生态学报),2007,18(12):2654-2660 (in Chinese)
[34] Hobbie EA,Werner RA.Intramolecular,compound-specific,and bulk carbon isotope patterns in C3 and C4 plants:A review and synthesis.New Phytologist,2004,161:371-385
[35] Chai H (柴华),Zhong S-Z (钟尚志),Cui H-Y (崔海莹),et al.Variation in the carbon isotope composition of CO2 derived from plant autotrophic respiration.Acta Ecologica Sinica (生态学报),2018,38(8):2616-2624 (in Chinese)
[36] Marshall JD,Zhang J.Carbon isotope discrimination and water-use efficiency in native plants of the north-central Rockies.Ecology,1994,75:1887-1895
[37] K?rner C,Farquhar GD,Wong SC.Carbon isotope discrimination by plants follows latitudinal and altitudinal trends.Oecologia,1991,88:30-40
[38] Piao H-C (朴河春),Zhu J-M (朱建明),Zhu S-F (朱书法),et al.Altitudinal variations of nutrient concentrations and carbon isotope compositionsin a C3 plant and the effects of nutrient interactions on carbon isotope discrimination in limestone areas of southwest China.Advance in Earth Sciences (地球科学进展),2004,19(suppl.1):412-418 (in Chinese)
[39] Cernusak LA,Ubierna N,Winter K,et al.Environmental and physiological determinants of carbon isotope discrimination in terrestrial plants.New Phytologist,2013,200:950-965
[40] Vitousek PM,Field CB,Matson PA.Variation in foliar δ13C in Hawaiian Metrosideros polymorpha:A case of internal resistance?Oecologia,1990,84:362-370
[41] Wang C,Wei H,Liu D,et al.Depth profiles of soil carbon isotopes along a semi-arid grassland transect in northern China.Plant and Soil,2017,417:43-52