亚高寒草甸阳坡—阴坡梯度上植物功能性状及群落构建机制研究
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摘要
在阳坡-阴坡梯度上,由于地形学的缘故百米的距离可能就会造成环境因素极大的变化,植物群落组成、物种多样性和生产力等随之也发生了很大的改变。目前,国内外关于阳坡-阴坡梯度植物生态学的研究主要是描述性的,很少有研究从功能性状的角度去揭示群落的构建以及植物的功能性状对于不同坡向的响应,从而探讨不同坡向物种的共存机制以及生存策略。功能性状是指间接的通过影响物种的“生长-生殖-生存”三方面的个体表现,而影响物种适合度的任何一种“形态-生理-物候”(morpho-physio-phenological)性状,以功能性状为基础的方法,目前正广泛的应用于生态学研究中。
基于功能性状的生态学研究中,一个重要的方面是植物对环境的适应。物种对环境的适应性是植物生存策略的一个重要方面。植物在漫长的进化和发展过程中,与环境相互作用,逐渐形成了许多内在生理和外在形态方面的适应对策,以最大程度地减小环境的不利影响,这些适应对策的表现即为植物性状(Plant traits)。叶性状,如比叶面积、叶片氮含量,已经被广泛认为与植物的资源利用策略相关。除了资源利用策略之外,叶片性状还反映了不同的“投资回报“策略。随着环境的改变,物种会采取不同的策略以使得自身在不同的环境下更具竞争力,而这就会表现在功能性状的改变上。
基于功能性状的生态学研究中,另一个重要方面是探讨群落的构建。性状的研究使得生态学家从以种群统计为基础的群落学研究中脱离出来,可以研究多物种的共存。以功能性状为基础的群落构建法则有两种,一种是环境筛选,一种是生物间的相互作用。环境筛选的作用使在同一地区生存的物种具有相似的性状,而生物间的相互作用使得共存物种的性状趋异。这两种构建法则在不同环境中可能具有不同的作用力,进而影响了功能多样性。例如在环境资源较差的条件下物种间竞争强烈,生态位的分化也更加强烈,以使物种能充分互补的利用有限的资源,提高了群落的功能多样性。随着以性状为基础的群落生态学研究方法的发展,一些学者认为除了物种多样性,功能多样性也对生态系统功能发挥着重要的作用。功能多样性就是在一个群落中功能性状的值,范围和分布。
综合以上的研究背景,在本研究中我们主要从以下三个方面研究了阳坡-阴坡梯度上植物群落的变化和群落构建的机理。首先,探索阳坡-阴坡梯度上植被差异的主要限制性环境因子是我们探索群落构建等生态学问题的基础。我们沿着阳坡-阴坡梯度在两个山地设置了连续的九个样地,在每个样地测量了六个主要的环境因子并调查了植被结构。其次,我们通过研究植物的叶片功能性状,探索了不同坡向的植物资源利用策略和投资回报策略。我们在甘南合作的亚高寒草甸选取山地的阴坡和阳坡,分别测量了四个叶片功能性状：比叶面积,叶片氮含量,叶片磷含量和叶片氮磷比。再次,我们从研究群落功能多样性入手探讨阳坡-阴坡梯度上植物群落的构建和物种的共存机制。本研究我们选用了两个物种性状,比叶面积和高度,来研究坡向梯度上功能性状多样性和物种多样性的格局。
通过以上的实验研究我们得出了以下结果：1)土壤含水量从阳坡向阴坡逐渐提高,是该梯度上主要的限制性因素,可以解释整个植被差异的28%；物种多样性沿着该梯度显著的提高,与土壤含水量呈现显著的正相关性；从阳坡到阴坡,生产力却显著地降低,与土壤含水量呈现显著的负相关关系；物种多样性与生产力呈现显著的负相关关系；2)每个功能群的平均比叶面积和叶片磷含量都呈现出阳坡显著小于阴坡,叶片氮磷比是阴坡显著的小于阳坡；在阳坡禾草(禾本科+莎草科)是优势功能群,而在阴坡杂草(非豆科杂草)则是优势功能群,而且禾草的比叶面积,叶片营养含量都显著的低于杂草；土壤含水量与群落组成和叶性状的变化有显著的相关性；3)两种类型的多样性在此梯度上的变化并不一致。物种多样性沿着阳坡-阴坡梯度逐渐增大,而功能多样性则沿着此梯度逐渐降低；而功能多样性则由生态位分化和物种丰富度共同决定,并受土壤含水量的影响；物种多样性和功能多样性之间呈现显著的二次曲线的关系。
由以上实验结果可得出本研究主要结论如下：
1)在高寒草甸阴阳坡梯度上决定植被结构的主要限制性资源为土壤含水量。
2)沿着阳坡-阴坡梯度,物种共存策略并不一致,阳坡的植物增大了生态位的分化以利用阳坡较为贫瘠的资源,而阴坡植物的生态位分化程度较小。土壤水分决定了物种多样性,而物种丰富度和生态位分化则共同决定了功能多样性,同时也受到土壤含水量的影响。
3)植被的资源利用策略在阴坡和阳坡发生了很大的变化：阴坡植物采取了快速获取资源的快回报策略,而阳坡植物则采取了有效贮存资源的慢回报策略。这种变化可以从不同坡向植物的叶性状的差异和优势功能群的改变反映出来,土壤含水量的改变是影响策略改变的重要因素。
Although located only a few hundred meters apart and sharing the same macroclimatic zone, the microclimatic conditions on different slopes vary dramatically affecting species composition, biodiversity and productivity. Recently most studies along the south- to north-facing slope are focused on descriptions of the different vegetation; however, few studies investigated the mechanisms of the community assembly and strategies of plant resource use based on functional traits. The use of methods based on functional traits is growing in the recent ecology studies. Functional traits are defined as morpho-physio-phenological traits that impact fitness indirectly via their effects on growth, reproduction and survival, three components of individual performance.
In ecology studies based on functional traits, adaptation of vegetation to environment is one of the most important aspects. Adaptation to environment matters the survival strategy of plants. In the process of evolution and development, plants formed many physiological and morphological attributes adopting environment conditions in order to diminish the adverse effect. Especially, leaf functional traits, such as specific leaf area and leaf nitrogen content, are widely considered to associate with resource use strategies. Besides, leaf functional traits have also been suggested to respond the plant investment-return strategy.
Another important aspect in ecological studies based on functional traits is exploring the community assembly mechanism. The approach based on traits enables ecologists to distract from the population demography studying pairwise species and to rebuild community ecology studying more co-existing species whose abundance will be decreased or increased by their different traits. The mechanism of species co-existing associates with the niche space partitioning which also associates functional diversity. Some researchers have suggested that functional diversity may be what is important for ecosystem function. Functional diversity is the range and distribution of functional trait values in a community.
On the basis of the above-mentioned background, in our study we mainly do the researches on the vegetation adaptations and community assembly and obtain important results from the three aspects as following:firstly, in this study we measured soil factors and vegetation composition along this gradient to explore the limiting resource factors affecting vegetation distribution. Secondly, we explored the resource use strategies on south- and north- facing slopes. In this study, leaf functional traits [specific leaf area (SLA), leaf nitrogen content (LNC), leaf phosphorus content (LPC) and leaf N:P ratio] and vegetation composition were investigated on south- and north- facing slopes in a sub-alpine meadow. Thirdly,beginning with studying functional diversity, we explored the niche space partitioning in plant communities to investigate the community assembly and species coexistence on different slope aspects; we chose two functional traits (specific leaf area and plant height) to study the patterns of species diversity and functional diversity along the slope gradient.
Our results showed:1) Soil water content was the primary limiting factors in this gradient; species diversity increased but primary productivity declined or comparable from south- to north-facing slope; species diversity and productivity positively and negatively relates to soil water content, respectively; species diversity and productivity were negatively linearly related.2) both SLA and LPC of each functional group were significantly higher and N:P ratio was lower on north- than on south- facing slopes; furthermore, grasses dominated south-facing slopes, while north-facing slopes were dominated by forbs. In addition, soil water content was significantly related to the shift in functional groups along the south- to north- facing slope gradient; soil water availability significantly related vegetation composition and leaf functional traits.3) the pattern of species diversity and functional diversity are not similar along this gradient. Both species richness and Shannon-wiener index were increased, but oppositely functional diversity was slightly decreased or comparable from south- to north- facing slope; soil water content (SWC) was the limiting resource in this region and determined species diversity. While functional diversity was determined by both species diversity and niche differentiation with the influence of soil water content; species diversity and functional diversity formed a significant quadratic regression relationship.
The main conclusions we got are as followings:
1) Soil water content was the primary limiting factors in this gradient determining vegetation distribution.
2) The pattern of species diversity and functional diversity are not similar along this gradient. Species diversity increased, but functional diversity decreased from south- to north- facing slope; Soil water content (SWC) was the limiting resource in this region and determined species diversity. While functional diversity was determined by both species diversity and niche differentiation with the influence of soil water content; Species diversity and functional diversity formed a significant quadratic regression relationship.
3) The change of functional traits and dominant functional group implied a difference in resource-use strategy between the two slope aspects, reflecting a transition in investment-return strategy from slow on south-facing slopes to fast on north-facing slopes. Moreover, we concluded that soil water availability affected vegetation composition and leaf functional traits, resulting in the shift in resource-use strategy.
基于功能性状的生态学研究中,一个重要的方面是植物对环境的适应。物种对环境的适应性是植物生存策略的一个重要方面。植物在漫长的进化和发展过程中,与环境相互作用,逐渐形成了许多内在生理和外在形态方面的适应对策,以最大程度地减小环境的不利影响,这些适应对策的表现即为植物性状(Plant traits)。叶性状,如比叶面积、叶片氮含量,已经被广泛认为与植物的资源利用策略相关。除了资源利用策略之外,叶片性状还反映了不同的“投资回报“策略。随着环境的改变,物种会采取不同的策略以使得自身在不同的环境下更具竞争力,而这就会表现在功能性状的改变上。
基于功能性状的生态学研究中,另一个重要方面是探讨群落的构建。性状的研究使得生态学家从以种群统计为基础的群落学研究中脱离出来,可以研究多物种的共存。以功能性状为基础的群落构建法则有两种,一种是环境筛选,一种是生物间的相互作用。环境筛选的作用使在同一地区生存的物种具有相似的性状,而生物间的相互作用使得共存物种的性状趋异。这两种构建法则在不同环境中可能具有不同的作用力,进而影响了功能多样性。例如在环境资源较差的条件下物种间竞争强烈,生态位的分化也更加强烈,以使物种能充分互补的利用有限的资源,提高了群落的功能多样性。随着以性状为基础的群落生态学研究方法的发展,一些学者认为除了物种多样性,功能多样性也对生态系统功能发挥着重要的作用。功能多样性就是在一个群落中功能性状的值,范围和分布。
综合以上的研究背景,在本研究中我们主要从以下三个方面研究了阳坡-阴坡梯度上植物群落的变化和群落构建的机理。首先,探索阳坡-阴坡梯度上植被差异的主要限制性环境因子是我们探索群落构建等生态学问题的基础。我们沿着阳坡-阴坡梯度在两个山地设置了连续的九个样地,在每个样地测量了六个主要的环境因子并调查了植被结构。其次,我们通过研究植物的叶片功能性状,探索了不同坡向的植物资源利用策略和投资回报策略。我们在甘南合作的亚高寒草甸选取山地的阴坡和阳坡,分别测量了四个叶片功能性状：比叶面积,叶片氮含量,叶片磷含量和叶片氮磷比。再次,我们从研究群落功能多样性入手探讨阳坡-阴坡梯度上植物群落的构建和物种的共存机制。本研究我们选用了两个物种性状,比叶面积和高度,来研究坡向梯度上功能性状多样性和物种多样性的格局。
通过以上的实验研究我们得出了以下结果：1)土壤含水量从阳坡向阴坡逐渐提高,是该梯度上主要的限制性因素,可以解释整个植被差异的28%；物种多样性沿着该梯度显著的提高,与土壤含水量呈现显著的正相关性；从阳坡到阴坡,生产力却显著地降低,与土壤含水量呈现显著的负相关关系；物种多样性与生产力呈现显著的负相关关系；2)每个功能群的平均比叶面积和叶片磷含量都呈现出阳坡显著小于阴坡,叶片氮磷比是阴坡显著的小于阳坡；在阳坡禾草(禾本科+莎草科)是优势功能群,而在阴坡杂草(非豆科杂草)则是优势功能群,而且禾草的比叶面积,叶片营养含量都显著的低于杂草；土壤含水量与群落组成和叶性状的变化有显著的相关性；3)两种类型的多样性在此梯度上的变化并不一致。物种多样性沿着阳坡-阴坡梯度逐渐增大,而功能多样性则沿着此梯度逐渐降低；而功能多样性则由生态位分化和物种丰富度共同决定,并受土壤含水量的影响；物种多样性和功能多样性之间呈现显著的二次曲线的关系。
由以上实验结果可得出本研究主要结论如下：
1)在高寒草甸阴阳坡梯度上决定植被结构的主要限制性资源为土壤含水量。
2)沿着阳坡-阴坡梯度,物种共存策略并不一致,阳坡的植物增大了生态位的分化以利用阳坡较为贫瘠的资源,而阴坡植物的生态位分化程度较小。土壤水分决定了物种多样性,而物种丰富度和生态位分化则共同决定了功能多样性,同时也受到土壤含水量的影响。
3)植被的资源利用策略在阴坡和阳坡发生了很大的变化：阴坡植物采取了快速获取资源的快回报策略,而阳坡植物则采取了有效贮存资源的慢回报策略。这种变化可以从不同坡向植物的叶性状的差异和优势功能群的改变反映出来,土壤含水量的改变是影响策略改变的重要因素。
Although located only a few hundred meters apart and sharing the same macroclimatic zone, the microclimatic conditions on different slopes vary dramatically affecting species composition, biodiversity and productivity. Recently most studies along the south- to north-facing slope are focused on descriptions of the different vegetation; however, few studies investigated the mechanisms of the community assembly and strategies of plant resource use based on functional traits. The use of methods based on functional traits is growing in the recent ecology studies. Functional traits are defined as morpho-physio-phenological traits that impact fitness indirectly via their effects on growth, reproduction and survival, three components of individual performance.
In ecology studies based on functional traits, adaptation of vegetation to environment is one of the most important aspects. Adaptation to environment matters the survival strategy of plants. In the process of evolution and development, plants formed many physiological and morphological attributes adopting environment conditions in order to diminish the adverse effect. Especially, leaf functional traits, such as specific leaf area and leaf nitrogen content, are widely considered to associate with resource use strategies. Besides, leaf functional traits have also been suggested to respond the plant investment-return strategy.
Another important aspect in ecological studies based on functional traits is exploring the community assembly mechanism. The approach based on traits enables ecologists to distract from the population demography studying pairwise species and to rebuild community ecology studying more co-existing species whose abundance will be decreased or increased by their different traits. The mechanism of species co-existing associates with the niche space partitioning which also associates functional diversity. Some researchers have suggested that functional diversity may be what is important for ecosystem function. Functional diversity is the range and distribution of functional trait values in a community.
On the basis of the above-mentioned background, in our study we mainly do the researches on the vegetation adaptations and community assembly and obtain important results from the three aspects as following:firstly, in this study we measured soil factors and vegetation composition along this gradient to explore the limiting resource factors affecting vegetation distribution. Secondly, we explored the resource use strategies on south- and north- facing slopes. In this study, leaf functional traits [specific leaf area (SLA), leaf nitrogen content (LNC), leaf phosphorus content (LPC) and leaf N:P ratio] and vegetation composition were investigated on south- and north- facing slopes in a sub-alpine meadow. Thirdly,beginning with studying functional diversity, we explored the niche space partitioning in plant communities to investigate the community assembly and species coexistence on different slope aspects; we chose two functional traits (specific leaf area and plant height) to study the patterns of species diversity and functional diversity along the slope gradient.
Our results showed:1) Soil water content was the primary limiting factors in this gradient; species diversity increased but primary productivity declined or comparable from south- to north-facing slope; species diversity and productivity positively and negatively relates to soil water content, respectively; species diversity and productivity were negatively linearly related.2) both SLA and LPC of each functional group were significantly higher and N:P ratio was lower on north- than on south- facing slopes; furthermore, grasses dominated south-facing slopes, while north-facing slopes were dominated by forbs. In addition, soil water content was significantly related to the shift in functional groups along the south- to north- facing slope gradient; soil water availability significantly related vegetation composition and leaf functional traits.3) the pattern of species diversity and functional diversity are not similar along this gradient. Both species richness and Shannon-wiener index were increased, but oppositely functional diversity was slightly decreased or comparable from south- to north- facing slope; soil water content (SWC) was the limiting resource in this region and determined species diversity. While functional diversity was determined by both species diversity and niche differentiation with the influence of soil water content; species diversity and functional diversity formed a significant quadratic regression relationship.
The main conclusions we got are as followings:
1) Soil water content was the primary limiting factors in this gradient determining vegetation distribution.
2) The pattern of species diversity and functional diversity are not similar along this gradient. Species diversity increased, but functional diversity decreased from south- to north- facing slope; Soil water content (SWC) was the limiting resource in this region and determined species diversity. While functional diversity was determined by both species diversity and niche differentiation with the influence of soil water content; Species diversity and functional diversity formed a significant quadratic regression relationship.
3) The change of functional traits and dominant functional group implied a difference in resource-use strategy between the two slope aspects, reflecting a transition in investment-return strategy from slow on south-facing slopes to fast on north-facing slopes. Moreover, we concluded that soil water availability affected vegetation composition and leaf functional traits, resulting in the shift in resource-use strategy.
引文
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