黄土区辽东栎群落细根生物量对物种多样性及气候的响应
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摘要
以陕西黄龙、甘肃子午岭、宁夏六盘山和青海孟达辽东栎(Quercus liaotungensis)次生中龄林为研究对象,采用根钻取土芯法收集细根,分析群落细根生物量分布的差异,以及与群落物种多样性和气候的相互关系。结果表明,黄土区辽东栎分布中心的黄龙和子午岭地区群落细根生物量显著高于分布边缘的六盘山和孟达地区(P<0.05);群落细根主要集中在0~20cm表层土壤中,占细根总生物量的60%~80%。黄龙地区细根生物量最大(821.58g·m~(-2))、孟达地区细根生物量最小(495.30g·m~(-2))。分析还表明,辽东栎林0~20cm活细根生物量、0~20cm死细根生物量与群落丰富度指数、Shannon-Wiener多样性指数、Simpson多样性指数、Pielou均匀度指数和年均温呈正相关;与较高的降雨量、海拔呈负相关。不同地区辽东栎群落细根分布的差异表明群落地下生物量分配模式不同,需要在今后研究中深入探索分配差异的机理,为物种多样性影响生态系统碳循环的激发效应提供基础。
Fine roots(diameter≤2 mm)contribute significantly to carbon cycling in forest ecosystems because they compromise a large fraction of annual net primary production.However,effects of both species diversity and environmental gradients on fine root biomass have less been investigated.In this contribution,a set of 12 sample plots were conducted in middle-aged Quercus liaotungensis forest in 4 regions of Huanglong Mountains(Shaanxi),Ziwuling Mountains(Gansu),Liupanshan Mountains(Ningxia)and Mengda Nature Reserve(Qinghai)across the Loess region.Fine roots were collected using a cylindrical(9 cm diameter)soil corer in each sample plot.The vertical distribution of fine root biomass in different sites and their relationships with community diversity and environmental factors were analyzed.The results showed that the total fine root biomass of the central distribution area in Huanglong Mountains and Ziwuling Mountains were significantly higher than those in the marginal distribution area in Liupanshan Mountains and Mengda(P<0.05).Fine root biomass was mainly distributed in 0-20 cm soil layers,which accountedfor 60%-80% of the total fine root biomass.The highest fine root biomass was existed in Huanglong Mountains(821.58 g·m~(-2))while the lowest was in Mengda(495.30 g·m~(-2)).Redundancy analysis showed that there was a significant relationship between species richness,Shannon-Wiener index,Simpson index,Pielou index and 0-20 cm fine root biomass and necromass,whereas a negative relationship between precipitation,elevation and 0-20 cm fine root biomass.We concluded that patterns of belowground biomass allocation differed from regions,more attention should be paid to the mechanism about the effects of species diversity on the priming effect in forest ecosystem carbon cycling.
Fine roots(diameter≤2 mm)contribute significantly to carbon cycling in forest ecosystems because they compromise a large fraction of annual net primary production.However,effects of both species diversity and environmental gradients on fine root biomass have less been investigated.In this contribution,a set of 12 sample plots were conducted in middle-aged Quercus liaotungensis forest in 4 regions of Huanglong Mountains(Shaanxi),Ziwuling Mountains(Gansu),Liupanshan Mountains(Ningxia)and Mengda Nature Reserve(Qinghai)across the Loess region.Fine roots were collected using a cylindrical(9 cm diameter)soil corer in each sample plot.The vertical distribution of fine root biomass in different sites and their relationships with community diversity and environmental factors were analyzed.The results showed that the total fine root biomass of the central distribution area in Huanglong Mountains and Ziwuling Mountains were significantly higher than those in the marginal distribution area in Liupanshan Mountains and Mengda(P<0.05).Fine root biomass was mainly distributed in 0-20 cm soil layers,which accountedfor 60%-80% of the total fine root biomass.The highest fine root biomass was existed in Huanglong Mountains(821.58 g·m~(-2))while the lowest was in Mengda(495.30 g·m~(-2)).Redundancy analysis showed that there was a significant relationship between species richness,Shannon-Wiener index,Simpson index,Pielou index and 0-20 cm fine root biomass and necromass,whereas a negative relationship between precipitation,elevation and 0-20 cm fine root biomass.We concluded that patterns of belowground biomass allocation differed from regions,more attention should be paid to the mechanism about the effects of species diversity on the priming effect in forest ecosystem carbon cycling.
引文
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[3] PREGITZER,KURT S.Fine roots of trees–a new perspective[J].New Phytologist,2002,154(2):267-270.
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[6]荐圣淇,赵传燕,方书敏,等.陇中黄土高原主要造林树种细根生物量分布[J].应用生态学报,2014,25(7):1905-1911.JIAN S Q,ZHAO C Y,FANG S M,et al.Distribution of fine root biomass of main planting tree species in Loess Plateau,China[J].Chinese Journal of Applied Ecology,2014.25(7):1905-1911.(in Chinese)
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[8] LI J,JIANG X M,YIN H J,et al.Root exudates and soil microbes in three Picea asperata plantations with different stand ages[J].The Journal of Applied Ecology,2014,25(2):325-332.
[9] YUAN Z Y,CHEN H Y H.Fine root dynamics with stand development in the boreal forest[J].Functional Ecology,2012,26(4):991-998.
[10] LIAO Y,MCCORMACK M L,FAN H,et al.Relation of fine root distribution to soil C in a Cunninghamia lanceolata plantation in subtropical China[J].Plant and Soil,2014,381(1/2):225-234.
[11] ASAYE Z,ZEWDIE S.Fine root dynamics and soil carbon accretion under thinned and un-thinned Cupressus lusitanica stands in,southern Ethiopia[J].Plant and Soil,2013,366(1/2):261-271.
[12]杨勤业,张伯平,郑度.关于黄土高原空间范围的讨论[J].自然资源学报,1988,3(1):9-15.YANG Q Y,ZHANG B P,ZHENG D.On the boundary of the Loess Plateau[J].Journal of Natural Resources,1988.3(1):9-15.(in Chinese)
[13] ZHU X,LI Y,PENG X,et al.Soils of the Loess region in China[J].Geoderma,1983,29(3):237-255.
[14]刘东生.黄土与环境[M].北京:科学出版社,1985.
[15] YU X,ZHOU H,LUO T.Spatial and temporal variations in insect‐infested acorn fall in a Quercus liaotungensis forest in north China[J].Ecological Research,2003,18(2):155-164.
[16]康永祥.黄土高原辽东栎林群落生态研究[D].陕西杨陵:西北农林科技大学,2012.
[17]邓磊,张文辉,何景峰,等.不同采伐强度对辽东栎林幼苗更新的影响[J].西北林学院学报,2011.26(2):160-166.DENG L,ZHANG W H,HE J F,et al.Effects of different cutting intensity on seedling regeneration of Quercus liaotungensis[J].Journal of Northwest Forestry University,2011.26(2):160-166.(in Chinese)
[18]张仰渠.陕西森林[M].北京:中国林业出版社,1989.
[19] DENG L,ZHANG W H,GUAN J H.Seed rain and community diversity of Liaotung oak(Quercus liaotungensis Koidz)in Shaanxi,northwest China[J].Ecological Engineering,2014,67:104-111.
[20] MARTEINSDOTTIR B.Seed rain and seed bank reveal that seed limitation strongly influences plant community assembly in grasslands[J].Plos One,2014,9(7):103352.
[21] DU Y,MI X,LIU X,et al.The effects of ice storm on seed rain and seed limitation in an evergreen broad-leaved forest in east China[J].Acta Oecologica,2012,39:87-93.
[22] GALINDEZ G,ORTEGA-BAES P,SCOPEL A L,et al.The dynamics of three shrub species in a fire-prone temperate savanna:the interplay between the seed bank,seed rain and fire regime[J].Plant Ecology,2013,214(1):75-86.
[23]董瑞.黄土高原辽东栎种群动态与生殖生态[D].陕西杨陵:西北农林科技大学,2008.
[24]关晋宏.甘肃森林生态系统固碳特征及与物种多样性关系研究[D].北京:中国科学院,2015.
[25]马克平,刘玉明.生物群落多样性的测度方法[J].生物多样性,1994,2(4):231-239.MA K P,LIU Y M.Measurement methods of biological diversity[J].Chinese Biodiversity,1994.2(4):231-239.(in Chinese)
[26]贺金生,陈伟烈.中国中亚热带东部常绿阔叶林主要类型的群落多样性特征[J].植物生态学报,1998,22(4):303-311.HE J S,CHEN W L.Community diversity of the main types of the evergreen broad-leaved forest in the eastern part of the middle subtropical China[J].Acta Phytoecologica Sinica,1998.22(4):303-311.(in Chinese)
[27]单建平,陶大立,王淼,等.长白山阔叶红松林细根周转的研究[J].应用生态学报,1993,4(3):241-245.SHAN J P,THAO D L,WANG M,et al.Fine roots turnover in a broad-leaved Korean pine forest of Changbai Mountain[J].Chinese Journal of Applied Ecology,1993,4(3):241-245.(in Chinese)
[28]耿鹏飞,金光泽.小兴安岭4种森林类型细根生物量的时空格局[J].林业科学,2016,52(6):140-148.GENG P F,JIN G Z.Spatial and temporal patterns of fine root biomass in four forest types in Xiaoxing'an Mountains[J].Scientia Silvae Sinicae,2016,52(6):140-148.(in Chinese)
[29] ZHOU Y,SU J,JANSSENS I A,et al.Fine root and litterfall dynamics of three Korean pine(Pinus koraiensis)forests along an altitudinal gradient[J].Plant and Soil,2014,374(1/2):19-32.
[30]盛浩,周萍,李洁,等.中亚热带山区深层土壤有机碳库对土地利用变化的响应[J].生态学报,2014,34(23):7004-7012.SHENG H,ZHOU P,LI J,et al.Response of deep soil organic carbon storage to land-use changes in subtropical Hilly Region of China[J].Acta Ecologica Sinica,2014.34(23):7004-7012.(in Chinese)
[31]董慧霞,李贤伟,张健,等.退耕地三倍体毛白杨林地细根生物量及其与土壤水稳性团聚体的关系[J].林业科学,2007,43(5):24-29.DONG H X,LI X W,ZHANG J,et al.Biomass of fine root and its relationship with water_stable aggregate in compound lands of triploid Populus tomentosa on lands converted from agricultural lands[J].Scientia Silvae Sinicae,2007.43(5):24-29.(in Chinese)
[32]关晋宏,杜盛,程积民,等.甘肃省森林碳储量现状与固碳速率[J].植物生态学报,2016,40(4):304-317.GUAN J H,DU S,CHENG J M,et al.Current stocks and rate of sequestration of forest carbon in Gansu Province,China[J].Chinese Journal of Plant Ecology,2016.40(4):304-317.(in Chinese)
[33] LIU C,XIANG W,LEI P,et al.Standing fine root mass and production in four Chinese subtropical forests along a succession and species diversity gradient[J].Plant and Soil,2013,376(1/2):445-459.
[34] LAVOREL S,GARNIER E.Predicting changes in community composition and ecosystem functioning from plant traits:revisiting the Holy Grail[J].Functional Ecology,2002,16(5):545-556.
[35] KERN C C,MONTGOMERY R A,REICH P B,et al.Harvest-created canopy gaps increase species and functional trait diversity of the forest ground-layer community[J].Forest Science,2014,60(2):335-344.
[36] CORNWELL W K,ACKERLY D D.Community assembly and shifts in plant trait distributions across an environmental gradient in coastal California[J].Ecological Monographs,2009,79(1):109-126.
[37] SWENSON N G.Partitioning the phylogenetic,functional,environmental,and spatial components of community diversity[J].Functional and Phylogenetic Ecology in R:Springer,2014:173-187.
[38] WESTOBY M,WRIGHT I J.Land-plant ecology on the basis of functional traits[J].Trends in Ecology&Evolution,2006,21(5):261-268.
[39]张小鹏,王得祥,张鹏,等.抚育间伐对小陇山林区华山松林下植物多样性的影响[J].西北林学院学报,2017.32(2):37-42.ZHANG X P,WANG D X,ZHANG P,et al.Effects of thinning on the diversity of undergrowth of Pinus armandii plantation in Xiaolong Mountainous region[J].Journal of Northwest Forestry University,2017.32(2):37-42.(in Chinese)
[40] LEI P,SCHERER-LORENZEN M,BAUHUS J.The effect of tree species diversity on fine-root production in a young temperate forest[J].Oecologia,2012,169(4):1105-1115.
[41] BRASSARD B W,CHEN H Y H,CAVARD X,et al.Tree species diversity increases fine root productivity through increased soil volume filling[J].Journal of Ecology,2013,101(1):210-219.
[2] EISSENSTAT D M,YANAI R D.The ecology of root lifespan[J].Advances in Ecological Research,1997,27:1-60.
[3] PREGITZER,KURT S.Fine roots of trees–a new perspective[J].New Phytologist,2002,154(2):267-270.
[4] GUO D,MITCHELL R J,WITHINGTON J M,et al..Endogenous and exogenous controls of root life span,mortality and nitrogen flux in a longleaf pine forest:root branch order predominates[J].Journal of Ecology,2008,96(4):737-745.
[5] BENNETT J N,ANDREW B,PRESCOTT C E.Vertical fine root distributions of western redcedar,western hemlock,and salal in old-growth cedar-hemlock forests on northern Vancouver island[J].Canadian Journal of Forest Research,2002,32(7):1208-1216.
[6]荐圣淇,赵传燕,方书敏,等.陇中黄土高原主要造林树种细根生物量分布[J].应用生态学报,2014,25(7):1905-1911.JIAN S Q,ZHAO C Y,FANG S M,et al.Distribution of fine root biomass of main planting tree species in Loess Plateau,China[J].Chinese Journal of Applied Ecology,2014.25(7):1905-1911.(in Chinese)
[7]关晋宏,张克栋,邓磊,等.不同起源云杉林土壤有机碳密度及与物种多样性关系[J].水土保持研究,2016,23(2):49-54.GUAN J H,ZHANG K D,DENG L,et al.Soil organic carbon density and relationship with species diversity in pedigreed spruce at natural forest and plantation scales[J].Research of Soil and Water Conservation,2016.23(2):49-54.(in Chinese)
[8] LI J,JIANG X M,YIN H J,et al.Root exudates and soil microbes in three Picea asperata plantations with different stand ages[J].The Journal of Applied Ecology,2014,25(2):325-332.
[9] YUAN Z Y,CHEN H Y H.Fine root dynamics with stand development in the boreal forest[J].Functional Ecology,2012,26(4):991-998.
[10] LIAO Y,MCCORMACK M L,FAN H,et al.Relation of fine root distribution to soil C in a Cunninghamia lanceolata plantation in subtropical China[J].Plant and Soil,2014,381(1/2):225-234.
[11] ASAYE Z,ZEWDIE S.Fine root dynamics and soil carbon accretion under thinned and un-thinned Cupressus lusitanica stands in,southern Ethiopia[J].Plant and Soil,2013,366(1/2):261-271.
[12]杨勤业,张伯平,郑度.关于黄土高原空间范围的讨论[J].自然资源学报,1988,3(1):9-15.YANG Q Y,ZHANG B P,ZHENG D.On the boundary of the Loess Plateau[J].Journal of Natural Resources,1988.3(1):9-15.(in Chinese)
[13] ZHU X,LI Y,PENG X,et al.Soils of the Loess region in China[J].Geoderma,1983,29(3):237-255.
[14]刘东生.黄土与环境[M].北京:科学出版社,1985.
[15] YU X,ZHOU H,LUO T.Spatial and temporal variations in insect‐infested acorn fall in a Quercus liaotungensis forest in north China[J].Ecological Research,2003,18(2):155-164.
[16]康永祥.黄土高原辽东栎林群落生态研究[D].陕西杨陵:西北农林科技大学,2012.
[17]邓磊,张文辉,何景峰,等.不同采伐强度对辽东栎林幼苗更新的影响[J].西北林学院学报,2011.26(2):160-166.DENG L,ZHANG W H,HE J F,et al.Effects of different cutting intensity on seedling regeneration of Quercus liaotungensis[J].Journal of Northwest Forestry University,2011.26(2):160-166.(in Chinese)
[18]张仰渠.陕西森林[M].北京:中国林业出版社,1989.
[19] DENG L,ZHANG W H,GUAN J H.Seed rain and community diversity of Liaotung oak(Quercus liaotungensis Koidz)in Shaanxi,northwest China[J].Ecological Engineering,2014,67:104-111.
[20] MARTEINSDOTTIR B.Seed rain and seed bank reveal that seed limitation strongly influences plant community assembly in grasslands[J].Plos One,2014,9(7):103352.
[21] DU Y,MI X,LIU X,et al.The effects of ice storm on seed rain and seed limitation in an evergreen broad-leaved forest in east China[J].Acta Oecologica,2012,39:87-93.
[22] GALINDEZ G,ORTEGA-BAES P,SCOPEL A L,et al.The dynamics of three shrub species in a fire-prone temperate savanna:the interplay between the seed bank,seed rain and fire regime[J].Plant Ecology,2013,214(1):75-86.
[23]董瑞.黄土高原辽东栎种群动态与生殖生态[D].陕西杨陵:西北农林科技大学,2008.
[24]关晋宏.甘肃森林生态系统固碳特征及与物种多样性关系研究[D].北京:中国科学院,2015.
[25]马克平,刘玉明.生物群落多样性的测度方法[J].生物多样性,1994,2(4):231-239.MA K P,LIU Y M.Measurement methods of biological diversity[J].Chinese Biodiversity,1994.2(4):231-239.(in Chinese)
[26]贺金生,陈伟烈.中国中亚热带东部常绿阔叶林主要类型的群落多样性特征[J].植物生态学报,1998,22(4):303-311.HE J S,CHEN W L.Community diversity of the main types of the evergreen broad-leaved forest in the eastern part of the middle subtropical China[J].Acta Phytoecologica Sinica,1998.22(4):303-311.(in Chinese)
[27]单建平,陶大立,王淼,等.长白山阔叶红松林细根周转的研究[J].应用生态学报,1993,4(3):241-245.SHAN J P,THAO D L,WANG M,et al.Fine roots turnover in a broad-leaved Korean pine forest of Changbai Mountain[J].Chinese Journal of Applied Ecology,1993,4(3):241-245.(in Chinese)
[28]耿鹏飞,金光泽.小兴安岭4种森林类型细根生物量的时空格局[J].林业科学,2016,52(6):140-148.GENG P F,JIN G Z.Spatial and temporal patterns of fine root biomass in four forest types in Xiaoxing'an Mountains[J].Scientia Silvae Sinicae,2016,52(6):140-148.(in Chinese)
[29] ZHOU Y,SU J,JANSSENS I A,et al.Fine root and litterfall dynamics of three Korean pine(Pinus koraiensis)forests along an altitudinal gradient[J].Plant and Soil,2014,374(1/2):19-32.
[30]盛浩,周萍,李洁,等.中亚热带山区深层土壤有机碳库对土地利用变化的响应[J].生态学报,2014,34(23):7004-7012.SHENG H,ZHOU P,LI J,et al.Response of deep soil organic carbon storage to land-use changes in subtropical Hilly Region of China[J].Acta Ecologica Sinica,2014.34(23):7004-7012.(in Chinese)
[31]董慧霞,李贤伟,张健,等.退耕地三倍体毛白杨林地细根生物量及其与土壤水稳性团聚体的关系[J].林业科学,2007,43(5):24-29.DONG H X,LI X W,ZHANG J,et al.Biomass of fine root and its relationship with water_stable aggregate in compound lands of triploid Populus tomentosa on lands converted from agricultural lands[J].Scientia Silvae Sinicae,2007.43(5):24-29.(in Chinese)
[32]关晋宏,杜盛,程积民,等.甘肃省森林碳储量现状与固碳速率[J].植物生态学报,2016,40(4):304-317.GUAN J H,DU S,CHENG J M,et al.Current stocks and rate of sequestration of forest carbon in Gansu Province,China[J].Chinese Journal of Plant Ecology,2016.40(4):304-317.(in Chinese)
[33] LIU C,XIANG W,LEI P,et al.Standing fine root mass and production in four Chinese subtropical forests along a succession and species diversity gradient[J].Plant and Soil,2013,376(1/2):445-459.
[34] LAVOREL S,GARNIER E.Predicting changes in community composition and ecosystem functioning from plant traits:revisiting the Holy Grail[J].Functional Ecology,2002,16(5):545-556.
[35] KERN C C,MONTGOMERY R A,REICH P B,et al.Harvest-created canopy gaps increase species and functional trait diversity of the forest ground-layer community[J].Forest Science,2014,60(2):335-344.
[36] CORNWELL W K,ACKERLY D D.Community assembly and shifts in plant trait distributions across an environmental gradient in coastal California[J].Ecological Monographs,2009,79(1):109-126.
[37] SWENSON N G.Partitioning the phylogenetic,functional,environmental,and spatial components of community diversity[J].Functional and Phylogenetic Ecology in R:Springer,2014:173-187.
[38] WESTOBY M,WRIGHT I J.Land-plant ecology on the basis of functional traits[J].Trends in Ecology&Evolution,2006,21(5):261-268.
[39]张小鹏,王得祥,张鹏,等.抚育间伐对小陇山林区华山松林下植物多样性的影响[J].西北林学院学报,2017.32(2):37-42.ZHANG X P,WANG D X,ZHANG P,et al.Effects of thinning on the diversity of undergrowth of Pinus armandii plantation in Xiaolong Mountainous region[J].Journal of Northwest Forestry University,2017.32(2):37-42.(in Chinese)
[40] LEI P,SCHERER-LORENZEN M,BAUHUS J.The effect of tree species diversity on fine-root production in a young temperate forest[J].Oecologia,2012,169(4):1105-1115.
[41] BRASSARD B W,CHEN H Y H,CAVARD X,et al.Tree species diversity increases fine root productivity through increased soil volume filling[J].Journal of Ecology,2013,101(1):210-219.