黄土高原刺槐细根形态特征和生物量研究
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摘要
细根形态反映细根功能,以黄土高原陕西省永寿县15年生刺槐人工林为研究对象,采集刺槐完整根段,依据根序分成1～5级根,对其细根形态结构和生物量进行测定分析,为深入认识刺槐人工林细根形态特征提供参考。结果表明:1)1～2、2～3、3～4、4～5级根的平均分支比分别为4.52、2.06、4.03和4.17。其中,只有2～3级根的分支比与其他分支水平之间具有显著差异(P<0.05)。2)从1级根到5级根,随序级的升高,刺槐细根直径、根长呈升高趋势,而比根长呈降低趋势。各根序之间的细根直径、根长、比根长都具有显著差异(P<0.05)。3)刺槐1～5级细根生物量的排序为1级<2级<3级<5级<4级,且前4级之间的细根生物量具有显著差异(P<0.05)。1～5级细根总生物量为240.95g·m-2,4～5级根生物量之和占总生物量的75.88%,对总生物量有较大的贡献。1～5级细根各分支水平高级根与低级根的细根生物量之间具有显著的线性关系。
Fine root morphology reflects the function of the fine root.This study was conducted in 15-yearold Robinia pseudoacacia plantations of the Loess Plateau,located in Yongshou County,Shaanxi Province,China.Intact fine root segments of R.pseudoacacia were collected by excavation in the plantations.Individual roots were dissected according to the branching order,starting from the distal end of the root system that was numbered as the first order and then increasing sequentially with each branch from the first order to fifth order roots.The fine root morphology and biomass of the first five orders were measured and analyzed.The results showed that:1)the average ratios of branching(Rb)of 1-2,2-3,3-4 and 4-5 levels were 4.52,2.06,4.03 and 4.17 respectively.The ANOVA demonstrated that the Rb of 2-3 level was significantly significant from those of 1-2,3-4 and 4-5 levels(P<0.05).2)From first order to fifth order root,fine root diameter and root length showed increasing trend with the increase of order,while specific root length showed decreasing trend with the increase of order.The root diameter,length and specific root length were significantly different among orders(P<0.05).3)From the first to fifth order,fine root biomass was in an order of 1<2<3<5<4.And the fine root biomass between the first four grades was significantly different(P<0.05).The total fine root biomass of the first to fifth order was 240.95 g·m-2.The fourth to fifth order roots accounted for 75.88%of the total biomass,and had a greater contribution to the total biomass.From the first to fifth order,there was a significant linear relationship between the fine root biomass of lower level and higher level.
Fine root morphology reflects the function of the fine root.This study was conducted in 15-yearold Robinia pseudoacacia plantations of the Loess Plateau,located in Yongshou County,Shaanxi Province,China.Intact fine root segments of R.pseudoacacia were collected by excavation in the plantations.Individual roots were dissected according to the branching order,starting from the distal end of the root system that was numbered as the first order and then increasing sequentially with each branch from the first order to fifth order roots.The fine root morphology and biomass of the first five orders were measured and analyzed.The results showed that:1)the average ratios of branching(Rb)of 1-2,2-3,3-4 and 4-5 levels were 4.52,2.06,4.03 and 4.17 respectively.The ANOVA demonstrated that the Rb of 2-3 level was significantly significant from those of 1-2,3-4 and 4-5 levels(P<0.05).2)From first order to fifth order root,fine root diameter and root length showed increasing trend with the increase of order,while specific root length showed decreasing trend with the increase of order.The root diameter,length and specific root length were significantly different among orders(P<0.05).3)From the first to fifth order,fine root biomass was in an order of 1<2<3<5<4.And the fine root biomass between the first four grades was significantly different(P<0.05).The total fine root biomass of the first to fifth order was 240.95 g·m-2.The fourth to fifth order roots accounted for 75.88%of the total biomass,and had a greater contribution to the total biomass.From the first to fifth order,there was a significant linear relationship between the fine root biomass of lower level and higher level.
引文
[1]王政权,郭大立.根系生态学[J].植物生态学报,2008,32(6):1213-1216.
[2]熊德成,黄锦学,杨智杰,等.亚热带6种树种细根序级结构和形态特征[J].生态学报,2012,32(6):1888-1897.XIONG D C,HUANG J X,YANG Z J,et al.Fine root architecture and morphology among different branch orders of six subtropical tree species[J].Acta Ecologica Sinica,2012,32(6):1888-1897.(in Chinese)
[3]PREGITZER K S,JARED L,DEFOREST,et al.Fine root architecture of nine North American trees[J].Ecological Monographs,2002,72(2):293-309.
[4]GUO D L,MITCHELL R J,HENDRICKS J J.Fine root branch orders respond differentially to carbon source-sink manipulations in a longleaf pine forest[J].Oecologia,2004,140(3):450-457.
[5]卓鸣秀,闫晓俊,熊德成,等.福建建瓯万木林自然保护区8种樟科树种的细根形态特征[J].亚热带资源与环境学报,2018,13(2):22-27.ZHUO M X,YAN X J,XIONG D C,et al.Morphological characteristics of fine roots of 8lauraceae species in Jianou Wanmulin Forest Nature Reserve,Fujian Province[J].Journal of Subtropical Resources and Environment,2018,13(2):22-27.(in Chinese)
[6]王向荣,谷加存,梅莉,等.水曲柳和落叶松细根形态及母根与子根比例关系[J].生态学报,2006,26(6):1686-1692.WANG X R,GU J C,MEI L,et al.Fine root order morphology and proportion between mother roots and daughter roots in Fraxinus mandshurica and Larix gmelinii plantations[J].Acta Ecologica Sinica,2006,26(6):1686-1692.(in Chinese)
[7]师伟,王政权,刘金梁,等.帽儿山天然次生林20个阔叶树种细根形态[J].植物生态学报,2008,32(6):1217-1226.SHI W,WANG Z Q,LIU J L,et al.Fine root morphology of twenty hardwood species in Maoershan natural secondary forest in Northeastern China[J].Journal of Plant Ecology,2008,32(6):1217-1226.(in Chinese)
[8]张良德,徐学选,胡伟,等.黄土丘陵区燕沟流域人工刺槐林的细根空间分布特征[J].林业科学,2011,47(11):31-36.ZHANG L D,XU X X,HU W,et al.Spatial distribution of fine roots of a Robinia pseudoacacia plantation in Yangou watershed in the hilly region of the Loess Plateau[J].Scientia Silvae Sinicae,2011,47(11):31-36.(in Chinese)
[9]张良德,徐学选.延安燕沟流域刺槐根系分布特征[J].西北林学院学报,2011,26(2):9-14.ZHANG L D,XU X X.Distribution characters of robinia pseudoacacia root in Yangou watershed in Yanan[J].Journal of Northwest Forestry University,2011,26(2):9-14.(in Chinese)
[10]白世红,丁新景,马风云,等.黄河三角洲盐碱地人工刺槐混交林细根分布研究[J].中国生态农业学报,2018,26(1):116-124.BAI S H,DING X J,MA F X,et al.Fine root distribution in mixed Robinia pseudoacacia plantations in saline soils of the Yellow River Delta[J].Chinese Journal of Eco-agriculture,2018,26(1):116-124.(in Chinese)
[11]赵忠,成向荣,薛文鹏,等.黄土高原不同水分生态区刺槐细根垂直分布的差异[J].林业科学,2006,42(11):1-7.ZHAO Z,CHENG X R,XUE W P,et al.Difference of fine root vertical distribution of Robinia pseudoacacia under the different climate regions in the Loess Plateau[J].Scientia Silvae Sinicae,2006,42(11):1-7.(in Chinese)
[12]翟明普,蒋三乃,贾黎明.沙地杨树刺槐混交林细根动态[J].北京林业大学学报,2002,24(5-6):39-44.ZHAI M P,JIANG S N,JIA L M.Fine-root dynamics in mixed plantation of poplar and black locust[J].Journal of Beijing Forestry University 2002,24(5-6):39-44.(in Chinese)
[13]王迪海,赵忠,李剑.土壤水分对黄土高原主要造林树种细根表面积季节动态的影响[J].植物生态学报,2010,34(7):819-826.WANG D H,ZHAO Z,LI J.Impact of soil moisture on the seasonal dynamics of fine root surface area of major afforestation tree species on China’s Loess Plateau[J].Journal of Plant Ecology,2010,34(7):819-826.(in Chinese)
[14]蔡倩颖,王迪海.陕北枣树‘七月鲜’细根形态的季节变化[J].西北林学院学报,2017,32(3):30-34.CAI Q Y,WANG D H.Seasonal variations of the fine root morphology of“Qiyuexian”jujube in Northern Shaanxi[J].Journal of Northwest Forestry University,2017,32(3):30-34.(in Chinese)
[15]苏瑾,王迪海.黄土高原刺槐细根形态和生理生化指标季节动态研究[D].杨陵:西北农林科技大学,2016.
[16]李佳梅,朱启良,马璟.华北石质山地麻栎和刺槐混交林浅层细根特征[J].西北林学院学报,2018,33(1):37-42.LI J M,ZHU Q L,MA J.Comparison on fine root traits of Quercus accutissima and Robinia pseudoacaciain lower soil layer in rocky mountainous area of Northern China[J].Journal of Northwest Forestry University,2018,33(1):37-42.(in Chinese)
[17]WANG Z Q,GUO D L,WANG X R,et al.Fine root architecture,morphology,and biomass of different branch orders of two Chinese temperate tree species[J].Plant Soil,2006,288(1/2):155-171.
[18]WEST G B,BROWN J H,ENQUIST B J.A general model for the origin of allometric scaling laws in biology[J].Science,1997,276(5309):122-126.
[19]FITTER A H.Morphometric analysis of root systems:application of the technique and influence of soil fertility on root system development in two herbaceous species[J].Plant,Cell and Environment,1982,5(4):313-322.
[20]LYFORD W H.Rhizography of non-woody roots of trees in the forest floor.[M]//TORREY J G,CLARKSON D T.ed.The Development and Function of Roots.New York:Academic Press,1975:179-196.
[21]HISHI T,TAKEDA H.Dynamics of heterorhizic root systems:protoxylem groups within the fine-root system of Chamaecyparis obtuse[J].New Phytologist,2005,167(2):509-521.
[22]ESSENSTAT D M.On the relationship between specific root length and the rate of root proliferation:a field study using citrus rootstocks[J].New Phytologist,1991,118(1):63-68.
[23]KENRICH P.The Origin of Roots[M].3rd.WAISEL Y,ES-HEL A,KAFKAFI U,eds.Plant Root:the Hidden Half.New York:Baselr.2002:1-15.
[24]常文静,郭大立.中国温带、亚热带和热带森林45个常见树种细根直径变异[J].植物生态学报,2008,32(6):1248-1257.
[25]IDOL T W,POPE P E,PONDER F J.Fine root dynamics across a chronosequence of upland temperate deciduous forests[J].Forest Ecology and Management,2000,127(1/3):153-167.
[26]VOGT K A,VOGT D J,PALMIOTTO P A,et al.Review of root dynamics in forest ecosystems grouped by climate,forest type and species[J].Plant and Soil,1996,187:159-219.
[27]HUANG G,ZHAO X Y,ZHAO H L,et al.Linking root morphology,longevity and function to root branch order:a case study in three shrubs[J].Plant Soil,2010,336(1/2):197-208.
[28]许旸,谷加存,董雪云,等.海南岛4个热带阔叶树种前级细根的形态、解剖结构和组织碳氮含量[J].植物生态学报,2011,35(9):955-964.
[29]王维芳,董薪明,董小枫,等.森林生物量的空间自相关性研究[J].森林工程,2018,34(2):35-39.WANG W F,DONG X M,DONG X F,et al.Study on spatial autocorrelation of forest biomass[J].Forest Engineering,2018,34(2):35-39.(in Chinese)
[30]PERSSON H.The distribution and productivity of fine roots in boreal forests[J].Plant and Soil,1983,71:87-101.
[31]GILL RA,JACKSON R B.Global patterns of root turnover for terrestrial ecosystems[J].New Phytologist,2000,147:13-31.
[2]熊德成,黄锦学,杨智杰,等.亚热带6种树种细根序级结构和形态特征[J].生态学报,2012,32(6):1888-1897.XIONG D C,HUANG J X,YANG Z J,et al.Fine root architecture and morphology among different branch orders of six subtropical tree species[J].Acta Ecologica Sinica,2012,32(6):1888-1897.(in Chinese)
[3]PREGITZER K S,JARED L,DEFOREST,et al.Fine root architecture of nine North American trees[J].Ecological Monographs,2002,72(2):293-309.
[4]GUO D L,MITCHELL R J,HENDRICKS J J.Fine root branch orders respond differentially to carbon source-sink manipulations in a longleaf pine forest[J].Oecologia,2004,140(3):450-457.
[5]卓鸣秀,闫晓俊,熊德成,等.福建建瓯万木林自然保护区8种樟科树种的细根形态特征[J].亚热带资源与环境学报,2018,13(2):22-27.ZHUO M X,YAN X J,XIONG D C,et al.Morphological characteristics of fine roots of 8lauraceae species in Jianou Wanmulin Forest Nature Reserve,Fujian Province[J].Journal of Subtropical Resources and Environment,2018,13(2):22-27.(in Chinese)
[6]王向荣,谷加存,梅莉,等.水曲柳和落叶松细根形态及母根与子根比例关系[J].生态学报,2006,26(6):1686-1692.WANG X R,GU J C,MEI L,et al.Fine root order morphology and proportion between mother roots and daughter roots in Fraxinus mandshurica and Larix gmelinii plantations[J].Acta Ecologica Sinica,2006,26(6):1686-1692.(in Chinese)
[7]师伟,王政权,刘金梁,等.帽儿山天然次生林20个阔叶树种细根形态[J].植物生态学报,2008,32(6):1217-1226.SHI W,WANG Z Q,LIU J L,et al.Fine root morphology of twenty hardwood species in Maoershan natural secondary forest in Northeastern China[J].Journal of Plant Ecology,2008,32(6):1217-1226.(in Chinese)
[8]张良德,徐学选,胡伟,等.黄土丘陵区燕沟流域人工刺槐林的细根空间分布特征[J].林业科学,2011,47(11):31-36.ZHANG L D,XU X X,HU W,et al.Spatial distribution of fine roots of a Robinia pseudoacacia plantation in Yangou watershed in the hilly region of the Loess Plateau[J].Scientia Silvae Sinicae,2011,47(11):31-36.(in Chinese)
[9]张良德,徐学选.延安燕沟流域刺槐根系分布特征[J].西北林学院学报,2011,26(2):9-14.ZHANG L D,XU X X.Distribution characters of robinia pseudoacacia root in Yangou watershed in Yanan[J].Journal of Northwest Forestry University,2011,26(2):9-14.(in Chinese)
[10]白世红,丁新景,马风云,等.黄河三角洲盐碱地人工刺槐混交林细根分布研究[J].中国生态农业学报,2018,26(1):116-124.BAI S H,DING X J,MA F X,et al.Fine root distribution in mixed Robinia pseudoacacia plantations in saline soils of the Yellow River Delta[J].Chinese Journal of Eco-agriculture,2018,26(1):116-124.(in Chinese)
[11]赵忠,成向荣,薛文鹏,等.黄土高原不同水分生态区刺槐细根垂直分布的差异[J].林业科学,2006,42(11):1-7.ZHAO Z,CHENG X R,XUE W P,et al.Difference of fine root vertical distribution of Robinia pseudoacacia under the different climate regions in the Loess Plateau[J].Scientia Silvae Sinicae,2006,42(11):1-7.(in Chinese)
[12]翟明普,蒋三乃,贾黎明.沙地杨树刺槐混交林细根动态[J].北京林业大学学报,2002,24(5-6):39-44.ZHAI M P,JIANG S N,JIA L M.Fine-root dynamics in mixed plantation of poplar and black locust[J].Journal of Beijing Forestry University 2002,24(5-6):39-44.(in Chinese)
[13]王迪海,赵忠,李剑.土壤水分对黄土高原主要造林树种细根表面积季节动态的影响[J].植物生态学报,2010,34(7):819-826.WANG D H,ZHAO Z,LI J.Impact of soil moisture on the seasonal dynamics of fine root surface area of major afforestation tree species on China’s Loess Plateau[J].Journal of Plant Ecology,2010,34(7):819-826.(in Chinese)
[14]蔡倩颖,王迪海.陕北枣树‘七月鲜’细根形态的季节变化[J].西北林学院学报,2017,32(3):30-34.CAI Q Y,WANG D H.Seasonal variations of the fine root morphology of“Qiyuexian”jujube in Northern Shaanxi[J].Journal of Northwest Forestry University,2017,32(3):30-34.(in Chinese)
[15]苏瑾,王迪海.黄土高原刺槐细根形态和生理生化指标季节动态研究[D].杨陵:西北农林科技大学,2016.
[16]李佳梅,朱启良,马璟.华北石质山地麻栎和刺槐混交林浅层细根特征[J].西北林学院学报,2018,33(1):37-42.LI J M,ZHU Q L,MA J.Comparison on fine root traits of Quercus accutissima and Robinia pseudoacaciain lower soil layer in rocky mountainous area of Northern China[J].Journal of Northwest Forestry University,2018,33(1):37-42.(in Chinese)
[17]WANG Z Q,GUO D L,WANG X R,et al.Fine root architecture,morphology,and biomass of different branch orders of two Chinese temperate tree species[J].Plant Soil,2006,288(1/2):155-171.
[18]WEST G B,BROWN J H,ENQUIST B J.A general model for the origin of allometric scaling laws in biology[J].Science,1997,276(5309):122-126.
[19]FITTER A H.Morphometric analysis of root systems:application of the technique and influence of soil fertility on root system development in two herbaceous species[J].Plant,Cell and Environment,1982,5(4):313-322.
[20]LYFORD W H.Rhizography of non-woody roots of trees in the forest floor.[M]//TORREY J G,CLARKSON D T.ed.The Development and Function of Roots.New York:Academic Press,1975:179-196.
[21]HISHI T,TAKEDA H.Dynamics of heterorhizic root systems:protoxylem groups within the fine-root system of Chamaecyparis obtuse[J].New Phytologist,2005,167(2):509-521.
[22]ESSENSTAT D M.On the relationship between specific root length and the rate of root proliferation:a field study using citrus rootstocks[J].New Phytologist,1991,118(1):63-68.
[23]KENRICH P.The Origin of Roots[M].3rd.WAISEL Y,ES-HEL A,KAFKAFI U,eds.Plant Root:the Hidden Half.New York:Baselr.2002:1-15.
[24]常文静,郭大立.中国温带、亚热带和热带森林45个常见树种细根直径变异[J].植物生态学报,2008,32(6):1248-1257.
[25]IDOL T W,POPE P E,PONDER F J.Fine root dynamics across a chronosequence of upland temperate deciduous forests[J].Forest Ecology and Management,2000,127(1/3):153-167.
[26]VOGT K A,VOGT D J,PALMIOTTO P A,et al.Review of root dynamics in forest ecosystems grouped by climate,forest type and species[J].Plant and Soil,1996,187:159-219.
[27]HUANG G,ZHAO X Y,ZHAO H L,et al.Linking root morphology,longevity and function to root branch order:a case study in three shrubs[J].Plant Soil,2010,336(1/2):197-208.
[28]许旸,谷加存,董雪云,等.海南岛4个热带阔叶树种前级细根的形态、解剖结构和组织碳氮含量[J].植物生态学报,2011,35(9):955-964.
[29]王维芳,董薪明,董小枫,等.森林生物量的空间自相关性研究[J].森林工程,2018,34(2):35-39.WANG W F,DONG X M,DONG X F,et al.Study on spatial autocorrelation of forest biomass[J].Forest Engineering,2018,34(2):35-39.(in Chinese)
[30]PERSSON H.The distribution and productivity of fine roots in boreal forests[J].Plant and Soil,1983,71:87-101.
[31]GILL RA,JACKSON R B.Global patterns of root turnover for terrestrial ecosystems[J].New Phytologist,2000,147:13-31.
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