摘要
全球气候环境变化对陆地植物生态系统的影响愈来愈多的受到人们的关注。其中植物物候对气候环境变化的响应是当今生态学研究的热点问题之一。植物物候及其发生的变化是多个环境因子综合影响的结果。群落中植物之间物候上的差异对群落的构建有重要的作用。氮素是陆地生态系统植物生长的主要限制性因子,但是在全球变化背景下氮素输入陆地生态系统的方式和数量都发生了变化,这引起了氮素在陆地生态系统中的富集。草地生态系统是陆地生态系统重要的组成部分,长期以来关于氮素营养对草地植物群落影响的研究大多集中在群落生产力和多样性以及群落结构的变化等方面。近年来的研究表明,草地群落中氮素的增加会引起植物物候的变化,尤其是植物花期物候的变化。由于全球环境变化对草地植物群落的影响日益明显,有关草地植物物候的研究受到了广泛的关注。但大多数草地植物物候的研究集中在低海拔地区进行,而在海拔高、气候寒冷、生态系统敏感和脆弱的青藏高原高寒草甸开展的研究较少。高寒草甸植物群落通常被短暂的生长季节和严酷的自然环境所限制,这使得高寒草甸植物更容易受到环境变化的影响。本研究以青藏高原东缘兰州大学高寒草甸与湿地生态系统定位研究站—玛曲县(阿孜)实验点的天然草甸为研究对象。我们设置了三个氮素添加梯度(对照处理CK、低氮添加处理LN,高氮添加处理HN),从2009年到2011年,进行了为期三年的氮素添加实验。通过对不同氮素添加处理下的植物群落物候的定点定时观测以及对群落中常见组分种功能性状的测定,研究了氮素添加对青藏高原东缘高寒草甸植物群落花期物候、植物性状和植物群落结构的影响。我们得出了如下的结果：
1.我们在研究区的自然对照群落中,2009年观测到58个物种开花,2010年观测到51个物种开花,2011年观测到55个物种开花。2009,2010,2011年群落中物种的开花序列分布表明,高寒草甸植物群落中物种的花期物候从每年的4月下旬开始至9月下旬结束,群落中各物种的始花时间、开花峰值期及花期持续期不同；
2.高寒草甸植物群落中不同氮素添加处理下植物的始花时间(first flowering date FFD)和开花持续时间(flowering duration date, FDD)成负相关。不同氮素添加处理之间群落的始花期和开花持续期之间没有差异,氮素添加对群落中物种的始花时间和开花持续期的负相关关系没有影响；
3.2010年群落中38个常见组分种,2011年34个常见组分种的始花期对氮素添加的响应显示,禾草类植物和双子叶杂草类植物对氮素添加的响应不同,禾草类植物的始花期推迟,而大多数双子叶植物的始花期提前。氮素添加处理中,这些常见组分种在科间的始花期变化表现为豆科始花期无变化；禾本科、莎草科、菊科的始花期推迟；毛莨科、龙胆科、玄参科的始花期提前；
4.高寒草甸植物群落中常见组分种的始花期和繁殖分配对氮素添加的响应显示,始花期和繁殖分配对氮添加的响应,不论是从响应方向上还是响应程度上都不同。禾草类物种的始花期推迟,繁殖分配下降,而大多数双子叶杂草始花期提前,繁殖分配增加；
5.高寒草甸植物群落中主要组分种的始花期和繁殖分配对氮素添加的响应变化显示,始花期提前的物种繁殖分配趋向于增大,始花期推迟的物种繁殖分配趋向于减小,氮素添加后群落中植物的始花期变化和繁殖分配变化之间成显著的正相关；
6.群落中植物的花期植株高度和茎分配与开花峰值期在不同的氮素添加处理中成正相关；叶分配与开花峰值期成负相关。在氮素处理中开花峰值期和花期植株高度、茎分配,叶分配的相关程度增加；
7.通过非度量多维尺度分析(NMDS)排序显示,对照、低氮和高氮群落之间的相似性逐渐降低,随着氮素添加量的增加,群落花期物候和群落物种组成的变化越剧烈。
The impcat of global climate and environment change on terrestrial plant ecosystems have been increasingly concerned by scientists. In particular, the response of plant phenological events to changes in global climate has been a critical concern in ecology. Different phenological time among plants have an important role in community assembly. Plant phenological events change is the result of the combined effects of multiple environmental factors. Nitrogen has traditionally been considered the primary limiting nutrient for plant growth in terrestrial ecosystems, but in the context of global climate change the rate of nutrient inputs in natural systems has been changed, which resulted in nitrogen (N) enrichment in terrestrial ecosystems. Grassland ecosystem is an important component of terrestrial ecosystems, the study on the impact of nitrogen nutrition on grassland communities are mostly concentrated in the community productivity, species diversity and community structure changes for a long time. Recent studies show that nitrogen addition could affect plant phenology, especially affect flowering phenology. Environmental change of many ecosystems throughout the globe has important ramifications for grassland communities, and the research on grassland plant phenology have received widespread attention. But the majority of plant phenology research has been conducted in lowland areas, alpine meadow ecosystems may be particularly sensitive bellwethers of environmental change. Alpine plants are limited by a short growing season and extreme environmental conditions, which making them highly vulnerable to modification of limiting factors. This suty was conducted in an alpine meadow at The Research Station of Alpine Meadow and Wetland Ecosystems of Lanzhou University, in Maqu (azi)(N33°40'E101°52', altitude3550m). We set up three levels of nitrogen addition (control, low nitrogen addition, high nitrogen addition), from2009to2011, plant phenology was observed in fixed plots and fixed time (from early May to late-September in each year), and measured plant functional traits (plant height, SLA, individual plant biomass allocation)for most common species in nitrogen addition and control communities, to study the effects of nitrogen addition on flowering phenology, functional traits and community structure of alpine meadow in the eastern Qinghai-Tibet Plateau.
We gained following results
1. In natural community, we observed58,51,55, species flowered in2009,2010,2011respectively. The flowering sequence distribution of species in the community show that, species flowering phenology started from the late April to the end of September. The frist flowering date (FFD), peak flowering date (PFD) and flowering duration day (FDD) are different among species.
2. In alpine meadow community, the first flowering date is negatively correlated with the flowering duration day in different nitrogen addition treatments.
3. We tested the effect of nitrogen addition on first flowering date for38common species in2010,34common species in2011. Nitrogen addition significantly delayed the first flowering date for all graminoid species, but accelerated first flowering date for most forb species. In response to nitrogen addition, species of Gramineae, Sedge and Compositae delayed first flowering date; species of Ranunculaceae, Gentianaceae and Scrophulariaceae accelerated first flowering date; legumes did not change in flowering time.
4. For most species, both first flowering date and reproductive allocation (RA) significantly responded to nitrogen addition. Species are different either in the response direction or in the change of the FFD and RA. All graminoid species trends to reduce RA in response to nitrogen addition, The response of forb species to nitrogen addition was more variable. Nitrogen addition significantly delayed the FFD and reduced the RA for all graminoid species, but accelerated flowering and increased RA for most forb species.
5. Species that advanced their flowering time with nitrogen addition increased their reproductive allocation, whereas those that delayed flowering time tended to decline in reproductive allocation with nitrogen addition. That changes in FFD significantly negatively correlated with the changes in RA over species in response to nitrogen, which indicated a positive relationship between flowering response and plant performance in reproductive allocation.
6. In alpine meadow community, the plant height and plant stem allocation is positively correlated with peak flowering date in different nitrogen addition treatments. This positive relevance increased in nitrogen added plots.
7. Nonmetric multidimensional scaling (NMDS) ordination of phenologically-weighed community similarity among nitrogen addition gradient showed that, the community level phenology similarity and species composition similarity were significantly decreased after nitrogen addition. With the nitrogen addition gradient the change of both community level phenology and community structure were more intense.
引文
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