南北样带温带区主要树种功能性状地理分异研究
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摘要
植物地理分布格局一直以来是生态学研究的重点内容。植物功能性状不但反映了植物对生境条件的响应和适应，而且能够将环境、植物和生态系统结构、过程与功能联系起来。在全球气候变化的背景下，通过分析植物功能性状与环境的关系，可以更好认识植物分布格局变化的环境驱动机制，为预测未来植被的发展趋势提供理论依据。中国东部南北样带作为全球15条陆地样带之一，环境梯度的变化造就了其特有的植被分布格局，而在南北样带温带区，栎属不同树种由北至南呈现出地理替代分布现象。本研究的目的就是在总体了解南北样带温带区主要树种功能性状地理分异特征的基础上，以南北样带温带区优势栎属树种为主要研究对象，探讨栎属树种生活史不同阶段的功能性状沿环境梯度的变化格局，揭示栎属树种地理替代分布的环境驱动机制。基于此，开展了4个方面的研究：(1)南北样带温带区主要树种功能性状与环境条件的关系；(2)南北样带温带区栎属树种功能性状沿环境梯度的变化格局；(3)南北样带温带区栎属树种种子性状对环境条件的响应和适应；(4)南北样带温带区栎属树种幼苗功能性状的变化差异。主要研究方法和结果如下：
(1)选取南北样带温带区11个地点的55种主要分布树种为研究对象，采用简单线性回归和多元线性逐步回归方法，分析主要树种功能性状与环境因子之间的关系，同时采用标准化主轴法分析功能性状间的关系，对南北样带温带区主要树种功能性状的地理分异特征进行探讨。结果表明：气候条件和土壤因子分别从不同方面驱动树种功能性状的变化分异，气候因子相对于土壤因子解释了更多功能性状的变化。随降水量的降低和土壤磷含量的增加，叶片干物质含量和比叶重均表现为增加的趋势；随日照时数的增加和土壤氮含量的降低，叶片单位重量氮含量增加；随土壤有机质含量和日照时数的增加，叶片单位重量磷含量增加；温度是制约树皮胸径比变化的唯一因素，随温度降低树皮厚度增加；而叶片钾含量在研究区域内没有显著变化。同时，功能性状间的关系更好的反映了主要树种的适应策略，叶片干物质含量和比叶重较高的树种能很好的生存在水分资源相对匮乏的生境里；叶片氮含量、磷含量较高的树种，用快速的养分循环来适应环境；而叶片钾含量也是南北样带温带区主要树种生长的关键因子。
(2)通过查阅栎属树种的标本，提取南北样带温带区23个地点的12种栎属树种的叶片形态信息，采用相关分析和标准化主轴分析方法，探讨了南北样带温带区栎属树种叶片形态性状沿气候梯度的变化，并结合已有对南北样带温带区栎属树种功能性状的研究，进一步讨论了成年林木阶段栎属树种地理分异的环境驱动机制。总体上，在南北样带温带区，栎属树种的叶片形态性状沿气候梯度发生显著的变化。叶片大小和叶片分裂程度随温度的降低和日照时数的增加而增加；随温度的增加和日照时数的减少，叶脉密度呈增加的趋势；随降水的增加，叶脉密度也逐渐增加。而且，栎属树种叶片形态性状相互关联，在有限资源下采用最优的叶片形态以最大化生长，叶片大小是关键的形态性状，与叶柄长度和叶片分裂程度显著正相关，与叶脉密度显著负相关；而随叶片倾向于长条状发展，叶柄长度和叶脉密度也随之增加。综合多个研究结果表明，气候条件的制约作用造成了南北样带温带区栎属树种的地理分布格局，栎属树种通过调节形态、结构和生理等性状的变化适应环境。
(3)选取南北样带温带区9个核心分布区的优势栎属树种作为研究对象，野外实地采集多种栎属树种的种子，通过测定种子的质量和化学组成，分析种子性状沿环境梯度的变化格局及其种子化学组成间的关系，结果发现：气候条件是驱动栎属树种种子性状变化的主要因素，这与栎属树种叶片功能性状与环境关系类似，而气候因子中温度的驱动作用较大，日照时数次之。种子质量较大的栎属树种更倾向分布于低温和长日照的样带北部区域；随温度降低，种子中淀粉含量积累减少的同时，多种抗寒物质如蛋白质、可溶性糖和脯氨酸等逐渐增加。而且，栎属树种为适应环境而调整种子的化学组成具有同步性，蛋白质、可溶性总糖和脯氨酸者相互正相关，而三者均与淀粉负相关。
(4)对采集于不同核心分布区的优势栎属树种的种子进行田间栽培实验，选择南北样带温带区的中部作为田间实验点，保证均一环境条件的同时，尽量减小环境胁迫对实验的影响。以均一环境下1年生幼苗为对象，研究比较种内及种间功能性状的变化差异，分析幼苗功能性状与成熟林木功能性状之间的关系，进一步讨论种子质量对幼苗生长性状的影响作用。结果发现：栎属树种功能性状的种内差异较小，种间差异性较大，种间变异系数高于种内，而且相同环境下幼苗的功能性状与原生地成年林木功能性状的相关性较低。种子质量对幼苗叶片功能性状的影响较小，而对生物量及其分配(根冠比)的影响较大。
本研究全面分析了南北样带温带区主要树种和栎属树种功能性状的地理分异特征，结果表明即使在相同尺度上，不同研究对象也存在不同的响应和适应机制，而对植物生活史的不同阶段，环境条件的影响作用也存在差异。这证实了植物物种分布格局的形成和变化并不是随机的和偶然的，环境条件在影响植物的同时，植物通过表观形态、结构组成和生理功能等一系列的权衡调节对环境进行响应，不同生活史阶段也形成不同的适应策略，从而在适宜区域进行定居、生长和繁衍。
The study of geographical distribution pattern has long been important in ecology. Plantfunctional traits not only reflect the response and adopt of plants to habitat conditions, but alsoconnect the environment to plants and the structure, process and function of ecological system.Under the background of global climate change, studies on the relationships between plantfunctional traits and environment can better understand the driving mechanism of theenvironment to the changes of plant distribution pattern, and provide the theoretical basis topredict the future trend of vegetation development. The North-South Transect of Eastern China(NSTEC) is one of15land belts in the world, with the unique vegetation distribution patternhere created by the environmental change. In temperate zone of NSTEC, the distribution ofQuercus trees presents a geographical vicarism pattern.
Hence, the purpose of this study was to analyze the pattern of functional traits change ofmain tree species along the environmental gradient, discuss the adaptive strategy of main treespecies, and then reveal the mechanism of the Quercus trees geographical vicarism at differentlife-history stages. This study included four aspects:(1) the relationships between theenvironmental conditions and functional traits of main tree species;(2) the change pattern offunctional traits of Quercus trees along the environmental gradient;(3) the response andadaptation of seeds traits of Quercus trees to the environmental conditions;(4) thedifferentiation of functional traits of Quercus trees seedling. The study was conducted intemperate zone of NSTEC, which includes two climatic zones, so the change of climate isdramatic.
The main research methods and results were as follows:
(1) To study the geographic differentiation of functional traits of main tree species intemperate zone of NSTEC,55kinds of main tree species at11sites were sampled. SimpleLinear Regression and Multiple Linear Stepwise Regression were used to analysis relationships between the functional traits and environmental factors, and Standardised Major Axis was usedto analysis relationships among the functional traits. The results showed that compared to thesoil factor, the climate factors play a greater role in driving the change of traits. With thedecrease of precipitation and the increase of soil phosphorus content, leaf mass per area (LMA)and leaf dry matter content (LDMC) tended to increase; leaf nitrogen content (Nmass) increasedwith the increase of sunlight time and the decrease of soil nitrogen content; leaf phosphoruscontent (Pmass) increased with the increase of soil organic matter content and sunlight time;temperature was the only factor to drive the change of ratio of bark to diameter at breast height,the thickness of bark increased with the decrease of temperature; leaf potassium content (Kmass)didn’t change significantly in the study area. In addition, the relationships among functionaltraits reflected the adaptative strategies of main tree species, such as the tree species of highLMA and LDMC could live better in the habitat of lack of water, as their ability to useenvironmental resources are stronger; tree species which are higher Nmassand Pmass, adapte tothe environment with slower speed of nutrient circulation; and Kmass was also the key factorwhich affect the growth of main tree species.
(2) Through refering to plant specimens, the leaf morphological information of12kindsof Quercus species at23sites was got. Pearson Correlation analysis was used to analysis thechange of leaf morphological traits of Quercus trees along the gradient of climate, andStandardised Major Axis method was used to discuss the relationships among leafmorphological traits. In addition, the other studies of functional traits of Quercus trees wereused to discuss the environment driving mechanism of Quercus trees geographic differentiation.On the whole, leaf morphological traits of Quercus trees changed significantly along thegradient of climate. Leaf areas and dissection of leaf margin increased with temperaturedropping and sunlight time increasing; leaf vein density increased with the increases oftemperature and precipitation，while decreased with the increase of sunlight time. Moreover,leaf morphological traits were relative to each other, and Quercus trees develop the optimalleaf morphology to grow maximumly with limited resources. The leaf size was the most important morphological traits: it correlated positively with petiole length and dissection ofleaf margin, but negatively with leaf vein density; when leaf tend to become the elongatedshape, the petiole length and leaf vein density also increased. The integration of several studiesindicated that the distribution pattern of Quercus trees is restricted primarily by climate,Quercus trees are adapted to the environment by adjusting the change of traits of morphology,structure and physiology.
(3) In order to reveal the change pattern of seed traits of Quercus trees along environmentgradient and the relationship among seed chemical compositions, Quercus species from9coredistribution areas were selected in temperate zone of NSTEC. Seeds were collected in the field,and then seed mass and chemical compositions were measured. The results showed that climatewas the main affective condition on the change of seed traits, while temperature was the mostimportant climate factor, and the sunlight time took second place. The kind of Quercus treeswhich seed mass is heavier tended to distribute in northern area of temperate zone of NSTECof low temperature and long sunlight time. With decreasing of temperature, the accumulationof starch content decreased, but some cold-tolerant substances such as protein, soluble sugarand proline increased gradually. Furthermore, the adaptive strategy of Quercus trees byadjusting changes of seed chemical compositions was also synchronized, as there were positiverelationships among the content of soluble sugar, protein and praline, and all of them werenegative relations with starch.
(4) The seeds which collected from different core distribution areas of Quercus trees werepretreated and sown in the same environment. In order to minimize effect of environmentalstress on the study, the central site in temperate zone of NSTEC was choose as the site ofcommon garden experiments.1year seedlings of Quercus trees are acted as research objects tocompare the difference and variation of different functional traits in intra-species andinter-species, and the relationships between functional traits of seedlings and mature trees areanalyzed, and the relationships between seed mass and seedlings traits are discussed. Theintraspecies differences of all functional traits were smaller than the interspecies difference, and so were the variation coefficients, and all functional traits of seedling were weaklycorrelation with mature trees. Seed mass was little effect on leaf functional traits of seedling,but important influence on the biomass and its allocation (RSR).
As all the results proved, the formation and change of plant species distribution pattern arenot random and accident, the environment can influence plants functional traits. Meanwhile,plants respond and adapt to environment through a series of modulation of morphology,structure and physiological function, and develop different adaptation strategies in differentgrowth phases, so plants are able to establish, grow and reproduce in the appropriate location.
(1)选取南北样带温带区11个地点的55种主要分布树种为研究对象，采用简单线性回归和多元线性逐步回归方法，分析主要树种功能性状与环境因子之间的关系，同时采用标准化主轴法分析功能性状间的关系，对南北样带温带区主要树种功能性状的地理分异特征进行探讨。结果表明：气候条件和土壤因子分别从不同方面驱动树种功能性状的变化分异，气候因子相对于土壤因子解释了更多功能性状的变化。随降水量的降低和土壤磷含量的增加，叶片干物质含量和比叶重均表现为增加的趋势；随日照时数的增加和土壤氮含量的降低，叶片单位重量氮含量增加；随土壤有机质含量和日照时数的增加，叶片单位重量磷含量增加；温度是制约树皮胸径比变化的唯一因素，随温度降低树皮厚度增加；而叶片钾含量在研究区域内没有显著变化。同时，功能性状间的关系更好的反映了主要树种的适应策略，叶片干物质含量和比叶重较高的树种能很好的生存在水分资源相对匮乏的生境里；叶片氮含量、磷含量较高的树种，用快速的养分循环来适应环境；而叶片钾含量也是南北样带温带区主要树种生长的关键因子。
(2)通过查阅栎属树种的标本，提取南北样带温带区23个地点的12种栎属树种的叶片形态信息，采用相关分析和标准化主轴分析方法，探讨了南北样带温带区栎属树种叶片形态性状沿气候梯度的变化，并结合已有对南北样带温带区栎属树种功能性状的研究，进一步讨论了成年林木阶段栎属树种地理分异的环境驱动机制。总体上，在南北样带温带区，栎属树种的叶片形态性状沿气候梯度发生显著的变化。叶片大小和叶片分裂程度随温度的降低和日照时数的增加而增加；随温度的增加和日照时数的减少，叶脉密度呈增加的趋势；随降水的增加，叶脉密度也逐渐增加。而且，栎属树种叶片形态性状相互关联，在有限资源下采用最优的叶片形态以最大化生长，叶片大小是关键的形态性状，与叶柄长度和叶片分裂程度显著正相关，与叶脉密度显著负相关；而随叶片倾向于长条状发展，叶柄长度和叶脉密度也随之增加。综合多个研究结果表明，气候条件的制约作用造成了南北样带温带区栎属树种的地理分布格局，栎属树种通过调节形态、结构和生理等性状的变化适应环境。
(3)选取南北样带温带区9个核心分布区的优势栎属树种作为研究对象，野外实地采集多种栎属树种的种子，通过测定种子的质量和化学组成，分析种子性状沿环境梯度的变化格局及其种子化学组成间的关系，结果发现：气候条件是驱动栎属树种种子性状变化的主要因素，这与栎属树种叶片功能性状与环境关系类似，而气候因子中温度的驱动作用较大，日照时数次之。种子质量较大的栎属树种更倾向分布于低温和长日照的样带北部区域；随温度降低，种子中淀粉含量积累减少的同时，多种抗寒物质如蛋白质、可溶性糖和脯氨酸等逐渐增加。而且，栎属树种为适应环境而调整种子的化学组成具有同步性，蛋白质、可溶性总糖和脯氨酸者相互正相关，而三者均与淀粉负相关。
(4)对采集于不同核心分布区的优势栎属树种的种子进行田间栽培实验，选择南北样带温带区的中部作为田间实验点，保证均一环境条件的同时，尽量减小环境胁迫对实验的影响。以均一环境下1年生幼苗为对象，研究比较种内及种间功能性状的变化差异，分析幼苗功能性状与成熟林木功能性状之间的关系，进一步讨论种子质量对幼苗生长性状的影响作用。结果发现：栎属树种功能性状的种内差异较小，种间差异性较大，种间变异系数高于种内，而且相同环境下幼苗的功能性状与原生地成年林木功能性状的相关性较低。种子质量对幼苗叶片功能性状的影响较小，而对生物量及其分配(根冠比)的影响较大。
本研究全面分析了南北样带温带区主要树种和栎属树种功能性状的地理分异特征，结果表明即使在相同尺度上，不同研究对象也存在不同的响应和适应机制，而对植物生活史的不同阶段，环境条件的影响作用也存在差异。这证实了植物物种分布格局的形成和变化并不是随机的和偶然的，环境条件在影响植物的同时，植物通过表观形态、结构组成和生理功能等一系列的权衡调节对环境进行响应，不同生活史阶段也形成不同的适应策略，从而在适宜区域进行定居、生长和繁衍。
The study of geographical distribution pattern has long been important in ecology. Plantfunctional traits not only reflect the response and adopt of plants to habitat conditions, but alsoconnect the environment to plants and the structure, process and function of ecological system.Under the background of global climate change, studies on the relationships between plantfunctional traits and environment can better understand the driving mechanism of theenvironment to the changes of plant distribution pattern, and provide the theoretical basis topredict the future trend of vegetation development. The North-South Transect of Eastern China(NSTEC) is one of15land belts in the world, with the unique vegetation distribution patternhere created by the environmental change. In temperate zone of NSTEC, the distribution ofQuercus trees presents a geographical vicarism pattern.
Hence, the purpose of this study was to analyze the pattern of functional traits change ofmain tree species along the environmental gradient, discuss the adaptive strategy of main treespecies, and then reveal the mechanism of the Quercus trees geographical vicarism at differentlife-history stages. This study included four aspects:(1) the relationships between theenvironmental conditions and functional traits of main tree species;(2) the change pattern offunctional traits of Quercus trees along the environmental gradient;(3) the response andadaptation of seeds traits of Quercus trees to the environmental conditions;(4) thedifferentiation of functional traits of Quercus trees seedling. The study was conducted intemperate zone of NSTEC, which includes two climatic zones, so the change of climate isdramatic.
The main research methods and results were as follows:
(1) To study the geographic differentiation of functional traits of main tree species intemperate zone of NSTEC,55kinds of main tree species at11sites were sampled. SimpleLinear Regression and Multiple Linear Stepwise Regression were used to analysis relationships between the functional traits and environmental factors, and Standardised Major Axis was usedto analysis relationships among the functional traits. The results showed that compared to thesoil factor, the climate factors play a greater role in driving the change of traits. With thedecrease of precipitation and the increase of soil phosphorus content, leaf mass per area (LMA)and leaf dry matter content (LDMC) tended to increase; leaf nitrogen content (Nmass) increasedwith the increase of sunlight time and the decrease of soil nitrogen content; leaf phosphoruscontent (Pmass) increased with the increase of soil organic matter content and sunlight time;temperature was the only factor to drive the change of ratio of bark to diameter at breast height,the thickness of bark increased with the decrease of temperature; leaf potassium content (Kmass)didn’t change significantly in the study area. In addition, the relationships among functionaltraits reflected the adaptative strategies of main tree species, such as the tree species of highLMA and LDMC could live better in the habitat of lack of water, as their ability to useenvironmental resources are stronger; tree species which are higher Nmassand Pmass, adapte tothe environment with slower speed of nutrient circulation; and Kmass was also the key factorwhich affect the growth of main tree species.
(2) Through refering to plant specimens, the leaf morphological information of12kindsof Quercus species at23sites was got. Pearson Correlation analysis was used to analysis thechange of leaf morphological traits of Quercus trees along the gradient of climate, andStandardised Major Axis method was used to discuss the relationships among leafmorphological traits. In addition, the other studies of functional traits of Quercus trees wereused to discuss the environment driving mechanism of Quercus trees geographic differentiation.On the whole, leaf morphological traits of Quercus trees changed significantly along thegradient of climate. Leaf areas and dissection of leaf margin increased with temperaturedropping and sunlight time increasing; leaf vein density increased with the increases oftemperature and precipitation，while decreased with the increase of sunlight time. Moreover,leaf morphological traits were relative to each other, and Quercus trees develop the optimalleaf morphology to grow maximumly with limited resources. The leaf size was the most important morphological traits: it correlated positively with petiole length and dissection ofleaf margin, but negatively with leaf vein density; when leaf tend to become the elongatedshape, the petiole length and leaf vein density also increased. The integration of several studiesindicated that the distribution pattern of Quercus trees is restricted primarily by climate,Quercus trees are adapted to the environment by adjusting the change of traits of morphology,structure and physiology.
(3) In order to reveal the change pattern of seed traits of Quercus trees along environmentgradient and the relationship among seed chemical compositions, Quercus species from9coredistribution areas were selected in temperate zone of NSTEC. Seeds were collected in the field,and then seed mass and chemical compositions were measured. The results showed that climatewas the main affective condition on the change of seed traits, while temperature was the mostimportant climate factor, and the sunlight time took second place. The kind of Quercus treeswhich seed mass is heavier tended to distribute in northern area of temperate zone of NSTECof low temperature and long sunlight time. With decreasing of temperature, the accumulationof starch content decreased, but some cold-tolerant substances such as protein, soluble sugarand proline increased gradually. Furthermore, the adaptive strategy of Quercus trees byadjusting changes of seed chemical compositions was also synchronized, as there were positiverelationships among the content of soluble sugar, protein and praline, and all of them werenegative relations with starch.
(4) The seeds which collected from different core distribution areas of Quercus trees werepretreated and sown in the same environment. In order to minimize effect of environmentalstress on the study, the central site in temperate zone of NSTEC was choose as the site ofcommon garden experiments.1year seedlings of Quercus trees are acted as research objects tocompare the difference and variation of different functional traits in intra-species andinter-species, and the relationships between functional traits of seedlings and mature trees areanalyzed, and the relationships between seed mass and seedlings traits are discussed. Theintraspecies differences of all functional traits were smaller than the interspecies difference, and so were the variation coefficients, and all functional traits of seedling were weaklycorrelation with mature trees. Seed mass was little effect on leaf functional traits of seedling,but important influence on the biomass and its allocation (RSR).
As all the results proved, the formation and change of plant species distribution pattern arenot random and accident, the environment can influence plants functional traits. Meanwhile,plants respond and adapt to environment through a series of modulation of morphology,structure and physiological function, and develop different adaptation strategies in differentgrowth phases, so plants are able to establish, grow and reproduce in the appropriate location.
引文
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