高寒草甸坡向梯度上植物群落组成及其氮磷化学计量学特征的研究
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摘要
坡向是青藏高原山地上决定植物群落组成变化的主导因子之一。坡向影响着地表接受的太阳辐射量,而太阳辐射在不同坡向上的再分配决定了土壤温度、土壤蒸发量、土壤含水量等生态因子的分配。南坡(阳坡)接受的太阳辐射较多,较热、干旱,而北坡(阴坡)接受的太阳辐射较少,故而较湿、阴冷。由于不同坡向光、热、水、土等自然因素的不同,植被的组成会产生相应的变化。这种在坡向的生境梯度上,植物群落组分结构上的变化如何影响土壤理化特征、植物的化学计量特征、抗逆生理特征等,而这些又如何响应于群落组成的变化,这是高寒草甸地区群落学研究中少有涉及的论题。
本文以青藏高原东部的高寒草甸为例,分析了阴阳坡梯度上土壤因子和植物群落组成的变化特征,研究了植物叶片氮磷化学计量特征对坡向的响应以及植物物种生理因子的坡向差异。
本文的主要结果为：
(1)从阳坡到阴坡,环境因子的变化较大,日平均土壤温度和光照度的变化趋势为阳坡>半阴半阳坡>阴坡,土壤含水量的变化趋势为阴坡>半阴半阳坡>阴坡。土壤养分的变化趋势为：土壤全磷和有机质总体表现为阴坡大于阳坡,而土壤全氮和速效氮、速效磷在坡向梯度上变化不显著。
(2)坡向梯度上植物组成结构发生了明显变化,阳坡主要由禾草(禾本科和莎草科)组成,阴坡则主要由杂草(非豆科杂草)组成。阴坡-阳坡梯度上群落物种多样性指数和丰富度指数的变化显著,其均表现为阴坡>半阴半阳坡>阳坡。
(3)在阳坡-阴坡梯度上,植物叶片的氮磷化学计量特征也发生了变化。在阳坡,植物叶片氮磷含量都小于阴坡,而磷含量的变化尤为显著；阳坡的叶片氮磷比则显著的高于阴坡。另外,植物叶片功能群的氮磷计量特征同样发生了变化,禾草类的平均叶片氮磷含量显著低于杂草类,而豆科的叶片氮含量和氮磷比在不同坡向均高于禾草和杂草类。
(4)在阳坡-阴坡梯度上,阳坡植物的氮磷比明显大于16,而阴坡则小于14,根据限制性元素计量比值判断,不同坡向上限制性元素发生了变化,即阳坡为磷限制,而阴坡则为氮限制。
(5)在阳坡-阴坡梯度上,植物的生长率与N：P呈显著负相关,与植物叶片磷含量呈显著正相关,这支持生长速率假说。另外,随着海拔高度的上升,植物叶片氮磷含量和氮磷比都呈现明显的下降趋势,这一现象支持生物-地球化学假说。
(6)在阴坡-阳坡梯度上,随着土壤含水量的下降和土壤温度、光照度的增加,植物物种脯氨酸与可溶性糖含量呈增加趋势,其大小顺序为：棘豆(Oxytropis kansuensis)>狼毒(Stellera chamaejweisme)>火绒草(Leontopodium leontopodioides)>莓叶委陵菜(Otentilla fragarioides)>矮嵩草(Kobresia humilis)>金露梅(Potentilla fruticosa),而植物叶绿素呈减少趋势,其大小顺序为：棘豆(Oxytropis kansuensis)<矮嵩草(Kobresia humilis)<莓叶委陵菜(Otentilla fragarioides)<火绒草(Leontopodium leontopodioides)<金露梅(Potentilla fruticosa)<狼毒(Stellera chamaejweisme),这些表明随着逆境胁迫的加剧,植物通过增加自身细胞的渗透调节物质来应对外界胁迫的能力。
本研究主要结论如下：
(1)在高寒草甸阴坡-阳坡梯度上,植物群落组成和物种多样性均发生了变化,引起这种变化的主导因素为土壤含水量。
(2)在阴坡-阳坡梯度上,植物叶片的氮磷化学计量特征并不一致,阴坡的植物叶片氮磷含量都显著高于阳坡,而其氮磷比则显著小于阳坡。限制性元素发生了变化,阴坡为氮限制,而阳坡则为磷限制,土壤含水量和土壤温度共同决定了这一变化。
(3)由阴坡至阳坡的生境梯度上,随着土壤含水量的下降和土壤温度、光照度的增加,逆境胁迫的增强,植物物种的脯氨酸和可溶性糖含量总体呈增加趋势,而叶绿素含量总体呈下降趋势,这是植物对逆境胁迫的一种生理适应,表明在阳坡植物具有较强的抗逆性。
Slope aspect is one of the key factors in determining plant community composition on terrestrial ecosystem of the Qinghai-Tibet Plateau Mountain. Slope aspect greatly influence the coming solar radiation, which determine soil temperature, soil evaporation, soil water content and other ecological factors. South-facing slope (sunny slope) absorbs more solar radiation, hence it is hotter and drier, while north-facing slope (shady slope) absorbs less solar radiation, so it is wetter and colder. Thus, vegetation responded with different structure and compositon in different slope aspects. Most studies on the large scale have shown that the distributions of light, heat, water affect vegetation structure and community composition on terrestrial ecosystem, many studies did on aspect impects-less on high altitude. Here we analyzed the variation characteristic of soil factors and plant community composition along north-south-facing slope gradient, studied the response of plant leaf N and P stoichiometry characteristic to slope aspect, and discussed the nitrogen, phosphorus stoichiometry differences in different functional groups and the physiological changes of species at north-and south-facing slope on a alpine meadow of the eastern Tibetan Plateau.
The main results of the study showed:
1. From the south-facing slopes to north-facing slope, the change of environmental factors was larger, daily average soil temperature and light intensity were south-facing slope> west-facing slope> north-facing slope, soil water content was north-facing slope> west-facing slope> south-facing slope. Soil total phosphorus and organic matter were greater in north-facing slope than south-facing slope. Soil total nitrogen, available nitrogen and available phosphorus didn't show a significant change in the aspect gradients.
2. Significant changes in plant composition occurred in the different slope aspects where the south-facing slope was dominated mainly by grasses (Poaceae and Cyperaceae) and the north-facing slope was dominated by mainly forbs (non-legumes forbs). Species diversity and richness changes significantly in the different slope aspects, it showed north-facing slope> west-facing slope> south-facing slope.
3. Nitrogen and phosphorus stoichiometry in plant leaves also changed dramatically from the south-facing slopes to north-facing slope, in the south-facing slope species average leaf nitrogen and phosphorus content were significantly less than the north-facing slope, but Leaf N:P ratio was significantly higher in south-facing slope than the north-facing slope. In addition, we also found that the nitrogen and phosphorus stoichiometry in the plant functional groups also changed, average plant leaf nitrogen and phosphorus content of grasses was significantly lower than the forbs, and the legume leaf nitrogen content and N:P ratio in different aspects higher than that of grasses and forbs.
4. The limiting elements changed from the south-facing slope to north-facing slope. In the south-facing slope the N:P ratio of plant leaf was greater dramaticlly than16, while the north-facing slope was less than14. Therefore, according to the limit elements stoichiometry ratio theory, south-facing slope was phosphorus limitation, while north-facing slope was nitrogen limitation.
5. From the south-facing slopes to north-facing slope, the plant growth rate was significantly negatively correlated with N:P, while it was significantly positively correlated with phosphorus content of plant leaves, which supported the growth rate hypothesis. Nitrogen and phosphorus content in plant leaves and N:P ratio showed a significantly decline trend with increasing of altitude, which supported the biogeochemical hypothesis.
6. Along the north-facing slope to south-facing slopes, soil water content declined and soil temperature, light intensity increased, the trend of proline and soluble sugar content of plant species were increased. The order of proline and soluble sugar content:Oxytropis kansuensis> Stellera chamaejweisme> Otentilla fragarioid.es> Kobresia humilis> Potentilla fruticosa, while the trend of plant chlorophyll content were declined, and the order plant chlorophyll content:Oxytropis kansuensis According to these results, we draw the following conclusions:
1. Soil water content was the key factor determining vegetation composition and species diversity on the north-and south-facing slope gradients in alpine meadow.
2. Plant leaf nitrogen and phosphorus stoichiometry was not consistent along the south-facing slope to north-facing slope gradients, the nitrogen and phosphorus content of the plant leaves on north-facing slope were significantly higher than south-facing, N:P ratio was significantly less than the south-facing slope. These suggested that these stoichiometry changes were controlled by soil water content and soil temperature.
3. Along the north-facing slope to south-facing slope gradients, soil water content declined and soil temperature, light intensity increased. With the increasing of stress, plant species of proline and soluble sugar content was significantly increased, while chlorophyll content decreased, and this was physiological adaptation of plant to stress, it shown that the stress resistance of plants was stronger in south-facing slope.
本文以青藏高原东部的高寒草甸为例,分析了阴阳坡梯度上土壤因子和植物群落组成的变化特征,研究了植物叶片氮磷化学计量特征对坡向的响应以及植物物种生理因子的坡向差异。
本文的主要结果为：
(1)从阳坡到阴坡,环境因子的变化较大,日平均土壤温度和光照度的变化趋势为阳坡>半阴半阳坡>阴坡,土壤含水量的变化趋势为阴坡>半阴半阳坡>阴坡。土壤养分的变化趋势为：土壤全磷和有机质总体表现为阴坡大于阳坡,而土壤全氮和速效氮、速效磷在坡向梯度上变化不显著。
(2)坡向梯度上植物组成结构发生了明显变化,阳坡主要由禾草(禾本科和莎草科)组成,阴坡则主要由杂草(非豆科杂草)组成。阴坡-阳坡梯度上群落物种多样性指数和丰富度指数的变化显著,其均表现为阴坡>半阴半阳坡>阳坡。
(3)在阳坡-阴坡梯度上,植物叶片的氮磷化学计量特征也发生了变化。在阳坡,植物叶片氮磷含量都小于阴坡,而磷含量的变化尤为显著；阳坡的叶片氮磷比则显著的高于阴坡。另外,植物叶片功能群的氮磷计量特征同样发生了变化,禾草类的平均叶片氮磷含量显著低于杂草类,而豆科的叶片氮含量和氮磷比在不同坡向均高于禾草和杂草类。
(4)在阳坡-阴坡梯度上,阳坡植物的氮磷比明显大于16,而阴坡则小于14,根据限制性元素计量比值判断,不同坡向上限制性元素发生了变化,即阳坡为磷限制,而阴坡则为氮限制。
(5)在阳坡-阴坡梯度上,植物的生长率与N：P呈显著负相关,与植物叶片磷含量呈显著正相关,这支持生长速率假说。另外,随着海拔高度的上升,植物叶片氮磷含量和氮磷比都呈现明显的下降趋势,这一现象支持生物-地球化学假说。
(6)在阴坡-阳坡梯度上,随着土壤含水量的下降和土壤温度、光照度的增加,植物物种脯氨酸与可溶性糖含量呈增加趋势,其大小顺序为：棘豆(Oxytropis kansuensis)>狼毒(Stellera chamaejweisme)>火绒草(Leontopodium leontopodioides)>莓叶委陵菜(Otentilla fragarioides)>矮嵩草(Kobresia humilis)>金露梅(Potentilla fruticosa),而植物叶绿素呈减少趋势,其大小顺序为：棘豆(Oxytropis kansuensis)<矮嵩草(Kobresia humilis)<莓叶委陵菜(Otentilla fragarioides)<火绒草(Leontopodium leontopodioides)<金露梅(Potentilla fruticosa)<狼毒(Stellera chamaejweisme),这些表明随着逆境胁迫的加剧,植物通过增加自身细胞的渗透调节物质来应对外界胁迫的能力。
本研究主要结论如下：
(1)在高寒草甸阴坡-阳坡梯度上,植物群落组成和物种多样性均发生了变化,引起这种变化的主导因素为土壤含水量。
(2)在阴坡-阳坡梯度上,植物叶片的氮磷化学计量特征并不一致,阴坡的植物叶片氮磷含量都显著高于阳坡,而其氮磷比则显著小于阳坡。限制性元素发生了变化,阴坡为氮限制,而阳坡则为磷限制,土壤含水量和土壤温度共同决定了这一变化。
(3)由阴坡至阳坡的生境梯度上,随着土壤含水量的下降和土壤温度、光照度的增加,逆境胁迫的增强,植物物种的脯氨酸和可溶性糖含量总体呈增加趋势,而叶绿素含量总体呈下降趋势,这是植物对逆境胁迫的一种生理适应,表明在阳坡植物具有较强的抗逆性。
Slope aspect is one of the key factors in determining plant community composition on terrestrial ecosystem of the Qinghai-Tibet Plateau Mountain. Slope aspect greatly influence the coming solar radiation, which determine soil temperature, soil evaporation, soil water content and other ecological factors. South-facing slope (sunny slope) absorbs more solar radiation, hence it is hotter and drier, while north-facing slope (shady slope) absorbs less solar radiation, so it is wetter and colder. Thus, vegetation responded with different structure and compositon in different slope aspects. Most studies on the large scale have shown that the distributions of light, heat, water affect vegetation structure and community composition on terrestrial ecosystem, many studies did on aspect impects-less on high altitude. Here we analyzed the variation characteristic of soil factors and plant community composition along north-south-facing slope gradient, studied the response of plant leaf N and P stoichiometry characteristic to slope aspect, and discussed the nitrogen, phosphorus stoichiometry differences in different functional groups and the physiological changes of species at north-and south-facing slope on a alpine meadow of the eastern Tibetan Plateau.
The main results of the study showed:
1. From the south-facing slopes to north-facing slope, the change of environmental factors was larger, daily average soil temperature and light intensity were south-facing slope> west-facing slope> north-facing slope, soil water content was north-facing slope> west-facing slope> south-facing slope. Soil total phosphorus and organic matter were greater in north-facing slope than south-facing slope. Soil total nitrogen, available nitrogen and available phosphorus didn't show a significant change in the aspect gradients.
2. Significant changes in plant composition occurred in the different slope aspects where the south-facing slope was dominated mainly by grasses (Poaceae and Cyperaceae) and the north-facing slope was dominated by mainly forbs (non-legumes forbs). Species diversity and richness changes significantly in the different slope aspects, it showed north-facing slope> west-facing slope> south-facing slope.
3. Nitrogen and phosphorus stoichiometry in plant leaves also changed dramatically from the south-facing slopes to north-facing slope, in the south-facing slope species average leaf nitrogen and phosphorus content were significantly less than the north-facing slope, but Leaf N:P ratio was significantly higher in south-facing slope than the north-facing slope. In addition, we also found that the nitrogen and phosphorus stoichiometry in the plant functional groups also changed, average plant leaf nitrogen and phosphorus content of grasses was significantly lower than the forbs, and the legume leaf nitrogen content and N:P ratio in different aspects higher than that of grasses and forbs.
4. The limiting elements changed from the south-facing slope to north-facing slope. In the south-facing slope the N:P ratio of plant leaf was greater dramaticlly than16, while the north-facing slope was less than14. Therefore, according to the limit elements stoichiometry ratio theory, south-facing slope was phosphorus limitation, while north-facing slope was nitrogen limitation.
5. From the south-facing slopes to north-facing slope, the plant growth rate was significantly negatively correlated with N:P, while it was significantly positively correlated with phosphorus content of plant leaves, which supported the growth rate hypothesis. Nitrogen and phosphorus content in plant leaves and N:P ratio showed a significantly decline trend with increasing of altitude, which supported the biogeochemical hypothesis.
6. Along the north-facing slope to south-facing slopes, soil water content declined and soil temperature, light intensity increased, the trend of proline and soluble sugar content of plant species were increased. The order of proline and soluble sugar content:Oxytropis kansuensis> Stellera chamaejweisme> Otentilla fragarioid.es> Kobresia humilis> Potentilla fruticosa, while the trend of plant chlorophyll content were declined, and the order plant chlorophyll content:Oxytropis kansuensis
1. Soil water content was the key factor determining vegetation composition and species diversity on the north-and south-facing slope gradients in alpine meadow.
2. Plant leaf nitrogen and phosphorus stoichiometry was not consistent along the south-facing slope to north-facing slope gradients, the nitrogen and phosphorus content of the plant leaves on north-facing slope were significantly higher than south-facing, N:P ratio was significantly less than the south-facing slope. These suggested that these stoichiometry changes were controlled by soil water content and soil temperature.
3. Along the north-facing slope to south-facing slope gradients, soil water content declined and soil temperature, light intensity increased. With the increasing of stress, plant species of proline and soluble sugar content was significantly increased, while chlorophyll content decreased, and this was physiological adaptation of plant to stress, it shown that the stress resistance of plants was stronger in south-facing slope.
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