不同物候时期华西雨屏区四个树种新鲜凋落物可提取腐殖质碳动态
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摘要
凋落物的可提取腐殖质碳可随着植物生长节律及物候时期的变化而变化,并进而影响物质循环的过程,为深入了解以凋落物为载体的生态系统物质循环特征,该研究以华西雨屏区麻栎(Quercus acutissima)、楠木(Phoebe zhennan)、柳杉(Cryptomeria fortune)和喜树(Camptotheca acuminate)为对象,通过定点动态收集萌芽期、展叶期、盛叶期和落叶期的不同类型凋落物,分析其可提取腐殖质碳(extractable humus carbon,HC)、胡敏酸碳(humic acid carbon,HAC)、富里酸碳(fulvic acid carbon,FAC)以及胡敏酸碳/富里酸碳(humic acid carbon/fulvic acid carbon,HAC/FAC)的差异。结果表明:相对于其他凋落器官,在同一物候时期凋落叶中的HC和HAC含量都最高,大致都表现为凋落叶>凋落枝>凋落果,且在落叶树种中更为显著;相对于其他时期,展叶期四个树种凋落叶均表现出较高的FAC含量,但不同物候期凋落枝和凋落果的HAC含量以及FAC含量受物种的影响较大。尽管HAC/FAC在各器官间差异并不显著,但落叶树种相同器官的HAC/FAC低于常绿树种,表明落叶树种凋落物富里酸相对含量较高,形成速度相对较快,难降解程度也相对较大。统计分析表明关键时期、物种类型、器官类型及其相互作用对凋落物中HC、HAC、FAC含量和HAC/FAC均具有不同程度的影响,这为进一步深入认识区域生态系统以凋落物为载体的物质循环过程提供了理论依据及新的思路。
The extractable humus carbon of the litter can change with the rhythm of the plant growth and the phenological period,and then affect the material circulation process. In order to fully understand the features of the litterfall-supported regional ecosystem cycle,Quercus acutissima,Phoebe zhennan,Cryptomeria fortune,and Camptotheca acuminatain were selected in the Rainy Area of West China. Different types of litters in the leaf budding period,leaf expanding period,leaf maturing period,and leaf falling period were dynamically collected through fixed-point analysis of the differences in the extractable humus carbon( HC),humic acid carbon( HAC),fulvic acid carbon( FAC),and humic acid carbon/fulvic acid carbon( HAC/FAC). The results were as follows: The contents of HC and HAC in leaf litter were the highest in the same critical period,and they all showed as leaf litter> twig litter> fruit litter,and also showed that the evergreen tree species was not significant but the deciduous tree species was significant; Compared with other periods,leaf litter of the four tree species exhibited high FAC content during the leaf expanding period,However,HAC content and FAC content of twig litter and fruit litter in phenological stages are greatly affected by species. Humic acid carbon/Fulvic acid carbon( HAC/FAC) in leaf litter of four tree species to characterize the relative content and formation rate of humic acid,fulvic acid during the humification of leaf litter,there was no significant difference between organs,showing that the deciduous tree species had lower HAC/FAC than evergreen tree species. The present study indicates that stage types,species types,organ types and their interactions have different degrees of influence on HC,HAC,FAC content and HAC/FAC in litter. The results provide a theoretical information and a new direction for further understanding the material circulation process of litterfall in the regional ecosystem.
The extractable humus carbon of the litter can change with the rhythm of the plant growth and the phenological period,and then affect the material circulation process. In order to fully understand the features of the litterfall-supported regional ecosystem cycle,Quercus acutissima,Phoebe zhennan,Cryptomeria fortune,and Camptotheca acuminatain were selected in the Rainy Area of West China. Different types of litters in the leaf budding period,leaf expanding period,leaf maturing period,and leaf falling period were dynamically collected through fixed-point analysis of the differences in the extractable humus carbon( HC),humic acid carbon( HAC),fulvic acid carbon( FAC),and humic acid carbon/fulvic acid carbon( HAC/FAC). The results were as follows: The contents of HC and HAC in leaf litter were the highest in the same critical period,and they all showed as leaf litter> twig litter> fruit litter,and also showed that the evergreen tree species was not significant but the deciduous tree species was significant; Compared with other periods,leaf litter of the four tree species exhibited high FAC content during the leaf expanding period,However,HAC content and FAC content of twig litter and fruit litter in phenological stages are greatly affected by species. Humic acid carbon/Fulvic acid carbon( HAC/FAC) in leaf litter of four tree species to characterize the relative content and formation rate of humic acid,fulvic acid during the humification of leaf litter,there was no significant difference between organs,showing that the deciduous tree species had lower HAC/FAC than evergreen tree species. The present study indicates that stage types,species types,organ types and their interactions have different degrees of influence on HC,HAC,FAC content and HAC/FAC in litter. The results provide a theoretical information and a new direction for further understanding the material circulation process of litterfall in the regional ecosystem.
引文
ABAKUMOV EV,AJTHAML T,BRUS J,et al.,2013.Humus accumulation,humification,and humic acid composition in soils of two post-mining chronosequences after coal mining[J].J Soils Sed,13(3):491-500.
ADAIN F,SPAGNOL M,NIEROP KGJ,2007.Biochemical origin and refractory properties of humic acid extracted from maize plants:The contribution of lignin[J].Biogeochemistry,82(1):55-65.
BERG B,MCCLAUGHERTY C,2008.Plant Litter.Decomposition,Humus Formation,Carbon Sequestration[M].Berlin:Springer-Verlag.
CATOVSKY S,HOLBROOK NM,BAZZAZ FA,2002.Coupling whole-tree transpiration and canopy photosynthesis in coniferous and broad-leaved tree species[J].Can J For Res,32(2):295-309.
COTRUFO MF,WALLENSTEIN MD,BOOT CM,et al.,2013.The microbial efficiency-matrix stabilization(MEMS)framework integrates plant litter decomposition with soil organic matter stabilization:Do labile plant inputs form stable soil organic matter?[J].Glob Change Biol,19(4):988-995.
CHEN CD,2000.The Du jiangyan Region-Pivot sector of assemblage,differentiation and maintenance of biodiversity in northern part of Hengduan Mountain[J].Acta Ecol Sin,20(1):28-34.[陈昌笃,2000.都江堰地区---横断山北段生物多样性交汇、分化和存留的枢纽地段[J].生态学报,20(1):28-34.]
DOU S,YU SQ,ZHANG JJ,2007.Effects of carbon dioxide concentration on humic substances formation in corn stalk decomposition[J].Acta Pedol Sin,44(3):458-466.[窦森,于水强,张晋京,2007.不同CO2浓度对玉米秸秆分解期间土壤腐殖质形成的影响[J].土壤学报,44(3):458-466.]
DOU S,TARDY Y,ZHANG JJ,et al.,2010.The rmodynamic stability of humic acid and fulvic acid in soil and its driving factors[J].Acta Pedol Sin,47(1):71-75.[窦森,YVESTARDY,张晋京,等,2010.土壤胡敏酸与富里酸热力学稳定性及其驱动因素初步研究[J].土壤学报,47(1):71-76.]
DONG M,1997.Survey and analysis of terrestrial biotopes[M].Beijing:China Standard Press.[董鸣,1997.陆地生物群落调查观测与分析[M].北京:中国标准出版社.]
FU C,YANG W,TAN B,et al.,2017.Seasonal dynamics of litterfall in a sub-alpine spruce-fir forest on the eastern Tibetan Plateau:Allometric scaling relationships based on one year of observations[J].Forests,8(9):314.
FANG X,TIAN DL,2006.Study on dynamics of C storage and C storage in Chinese fir plantation[J].Guihaia,26(5):516-522.[方晰,田大伦,2006.杉木人工林C库与C吸存的动态研究[J].广西植物,26(5):516-522.]
GOSZ JR,LIKENS GE,BORMANN FH,1973.Nutrient release from decomposing leaf and branch litter in the Hubbard Brook Forest,New Hampshire[J].Ecol Monogr,43(2):173-191.
GOSZ JR,LIKENS GE,BORMANN FH,1972.Nutrient content of litter fall on the Hubbard Brook experimental forest,New Hampshire[J].Ecology,53(5):769-784.
GE XM,TANG LZ,WANG RH,et al.,2017.Litterfall biomass and decomposition characteristics in a poplar plantation ecosystem[J].Ecol Environ,26(9):1457-1464.[葛晓敏,唐罗忠,王瑞华,等,2017.杨树人工林生态系统凋落物生物量及其分解特征[J].生态环境学报,26(9):1457-1464.]
KLINGE H,RODRIGUES WA,1968.Litter production in an area of Amazonian terra firme forest.Part I.Litter-fall,organic carbon and total nitrogen contents of litter[J].Amazoniana,1(4):287-302.
LIN B,LIU Q,WU Y,et al.,2004.Research progress in forest litters[J].Chin J Ecol,23(1):60-64.[林波,刘庆,吴彦,等,2004.森林凋落物研究进展[J].生态学杂志,23(1):60-64.]
MEIER CE,STANTURF JA,GARDINER ES,2006.Litterfall in the hardwood forest of a minor alluvial-floodplain[J].For Ecol Manag,234(1-3):60-77.
MA ZL,YANG WQ,WU FZ,et al.,2018.Degradation characteristics of leaf litter polyphenols in subtropical evergreen broad-leaved forest in Sichuan Basin during different rainfall periods[J].Acta Ecol Sin,38(9):3078-3085.[马志良,杨万勤,吴福忠,等,2018.四川盆地亚热带常绿阔叶林凋落叶多酚类物质在不同降雨期间的降解特征[J].生态学报,38(9):3078-3085.]
MA ZG,WANG JX,1993.Dynamics of forest litter in the habitat of giant pandas[J].J Plant Ecol,17(2):155-163.[马志贵,王金锡,1993.大熊猫栖息环境的森林凋落物动态研究[J].植物生态学报,17(2):155-163.]
NI XY,YANG WQ,XU LY,et al.,2014.Effects of winter snowpack on accumulation of humic acid and fulvic acid during humification of foliar litters in an alpine forest[J].Acta Pedol Sin,51(5):1138-1152.[倪祥银,杨万勤,徐李亚,等,2014.雪被斑块对高山森林凋落叶腐殖化过程中胡敏酸和富里酸累积的影响[J].土壤学报,51(5):1138-1152.]
NI XY,YANG WQ,TAN B,et al.,2015.Accelerated foliar litter humification in forest gaps:Dual feedbacks of carbon sequestration during winter and the growing season in an alpine forest[J].Geoderma:241-242,136-144.
PONGE JF,2013.Plant-soil feedbacks mediated by humus forms:A review[J].Soil Biol Biochem,57(2):1048-1060.
SUN F,YAN SW,LIN YM,et al.,2017.Distribution characteristics of soil organic carbon storage and carbon density from damaged vegetation under different climate types in Wenchuan earthquake region[J].Guihaia,37(12):1498-1507.[孙凡,严思维,林勇明,等,2017.汶川震区不同气候区受损植被土壤有机碳储量和碳密度分布特征[J].广西植物,37(12):1498-1507.]
SHI GY,LIANG SC,ZENG XB,et al.,2016.Comparative study on seasonal changes in physiological characteristics of Eucalyptus,broad-leaved and coniferous tree species[J].Guihaia,36(7):783-787.[石贵玉,梁士楚,曾小飚,等,2016.桉树与针叶树、阔叶树生理生化指标季节变化的比较[J].广西植物,36(7):783-787.]
WANG WJ,YANG WQ,TAN B,et al.,2013.The dynamics of soil fauna community during litter decomposition at different phenological stages in the subtropical evergreen broad-leaved forests in Sichuan basin[J].Acta Ecol Sin,33(18):5737-5750.[王文君,杨万勤,谭波,等,2013.四川盆地亚热带常绿阔叶林不同物候期凋落物分解与土壤动物群落结构的关系[J].生态学报,33(18):5737-5750.]
WANG Z,CHEN AL,CAO GQ,et al.,2017.Effects of different litter compositions on soil microbial biomass carbon and nitroger contents in Chinese fir plantation[J].J Southern Agric,48(10):1849-1857.[王珍,陈爱玲,曹光球,等,2017.不同凋落物配比对杉木土壤微生物量碳氮的影响[J].南方农业学报,48(10):1849-1857.]
ZHOU SX,HUANG CD,HAN BH,et al.,2017.Simulated nitrogen deposition significantly suppresses the decomposition of forest litter in a natural evergreen broadleaved forest in the rainy area of western China[J].Plant Soil,402(1-2):1-11.
ZHU LG,SUN J,ZHANG WJ,et al.,2017.Effects of pruning on hydrological characteristics of tea tree litter[J].JSouthern Agric,48(10):1795-1801.[朱留刚,孙君,张文锦,等,2017.修剪对茶树修剪凋落物水文特性的影响[J].南方农业学报,48(10):1795-1801.]
ADAIN F,SPAGNOL M,NIEROP KGJ,2007.Biochemical origin and refractory properties of humic acid extracted from maize plants:The contribution of lignin[J].Biogeochemistry,82(1):55-65.
BERG B,MCCLAUGHERTY C,2008.Plant Litter.Decomposition,Humus Formation,Carbon Sequestration[M].Berlin:Springer-Verlag.
CATOVSKY S,HOLBROOK NM,BAZZAZ FA,2002.Coupling whole-tree transpiration and canopy photosynthesis in coniferous and broad-leaved tree species[J].Can J For Res,32(2):295-309.
COTRUFO MF,WALLENSTEIN MD,BOOT CM,et al.,2013.The microbial efficiency-matrix stabilization(MEMS)framework integrates plant litter decomposition with soil organic matter stabilization:Do labile plant inputs form stable soil organic matter?[J].Glob Change Biol,19(4):988-995.
CHEN CD,2000.The Du jiangyan Region-Pivot sector of assemblage,differentiation and maintenance of biodiversity in northern part of Hengduan Mountain[J].Acta Ecol Sin,20(1):28-34.[陈昌笃,2000.都江堰地区---横断山北段生物多样性交汇、分化和存留的枢纽地段[J].生态学报,20(1):28-34.]
DOU S,YU SQ,ZHANG JJ,2007.Effects of carbon dioxide concentration on humic substances formation in corn stalk decomposition[J].Acta Pedol Sin,44(3):458-466.[窦森,于水强,张晋京,2007.不同CO2浓度对玉米秸秆分解期间土壤腐殖质形成的影响[J].土壤学报,44(3):458-466.]
DOU S,TARDY Y,ZHANG JJ,et al.,2010.The rmodynamic stability of humic acid and fulvic acid in soil and its driving factors[J].Acta Pedol Sin,47(1):71-75.[窦森,YVESTARDY,张晋京,等,2010.土壤胡敏酸与富里酸热力学稳定性及其驱动因素初步研究[J].土壤学报,47(1):71-76.]
DONG M,1997.Survey and analysis of terrestrial biotopes[M].Beijing:China Standard Press.[董鸣,1997.陆地生物群落调查观测与分析[M].北京:中国标准出版社.]
FU C,YANG W,TAN B,et al.,2017.Seasonal dynamics of litterfall in a sub-alpine spruce-fir forest on the eastern Tibetan Plateau:Allometric scaling relationships based on one year of observations[J].Forests,8(9):314.
FANG X,TIAN DL,2006.Study on dynamics of C storage and C storage in Chinese fir plantation[J].Guihaia,26(5):516-522.[方晰,田大伦,2006.杉木人工林C库与C吸存的动态研究[J].广西植物,26(5):516-522.]
GOSZ JR,LIKENS GE,BORMANN FH,1973.Nutrient release from decomposing leaf and branch litter in the Hubbard Brook Forest,New Hampshire[J].Ecol Monogr,43(2):173-191.
GOSZ JR,LIKENS GE,BORMANN FH,1972.Nutrient content of litter fall on the Hubbard Brook experimental forest,New Hampshire[J].Ecology,53(5):769-784.
GE XM,TANG LZ,WANG RH,et al.,2017.Litterfall biomass and decomposition characteristics in a poplar plantation ecosystem[J].Ecol Environ,26(9):1457-1464.[葛晓敏,唐罗忠,王瑞华,等,2017.杨树人工林生态系统凋落物生物量及其分解特征[J].生态环境学报,26(9):1457-1464.]
KLINGE H,RODRIGUES WA,1968.Litter production in an area of Amazonian terra firme forest.Part I.Litter-fall,organic carbon and total nitrogen contents of litter[J].Amazoniana,1(4):287-302.
LIN B,LIU Q,WU Y,et al.,2004.Research progress in forest litters[J].Chin J Ecol,23(1):60-64.[林波,刘庆,吴彦,等,2004.森林凋落物研究进展[J].生态学杂志,23(1):60-64.]
MEIER CE,STANTURF JA,GARDINER ES,2006.Litterfall in the hardwood forest of a minor alluvial-floodplain[J].For Ecol Manag,234(1-3):60-77.
MA ZL,YANG WQ,WU FZ,et al.,2018.Degradation characteristics of leaf litter polyphenols in subtropical evergreen broad-leaved forest in Sichuan Basin during different rainfall periods[J].Acta Ecol Sin,38(9):3078-3085.[马志良,杨万勤,吴福忠,等,2018.四川盆地亚热带常绿阔叶林凋落叶多酚类物质在不同降雨期间的降解特征[J].生态学报,38(9):3078-3085.]
MA ZG,WANG JX,1993.Dynamics of forest litter in the habitat of giant pandas[J].J Plant Ecol,17(2):155-163.[马志贵,王金锡,1993.大熊猫栖息环境的森林凋落物动态研究[J].植物生态学报,17(2):155-163.]
NI XY,YANG WQ,XU LY,et al.,2014.Effects of winter snowpack on accumulation of humic acid and fulvic acid during humification of foliar litters in an alpine forest[J].Acta Pedol Sin,51(5):1138-1152.[倪祥银,杨万勤,徐李亚,等,2014.雪被斑块对高山森林凋落叶腐殖化过程中胡敏酸和富里酸累积的影响[J].土壤学报,51(5):1138-1152.]
NI XY,YANG WQ,TAN B,et al.,2015.Accelerated foliar litter humification in forest gaps:Dual feedbacks of carbon sequestration during winter and the growing season in an alpine forest[J].Geoderma:241-242,136-144.
PONGE JF,2013.Plant-soil feedbacks mediated by humus forms:A review[J].Soil Biol Biochem,57(2):1048-1060.
SUN F,YAN SW,LIN YM,et al.,2017.Distribution characteristics of soil organic carbon storage and carbon density from damaged vegetation under different climate types in Wenchuan earthquake region[J].Guihaia,37(12):1498-1507.[孙凡,严思维,林勇明,等,2017.汶川震区不同气候区受损植被土壤有机碳储量和碳密度分布特征[J].广西植物,37(12):1498-1507.]
SHI GY,LIANG SC,ZENG XB,et al.,2016.Comparative study on seasonal changes in physiological characteristics of Eucalyptus,broad-leaved and coniferous tree species[J].Guihaia,36(7):783-787.[石贵玉,梁士楚,曾小飚,等,2016.桉树与针叶树、阔叶树生理生化指标季节变化的比较[J].广西植物,36(7):783-787.]
WANG WJ,YANG WQ,TAN B,et al.,2013.The dynamics of soil fauna community during litter decomposition at different phenological stages in the subtropical evergreen broad-leaved forests in Sichuan basin[J].Acta Ecol Sin,33(18):5737-5750.[王文君,杨万勤,谭波,等,2013.四川盆地亚热带常绿阔叶林不同物候期凋落物分解与土壤动物群落结构的关系[J].生态学报,33(18):5737-5750.]
WANG Z,CHEN AL,CAO GQ,et al.,2017.Effects of different litter compositions on soil microbial biomass carbon and nitroger contents in Chinese fir plantation[J].J Southern Agric,48(10):1849-1857.[王珍,陈爱玲,曹光球,等,2017.不同凋落物配比对杉木土壤微生物量碳氮的影响[J].南方农业学报,48(10):1849-1857.]
ZHOU SX,HUANG CD,HAN BH,et al.,2017.Simulated nitrogen deposition significantly suppresses the decomposition of forest litter in a natural evergreen broadleaved forest in the rainy area of western China[J].Plant Soil,402(1-2):1-11.
ZHU LG,SUN J,ZHANG WJ,et al.,2017.Effects of pruning on hydrological characteristics of tea tree litter[J].JSouthern Agric,48(10):1795-1801.[朱留刚,孙君,张文锦,等,2017.修剪对茶树修剪凋落物水文特性的影响[J].南方农业学报,48(10):1795-1801.]