青藏高原东部高寒草甸群落常见草本植物种子幼苗更新与生长研究
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摘要
幼苗是植物生活史中的一个重要阶段，植物通过幼苗更新才能使种群得以扩大、扩散和延续。从种子萌发到幼苗建成的更新过程与植物群落的物种分布、丰富度以及群落的更新演替是密切相关的，特别是高寒草甸作为多年寒中生植物草本群落，其组分种的种子作为植被更新的潜在动力，对植被的演替和更新具有很大的贡献。而适宜的生境是植物幼苗更新的外在条件，由于过度放牧和人为施肥等不同的管理措施可以导致高寒草甸群落中的微生境差异。但微生境如何影响群落组分种的种子萌发和幼苗生长，至今尚未见系统的研究报道。因此，我们利用人工模拟的方法，研究了青藏高原东部高寒草甸群落中常见的杂类草组分种在不同微生境(光照、养分)下的幼苗更新(从种子萌发到幼苗出土、存活至越冬)过程和生长(生长率、生物量积累和分配)特征。以期揭示高寒草甸群落中常见杂类草组分种的幼苗更新机制和过程，为草地的退化和恢复过程提供理论基础。我们的研究结果主要有以下几个方面：
1)光照微生境和种子大小的变化显著影响了这些杂类草组分种的萌发特征。本研究的20个物种间，小种子物种的萌发具有较大的光需求，而较大种子物种萌发具有相对较小的光需求，大部分物种的萌发是具有较小的或中性的光依赖性，其最适萌发的光照生境是中等光照和较低光照生境。大部分物种属于在较短时间内、集中性萌发并具有较大萌发率的物种，而较少物种属于在较长时间内、分散随机萌发并具有较低萌发率的物种。
2)光照和种子大小差异显著影响了幼苗的出土率、存活率和越冬率。幼苗出土率、存活率、越冬率均与光照强度呈显著的正相关关系，而与种子大小的相互关系则随着生境差异发生了变化。本研究的大部分物种的幼苗最大出土率、存活率和越冬率均出现在中等光照和高光照条件下。高养分生境可以显著提高一些物种的幼苗存活率和越冬率，而对出土率没有显著影响。
3)中等光照下的高养分微生境可以显著提高这些物种的幼苗更新补充概率。研究物种幼苗的更新补充率与光照呈显著正相关关系，养分差异也显著影响了物种内的更新补充率。在本研究的几个微生境下，种子幼苗平均更新补充概率约为10％～30％。
4)微生境和种子大小显著影响了幼苗的生长率和生物量分配特征。幼苗的相对生长率、生物量积累、根分配、生物量根冠比和形态根冠比与光照强度呈显著正相关关系；幼苗的叶分配、茎分配、比叶面积和叶面积比与光照强度均呈显著的负相关关系。种子大小与初始幼苗大小、80天时的幼苗生物量和茎分配之间呈现显著的正相关关系；而与幼苗相对生长率、叶分配及遮荫生境下的幼苗长度(形态)根冠比均表现为显著的负相关关系。幼苗的最大相对生长率和生物量一般都出现在高养分生境下的高光照和中等光照条件下。
5)较小种子物种幼苗具有较大的幼苗生长和更新补充可塑性。生境变化对较小种子物种的幼苗生长和更新补充产生更大的影响，而对较大种子物种的幼苗生长和更新补充则产生相对较小的影响。具有较小种子的物种幼苗比较大种子物种幼苗具有更大的生长可塑性和更新补充可塑性。
6)幼苗通过其异质微生境下的形态生长特征的可塑性来改变其更新补充。在变化的微生境下，具有较小生长可塑性的物种具有相对较大的幼苗存活率；而生物量、株高可塑性和平均可塑性较高的物种具有相对较大的幼苗越冬存活率；具有较大的生物量分配和相对生长率可塑性的物种幼苗具有相对较大的存活率。
7)放牧干扰下的高光照微生境有利于这些草甸常见组分种实生幼苗的更新补充，而施肥后草甸群落冠层底部的低光生境不利于这些物种的更新补充。不同种子大小的物种在不同微生境下表现了不同的幼苗更新特征，幼苗通过其形态特征可塑性调节来适应生境变化，提高其生存适合度内容，进而使其达到较大的出土、存活、越冬和成功更新补充概率。
Seedling period is the crucial stage during the whole plant life-history. Plants mainly make population extended, distributed and continued by seedling recruitment. The recruitment process from seed germination to seedling establishment is nearly related to species distribution, abundance and community succession in plant communities. Especially for alpine meadows which belong to perennial plant community, seeds of component species as potential dynamical of vegetation recruitment have significant contribution to vegetation recruitment and succession, because offspring recruitment may influence general patterns of succession and maintenance of diversity within communities and plant species richness and density in an area depends on a combination of patterns of arrival and survival of seeds, seedling emergence, and seedling survival. Fitting habitats were the extrinsic conditions for successful seedling recruitment in plant community. There are many different microsites in alpine meadow communities which were caused by different grassland management measures, e.g. overgrazing and fertilization. But. up to now. there is little systemic research on seed germination, seedling growth and establishment of component species in alpine meadow communities under different microsites were reported. So. we conducted this study on seedling recruitment process and growth performance of some familiar forbs component species under different microsites (light and nutrient availability), which were made by a simulative artificially method, in alpine meadow communities of the eastern Qinghai-Tibetan Plateau. Our objective was to study the seedling recruitment mechanism and process of familiar forbs component species in alpine meadow communities, and offer a theoretical foundation for understanding on grassland degradation and restoration process. Main results were following:
1) Seed germination characteristics were significantly affected by light availability and seed mass for these forbs component species in alpine meadow communities. Among twenty species for this study, the smaller-seeded species had a bigger light requirement for germination than the larger-seeded species. Most of the studied species presented a lesser or neutral light dependence and the optimum light microsites were the middle and low light availability. For these species, they mostly adopt short-term and concentrative germination strategy and presented a higher germination percent, but several species adopt long time, scattered and stochastic germination strategy with a lower germination percent.
2) Light availability and seed mass also affected significantly seedling emergence rate, seedling survival rate and overwinter survival rate. They all presented a significant positive correlation with light availability. The relationships between seed mass and seedling emergence rate, seedling survival rate and overwinter survival rate were altered by the microsites change. For studied species, the maximum seedling emergence rate, survival rate and overwinter survival rate mostly appeared in middle and higher light conditions. Higher nutrient availability could significantly advanced seedling survival rate and overwinter survival rate for some species, but no significant effect on seedling emergence rate.
3) Middle light and higher nutrient microsite can significantly advanced seedling recruitment probability for these species. For studied species, seedling recruitment rate was significant positively related to light availability, and nutrient difference also significantly affected seedling recruitment rate within species. For studied twenty species in ten microsites, the mean seedling recruitment percent (from seed to seedling overwinter survival) were about 10% to 30%.
4) Microsite and seed mass both significantly affected seedling growth rate and biomass allocation characteristics. Seedling relative growth rate, biomass accumulation, root allocation, root to shoot ratio on biomass and on length all were significant positively related to light availability, but leaf allocation, stem allocation, specific leaf area and leaf area ratio were significant negatively related to light availability. Seed mass presented significant positive relationships with initial seedling size, seedling biomass at 80d and stem allocation, but negative relationships with relative growth rate, leaf allocation and root to shoot ratio on length under shade. The maximum seedling relative growth rate and biomass mostly appeared in microsites with higher nutrient and higher or middle light availability.
5) The smaller-seeded species presented larger recruitment and growth plasticity than the larger-seeded species in this study. Microsites differences brought the larger influences on seedling growth and recruitment for the smaller-seeded species, but the lesser influences for the larger-seeded species.
6) Seedling advanced their recruitment probability by their morphological performance plasticity in heterogeneous microsites. Species with lesser growth plasticity showed a higher seedling survival rate under different micorsites. Meanwhile, species with larger plasticity in biomass and height, and mean plasticity index presented higher seedling overwinter survival rate. Additionally, species with larger plasticity in biomass allocation and relative growth rate also showed higher seedling survival rate.
7) All these suggested that higher light availability in grazed disturbed meadow community favor seedling recruitment for these familiar forbs component species, but bottom shade in fertilized meadow community restrain seedling recruitment for these species. Species with different seed mass show different seedling recruitment characteristics under variational microsites. Seedlings advance their fitness in emergence, survival. overwinter survival and successful recruitment by their morphological performance plasticity under variational microsites.
1)光照微生境和种子大小的变化显著影响了这些杂类草组分种的萌发特征。本研究的20个物种间，小种子物种的萌发具有较大的光需求，而较大种子物种萌发具有相对较小的光需求，大部分物种的萌发是具有较小的或中性的光依赖性，其最适萌发的光照生境是中等光照和较低光照生境。大部分物种属于在较短时间内、集中性萌发并具有较大萌发率的物种，而较少物种属于在较长时间内、分散随机萌发并具有较低萌发率的物种。
2)光照和种子大小差异显著影响了幼苗的出土率、存活率和越冬率。幼苗出土率、存活率、越冬率均与光照强度呈显著的正相关关系，而与种子大小的相互关系则随着生境差异发生了变化。本研究的大部分物种的幼苗最大出土率、存活率和越冬率均出现在中等光照和高光照条件下。高养分生境可以显著提高一些物种的幼苗存活率和越冬率，而对出土率没有显著影响。
3)中等光照下的高养分微生境可以显著提高这些物种的幼苗更新补充概率。研究物种幼苗的更新补充率与光照呈显著正相关关系，养分差异也显著影响了物种内的更新补充率。在本研究的几个微生境下，种子幼苗平均更新补充概率约为10％～30％。
4)微生境和种子大小显著影响了幼苗的生长率和生物量分配特征。幼苗的相对生长率、生物量积累、根分配、生物量根冠比和形态根冠比与光照强度呈显著正相关关系；幼苗的叶分配、茎分配、比叶面积和叶面积比与光照强度均呈显著的负相关关系。种子大小与初始幼苗大小、80天时的幼苗生物量和茎分配之间呈现显著的正相关关系；而与幼苗相对生长率、叶分配及遮荫生境下的幼苗长度(形态)根冠比均表现为显著的负相关关系。幼苗的最大相对生长率和生物量一般都出现在高养分生境下的高光照和中等光照条件下。
5)较小种子物种幼苗具有较大的幼苗生长和更新补充可塑性。生境变化对较小种子物种的幼苗生长和更新补充产生更大的影响，而对较大种子物种的幼苗生长和更新补充则产生相对较小的影响。具有较小种子的物种幼苗比较大种子物种幼苗具有更大的生长可塑性和更新补充可塑性。
6)幼苗通过其异质微生境下的形态生长特征的可塑性来改变其更新补充。在变化的微生境下，具有较小生长可塑性的物种具有相对较大的幼苗存活率；而生物量、株高可塑性和平均可塑性较高的物种具有相对较大的幼苗越冬存活率；具有较大的生物量分配和相对生长率可塑性的物种幼苗具有相对较大的存活率。
7)放牧干扰下的高光照微生境有利于这些草甸常见组分种实生幼苗的更新补充，而施肥后草甸群落冠层底部的低光生境不利于这些物种的更新补充。不同种子大小的物种在不同微生境下表现了不同的幼苗更新特征，幼苗通过其形态特征可塑性调节来适应生境变化，提高其生存适合度内容，进而使其达到较大的出土、存活、越冬和成功更新补充概率。
Seedling period is the crucial stage during the whole plant life-history. Plants mainly make population extended, distributed and continued by seedling recruitment. The recruitment process from seed germination to seedling establishment is nearly related to species distribution, abundance and community succession in plant communities. Especially for alpine meadows which belong to perennial plant community, seeds of component species as potential dynamical of vegetation recruitment have significant contribution to vegetation recruitment and succession, because offspring recruitment may influence general patterns of succession and maintenance of diversity within communities and plant species richness and density in an area depends on a combination of patterns of arrival and survival of seeds, seedling emergence, and seedling survival. Fitting habitats were the extrinsic conditions for successful seedling recruitment in plant community. There are many different microsites in alpine meadow communities which were caused by different grassland management measures, e.g. overgrazing and fertilization. But. up to now. there is little systemic research on seed germination, seedling growth and establishment of component species in alpine meadow communities under different microsites were reported. So. we conducted this study on seedling recruitment process and growth performance of some familiar forbs component species under different microsites (light and nutrient availability), which were made by a simulative artificially method, in alpine meadow communities of the eastern Qinghai-Tibetan Plateau. Our objective was to study the seedling recruitment mechanism and process of familiar forbs component species in alpine meadow communities, and offer a theoretical foundation for understanding on grassland degradation and restoration process. Main results were following:
1) Seed germination characteristics were significantly affected by light availability and seed mass for these forbs component species in alpine meadow communities. Among twenty species for this study, the smaller-seeded species had a bigger light requirement for germination than the larger-seeded species. Most of the studied species presented a lesser or neutral light dependence and the optimum light microsites were the middle and low light availability. For these species, they mostly adopt short-term and concentrative germination strategy and presented a higher germination percent, but several species adopt long time, scattered and stochastic germination strategy with a lower germination percent.
2) Light availability and seed mass also affected significantly seedling emergence rate, seedling survival rate and overwinter survival rate. They all presented a significant positive correlation with light availability. The relationships between seed mass and seedling emergence rate, seedling survival rate and overwinter survival rate were altered by the microsites change. For studied species, the maximum seedling emergence rate, survival rate and overwinter survival rate mostly appeared in middle and higher light conditions. Higher nutrient availability could significantly advanced seedling survival rate and overwinter survival rate for some species, but no significant effect on seedling emergence rate.
3) Middle light and higher nutrient microsite can significantly advanced seedling recruitment probability for these species. For studied species, seedling recruitment rate was significant positively related to light availability, and nutrient difference also significantly affected seedling recruitment rate within species. For studied twenty species in ten microsites, the mean seedling recruitment percent (from seed to seedling overwinter survival) were about 10% to 30%.
4) Microsite and seed mass both significantly affected seedling growth rate and biomass allocation characteristics. Seedling relative growth rate, biomass accumulation, root allocation, root to shoot ratio on biomass and on length all were significant positively related to light availability, but leaf allocation, stem allocation, specific leaf area and leaf area ratio were significant negatively related to light availability. Seed mass presented significant positive relationships with initial seedling size, seedling biomass at 80d and stem allocation, but negative relationships with relative growth rate, leaf allocation and root to shoot ratio on length under shade. The maximum seedling relative growth rate and biomass mostly appeared in microsites with higher nutrient and higher or middle light availability.
5) The smaller-seeded species presented larger recruitment and growth plasticity than the larger-seeded species in this study. Microsites differences brought the larger influences on seedling growth and recruitment for the smaller-seeded species, but the lesser influences for the larger-seeded species.
6) Seedling advanced their recruitment probability by their morphological performance plasticity in heterogeneous microsites. Species with lesser growth plasticity showed a higher seedling survival rate under different micorsites. Meanwhile, species with larger plasticity in biomass and height, and mean plasticity index presented higher seedling overwinter survival rate. Additionally, species with larger plasticity in biomass allocation and relative growth rate also showed higher seedling survival rate.
7) All these suggested that higher light availability in grazed disturbed meadow community favor seedling recruitment for these familiar forbs component species, but bottom shade in fertilized meadow community restrain seedling recruitment for these species. Species with different seed mass show different seedling recruitment characteristics under variational microsites. Seedlings advance their fitness in emergence, survival. overwinter survival and successful recruitment by their morphological performance plasticity under variational microsites.
引文
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