青杄功能性状及其在各器官间分配格局的时空变化
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摘要
为研究青杄在不同环境和发育阶段的适应性和生存策略,以关帝山典型森林分布区的青杄(Picea wilsonii)为对象,分析其叶片鲜干比、叶面积、叶绿素和C、N、P含量等重要功能性状在不同年龄(5、15、25、35、45 a)和不同海拔梯度(1 760、1 820、1 880、1 940、2 000 m)上的变化,以及C、N、P在各器官间分配格局随年龄和海拔的时空变化。结果表明:1)随树龄的增加,青杄叶面积、叶鲜干比呈增大趋势,叶P和根P含量呈下降趋势,叶有机碳含量下调,但根有机碳在上调。2)随海拔增加,叶绿素、叶面积显著增加,叶有机碳呈下降趋势,而根有机碳在增加。叶、枝、根中的N、P含量均呈先增加后减小趋势,相比之下,N、P随海拔的变化较有机碳更为显著。3)随树龄增加,C、N、P在各器官间分配格局表现为:P在幼年时于叶、根中分配比例相对较大,成年时主要集中在枝中。在研究的5～45 a范围内,N分配呈叶>枝>根的稳定格局,未随年龄而显著变化。有机碳则在各器官呈近乎均匀分配的基础上随年龄增加根中比例略有增大,相反叶中比例略有下降。C、N、P并未随海拔增加呈现各器官间分配格局上的显著变化。青杄各功能性状随年龄和海拔变化并通过在各器官间相互关联和协调分配来增强其生态适应性。
In order to study the adaptability and survival strategies of Picea wilsonii in different environments and development stages,P.wilsonii forest occurring in Guandi Mountain,a typical forest distribution area was selected as the research object to investigate the variations of some important functional traits,including leaf fresh-to-dry ratio,leaf area,contents of chlorophyll,carbon,nitrogen,and phosphorus at different ages(5,15,25,35,and 45 a) and different altitudes(1 760,1 820,1 880,1 940,and 2 000 m),and the spatial and temporal variations of distribution pattern of carbon,nitrogen and phosphorus in various organs with the changes of age and altitude.The results showed that 1) with the increasie of tree age,leaf area and leaf fresh-to-dry ratio presented an upward trend,while the contents of leaf phosphorus and root phosphorus were in a downward trend,leaf organic carbon tended to decrease,while root organic carbon tended to increase.2) With the increase of altitude,chlorophyll content and leaf area increased significantly,leaf organic carbon showed a downward trend,but root organic carbon increased.The nitrogen and phosphorus in the leaves,branches and roots increased first and then decreased.In contrast,the changes of nitrogen and phosphorus were more significant than those of organic carbon as altitude changed.3) With the increase of tree age,the distribution patterns of carbon,nitrogen and phosphorus in various organs showed that phosphorus accounted for a relatively large proportion in the leaves and roots at an early age,and distributed mostly in branches in adulthood.At the age of 5 to 45 a,the nitrogen was stable in the distribution pattern as leaf>branch>root and it did not change significantly with the change of tree age.With the increase of tree age,the organic carbon that distributed uniformly in various organs before increased a little bit in the roots,while decreased slightly in the leaves.Besides,carbon,nitrogen and phosphorus did not show significant changes in the distribution in various organs as altitude increased.The functional traits of the P.wilsonii were related to each other as age and altitude varied,which would enhance the ecological adaptability by coordination distribution.
In order to study the adaptability and survival strategies of Picea wilsonii in different environments and development stages,P.wilsonii forest occurring in Guandi Mountain,a typical forest distribution area was selected as the research object to investigate the variations of some important functional traits,including leaf fresh-to-dry ratio,leaf area,contents of chlorophyll,carbon,nitrogen,and phosphorus at different ages(5,15,25,35,and 45 a) and different altitudes(1 760,1 820,1 880,1 940,and 2 000 m),and the spatial and temporal variations of distribution pattern of carbon,nitrogen and phosphorus in various organs with the changes of age and altitude.The results showed that 1) with the increasie of tree age,leaf area and leaf fresh-to-dry ratio presented an upward trend,while the contents of leaf phosphorus and root phosphorus were in a downward trend,leaf organic carbon tended to decrease,while root organic carbon tended to increase.2) With the increase of altitude,chlorophyll content and leaf area increased significantly,leaf organic carbon showed a downward trend,but root organic carbon increased.The nitrogen and phosphorus in the leaves,branches and roots increased first and then decreased.In contrast,the changes of nitrogen and phosphorus were more significant than those of organic carbon as altitude changed.3) With the increase of tree age,the distribution patterns of carbon,nitrogen and phosphorus in various organs showed that phosphorus accounted for a relatively large proportion in the leaves and roots at an early age,and distributed mostly in branches in adulthood.At the age of 5 to 45 a,the nitrogen was stable in the distribution pattern as leaf>branch>root and it did not change significantly with the change of tree age.With the increase of tree age,the organic carbon that distributed uniformly in various organs before increased a little bit in the roots,while decreased slightly in the leaves.Besides,carbon,nitrogen and phosphorus did not show significant changes in the distribution in various organs as altitude increased.The functional traits of the P.wilsonii were related to each other as age and altitude varied,which would enhance the ecological adaptability by coordination distribution.
引文
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[17] 王海珍,韩路,徐雅丽,等.土壤水分梯度对灰胡杨光合作用与抗逆性的影响[J].生态学报,2017,37(2):432-442.WANG H Z,HAN L,XU Y L,et al.Effects of soil water gradient on photosynthetic characteristics and stress resistanc of Populus pruinosa in the tarim basin,China[J].Acta Ecologica Sinica,2017,37(2):432-442.(in Chinese)
[18] 杨士梭,温仲明,苗连朋,等.黄土丘陵区植物功能性状对微地形变化的响应[J].应用生态学报,2014,25(12):3413-3419.
[19] 谢锦,常顺利,张毓涛,等.天山北坡植物土壤生态化学计量特征的垂直地带性[J].生态学报,2016,36(14):4363-4372.XIE J,CHANG S L,ZHANG Y T,et al.Plant and soil ecological stoichiometry with vertical zonality on the northern slope ofthe middle Tianshan Mountains[J].Acta Ecologica Sinica,2016,36(14):4363-4372.(in Chinese)
[20] 温培才,盛茂银,王霖娇,等.西南喀斯特高原盆地石漠化环境植物群落结构与物种多样性时空动态[J].广西植物,2018,38(1):11-23.WEN P C,SHENG M Y,WANG L J,et al.Temporal-spatial dynamics for plant community structure and species diversity of rocky desertification ecosystem in karst plateau basin,southwest China [J].Guihaia,2018,38(1):11-23.(in Chinese)
[21] 张巧明,王得祥,龚明贵,等.秦岭火地塘林区不同海拔森林土壤理化性质[J].水土保持学报,2011,25(5):69-73.
[22] 孙娇,赵发珠,韩新辉,等.不同林龄刺槐林土壤团聚体化学计量特征及其与土壤养分的关系[J].生态学报,2016,36(21):6879-6888.SUN J,ZHAO F Z,HAN X H,et al.Ecological stoichiometry of soil aggregates and relationship with soil nutrients of different-aged Robinia pseudoacacia forests[J].Acta Ecologica Sinica,2016,36(21):6879-6888.(in Chinese)
[23] 杨秀清,史婵,王旭刚,等.关帝山云杉次生林土壤的空间异质性及其与地形相关性[J].中国水土保持科学,2017,15(4):16-24.
[24] 韦兰英,上官周平.黄土高原白羊草、沙棘和辽东栎细根比根长特性[J].生态学报,2006,26(12):4164-4170.
[25] 袁帅.长白内山区不同森林群落中主要树种植物功能性状与固碳能力初探[D].长春:东北师范大学,2011.
[2] 段媛媛,宋丽娟,牛素旗,等.不同林龄刺槐叶功能性状差异及其与土壤养分的关系[J].应用生态学报,2017,28(1):28-36.DUAN Y Y,SONG L J,NIU S Q,et al.Variation in leaf functional traits of different-aged Robinia pseudoacacia communitie and relationships with soil nutrients[J].Chinese Journal of Applied Ecology,2017,28(1):28-36 (in Chinese)
[3] 冯秋红,史作民,董莉莉.植物功能性状对环境的响应及其应用[J].林业科学,2008,44(4):125-131
[4] 谢益君.广西大明山常绿阔叶林20种优势植物的功能性状特征[D].南宁:广西大学,2013.
[5] COMELISSEN J H C,LAVOREL S,GAMIER E,et al.A handbook of protocols for standardized and easy measurement of plant functional traits worldwide[J].Australian Journal of Botany,2003,51:335-380.
[6] WRIGHT I J,REICH P B,WESTOBY M,et al.The worldwide leaf economics spectrum[J].Nature,2004,428(6985):821-827.
[7] 龚时慧,温仲明,施宇.延河流域植物群落功能性状对环境梯度的响应[J].生态学报,2011,31(20):6088-6097.GONG S H,WEN Z M,SHI Y.The response of community-weighted mean plant functional traits to environmental gradients in Yanhe river catchment[J].Acta Ecologica Sinica,2011,31(20):6088-6097.(in Chinese)
[8] 沈震,武玉荷,党鹏,等.黄土高原丘陵沟壑区2种起源油松林更新苗根际土壤酶活性和养分含量比较研究[J].西北林学院学报,2019,34(1):19-24.SHEN Z,WU Y H,DANG P,et al.Comparison of the enzyme activities and soil nutrients in the rhizosphere of Pinus tabuliformis seedlings of two origins in hilly and gully regions of loess plateau[J].Journal of Northwest Forestry University,2019,34(1):19-24.(in Chinese)
[9] 王迪海,赵忠,张彦.黄土高原刺槐细根与土壤水分特征[J].西北林学院学报,2012,27(1):1-5.WANG D H,ZHAO Z,ZHANG Y.The fine root of robinia pseudoacacia and soil moisture in the Loess Plateau[J].Journal of Northwest Forestry University,2012,27(1):1-5.(in Chinese)
[10] 廖良宁,卢姿瑾,李远发,等.桂西北细叶云南松天然林凋落物及土壤养分特征[J].西北林学院学报,2019,34(1):31-38.LIAO L N,LU Z J,LI Y F,et al.Litter and soil nutrient characteristics of Pinus yunnanensis var.tenuifolia natural forest in northwest Guangxi[J].Journal of Northwest Forestry University,2019,34(1):31-38.(in Chinese)
[11] 郑淑霞,上官周平.不同功能型植物光合特性及其与叶氮含量、比叶重的关系[J].生态学报,2007,27(1):171-181.
[12] GAUDET F,HODGSON J Q,EDEN A,et al.Induction of tumors in mice by genomic hypomethylation[J].Science,2003,300(5618):489-492.
[13] 金樑,杜晓光,侯扶江,等.黄土高原山地坡度对退耕农田生态系统天然植被演替初期的影响[J].草业科学,2007,168(7):66-71.
[14] 丁曼,温仲明,郑颖.黄土丘陵区植物功能性状的尺度变化与依赖[J].生态学报,2014,34(9):2308-2315.DING M,WEN Z M,ZHENG Y.Scale change and dependence of plant functional traits in hilly areas of the Loess Region,Shaanxi Province,China[J].Acta Ecologica Sinica,2014,34(9):2308-2315.(in Chinese)
[15] 施宇,温仲明,龚时慧.黄土丘陵区植物叶片与细根功能性状关系及其变化[J].生态学报,2011,31(22):6805-6814.SHI Y,WEN Z M,GONG S H.Comparisons of relationships between leaf and fine root traits in hilly area of the Loess Plateau,Yanhe river basin,Shaanxi Province,China[J].Acta Ecologica Sinica,2011,31(22):6805-6814.(in Chinese)
[16] 戚德辉,温仲明,杨士梭,等.基于功能性状的铁杆蒿对环境变化的响应与适应[J].应用生态学报,2015,26(7):1921-1927.
[17] 王海珍,韩路,徐雅丽,等.土壤水分梯度对灰胡杨光合作用与抗逆性的影响[J].生态学报,2017,37(2):432-442.WANG H Z,HAN L,XU Y L,et al.Effects of soil water gradient on photosynthetic characteristics and stress resistanc of Populus pruinosa in the tarim basin,China[J].Acta Ecologica Sinica,2017,37(2):432-442.(in Chinese)
[18] 杨士梭,温仲明,苗连朋,等.黄土丘陵区植物功能性状对微地形变化的响应[J].应用生态学报,2014,25(12):3413-3419.
[19] 谢锦,常顺利,张毓涛,等.天山北坡植物土壤生态化学计量特征的垂直地带性[J].生态学报,2016,36(14):4363-4372.XIE J,CHANG S L,ZHANG Y T,et al.Plant and soil ecological stoichiometry with vertical zonality on the northern slope ofthe middle Tianshan Mountains[J].Acta Ecologica Sinica,2016,36(14):4363-4372.(in Chinese)
[20] 温培才,盛茂银,王霖娇,等.西南喀斯特高原盆地石漠化环境植物群落结构与物种多样性时空动态[J].广西植物,2018,38(1):11-23.WEN P C,SHENG M Y,WANG L J,et al.Temporal-spatial dynamics for plant community structure and species diversity of rocky desertification ecosystem in karst plateau basin,southwest China [J].Guihaia,2018,38(1):11-23.(in Chinese)
[21] 张巧明,王得祥,龚明贵,等.秦岭火地塘林区不同海拔森林土壤理化性质[J].水土保持学报,2011,25(5):69-73.
[22] 孙娇,赵发珠,韩新辉,等.不同林龄刺槐林土壤团聚体化学计量特征及其与土壤养分的关系[J].生态学报,2016,36(21):6879-6888.SUN J,ZHAO F Z,HAN X H,et al.Ecological stoichiometry of soil aggregates and relationship with soil nutrients of different-aged Robinia pseudoacacia forests[J].Acta Ecologica Sinica,2016,36(21):6879-6888.(in Chinese)
[23] 杨秀清,史婵,王旭刚,等.关帝山云杉次生林土壤的空间异质性及其与地形相关性[J].中国水土保持科学,2017,15(4):16-24.
[24] 韦兰英,上官周平.黄土高原白羊草、沙棘和辽东栎细根比根长特性[J].生态学报,2006,26(12):4164-4170.
[25] 袁帅.长白内山区不同森林群落中主要树种植物功能性状与固碳能力初探[D].长春:东北师范大学,2011.