增温和氮素添加对松嫩草原羊草群落结构和功能的影响
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摘要
随着工业气体的排放和温室效应的加剧，全球气候变化已经成为不容置疑的事实。在环境温度和氮素改变的条件下，植物的生长以及生理生态特性将产生巨大的变化。松嫩草原不仅是我国面积最大的草甸草原，也是我国最具代表性的草地类型之一，在反映与调节全球气候变化过程中具有重要的作用。本研究以广泛分布于松嫩草原的羊草草甸为对象，在2006年到2009年采用红外线加热仪模拟增温效应、用氮素添加的方法模拟氮素沉降，研究了羊草草甸的群落结构特征、生产力、建群种物候期及种子生产能力与种子质量的变化规律，为预测全球气候变化对草地生态系统的影响提供科学依据。结果表明：
（1）增温和氮素添加处理后，物种组成发生了较为明显的改变，增温1.7℃使物种数量增加，施氮使物种数量减少。同时，群落的特征值发生明显变化，增温使禾草重要值降低，非禾草重要值升高，而氮素的效应与增温相反；增温后物种的丰富度和均匀度有所升高，而氮素导致物种丰富度和均匀度降低。最终，增温使物种多样性指数升高，氮素使多样性指数降低。
（2）增温对地上生物量生产有不利影响，对地下生物量的生产有利。在降水较多的年份，增温对地上生物量生产的抑制作用较小，而在干旱年份会产生明显的抑制作用。不论降雨的多少，氮素对羊草群落的地上和地下生物量生产均有明显的促进作用，二者的交互作用也使地上和地下生物量明显增加。
（3）增温对羊草的生殖物候期有明显的影响，表现为抽穗、开花和结实期提前，而种子成熟期没有明显的变化；对氮素添加的响应表现为抽穗、开花和结实期有延后的趋势，而种子成熟期会明显的延迟。
（4）增温对羊草的种子生产在不同的季节和降雨条件下有着不同的影响，果后营养期的增温会导致翌年单位面积的抽穗数量减少，开花期至种子成熟期这一阶段，由于降雨量较多，增温反而有利于种子的发育；增温会减少羊草单位面积的种子数和小种子的比例及小种子的发芽率和发芽速率，增加大种子的比例；但大种子的发芽率和发芽速率没有变化。这意味着在全球持续变暖的条件下羊草降低了种子生产能力，尽管大种子数量和比例的增加能够补偿一部分由于总籽粒数减少造成的成苗数量的减少，但是，由于单位面积可发芽的种子数量的减少，羊草种群的扩展会受到限制。
氮素添加能够明显的促进羊草的种子生产，使单位面积的可发芽种子数增加；同时，能够明显的减少羊草种子中小种子的比例，增加大种子的比例，并使不同大小种子的发芽率、发芽速率以及所产生幼苗的生长均受到促进。这暗示着氮素添加对羊草的有性生殖过程有促进作用，有利于羊草占据更多的生长空间。
综上所述，我们发现增温尽管维持了较高的生物多样性，但由于增温造成的干旱效应，生产力明显降低；氮素添加造成一些对氮素敏感物种的大量繁殖促进了净初级生产力，但导致群落的物种数量减少。即羊草群落的生长和结构对模拟增温和氮素添加的响应是敏感而迅速的，结果为解释和预测草原生态系统在全球变化背景下的变化趋势提供了依据。
Over the coming decades the signal of anthropogenic climate change is expectedto become increasingly apparent, with accelerated production of greenhouse gasesfurther warming the earth’s surface and modifying nitrogen deposition patterns.Global warming and nitrogen deposition may interact in their effects on the diversityand composition of natural communities. The two related goals of studying bioticresponses to climate change are to understand changes in species distributions andabundances and to understand ecosystem feedbacks to climate change. Songnenmeadow steppe is the largest, most representative grassland in China, which plays animportant role on response and regulation to global climate change. This studyevaluated Leymus chinensis plant communities, a dominant perennial grass widelydistributed in the Songnen plain, as a model to investigate the effect of increasingambient temperature on community composition, diversity, biomass, dominantspecies phenology, seed production from2006to2009. Our study showed potentialimportance of understanding ecosystem responses to climate change and nitrogendeposition in terms of species responses. The main results can be summarized asfollows:
(1) Warming and nitrogen addition may indirectly affect species compositionthrough species numbers variation. Warming increased species numbers, whereasnitrogen addition decreased it. At the same time, community characteristics changedobviously. Warming reduced important value of grass, while enhanced the importantvalue of herbs, the opposite result occurred by nitrogen addition. Both speciesrichness and evenness increased by warming, in contrast, the effect of nitrogenaddition decreased them. Finally, diversity index showed positive dependence onwarming but negative dependence on nitrogen addition.
(2) We found significant positive responses of warming to the undergroundbiomass but negative responses to the aboveground biomass. In particular, for theaboveground biomass, a negative impact in dry years and little affects in moreprecipitation years were found. Moreover, nitrogen addition significantly enhancedboth aboveground biomass and underground biomass regardless of rainfall. Theinteraction between them is the same result.
(3) Warming accelerated Leymus chinensis spiking stage, flowering stage andseed-setting stage, but had no effect on seed maturity stage. Nitrogen addition hadlittle effect on phenology score of spiking stage, but obviously delayed the floweringstage and setting stage, and seed maturing stage.
(4) Warming reduced the mean seed number per square meter and potentialgerminating seeds, and resulted in decreased proportion of light weight seeds butincreased the proportion of heavy weight seeds. The seed germination, germinationrate, and seedling shoot of light weight seeds were reduced by warming but the sameeffects were not observed in heavy weight seeds. Despite of increased proportion ofheavy weight seeds compensated decreased plant number because of reduced seednumber per square meter; a reduction in the potential germination seed number byincreased temperature may imply that climate warming will constrain populationexpansion depending on the viability implications for seeds under further globalclimate warming.
Nitrogen deposition cause a decreases in proportion of light weight seeds butresult an increases in heavy weight seeds, which lead to an improving of seedproduction, seed germination success, germination rate and growth of correspondingshoots in Leymus chinensis. These results implied that continued nitrogen depositionwill promote the sexual reproduction process and improve the growth of Leymuschinensis by occupy more space.
Overall, warming leaded a decrease in grassland productivity due to the soildrought from warming depspite warming maintained a high biodiversity. Nitrogendeposition leaded an increase in primary productivity, but a decrease of speciesnumber. And it was therefore that the growth and structure of the Leymus chinensiscommunity responded to global warming and nitrogen deposition were sensitive,which might provide a theoretical basis to explain and predict the trends of grasslandecosystems faced with further global changing.
（1）增温和氮素添加处理后，物种组成发生了较为明显的改变，增温1.7℃使物种数量增加，施氮使物种数量减少。同时，群落的特征值发生明显变化，增温使禾草重要值降低，非禾草重要值升高，而氮素的效应与增温相反；增温后物种的丰富度和均匀度有所升高，而氮素导致物种丰富度和均匀度降低。最终，增温使物种多样性指数升高，氮素使多样性指数降低。
（2）增温对地上生物量生产有不利影响，对地下生物量的生产有利。在降水较多的年份，增温对地上生物量生产的抑制作用较小，而在干旱年份会产生明显的抑制作用。不论降雨的多少，氮素对羊草群落的地上和地下生物量生产均有明显的促进作用，二者的交互作用也使地上和地下生物量明显增加。
（3）增温对羊草的生殖物候期有明显的影响，表现为抽穗、开花和结实期提前，而种子成熟期没有明显的变化；对氮素添加的响应表现为抽穗、开花和结实期有延后的趋势，而种子成熟期会明显的延迟。
（4）增温对羊草的种子生产在不同的季节和降雨条件下有着不同的影响，果后营养期的增温会导致翌年单位面积的抽穗数量减少，开花期至种子成熟期这一阶段，由于降雨量较多，增温反而有利于种子的发育；增温会减少羊草单位面积的种子数和小种子的比例及小种子的发芽率和发芽速率，增加大种子的比例；但大种子的发芽率和发芽速率没有变化。这意味着在全球持续变暖的条件下羊草降低了种子生产能力，尽管大种子数量和比例的增加能够补偿一部分由于总籽粒数减少造成的成苗数量的减少，但是，由于单位面积可发芽的种子数量的减少，羊草种群的扩展会受到限制。
氮素添加能够明显的促进羊草的种子生产，使单位面积的可发芽种子数增加；同时，能够明显的减少羊草种子中小种子的比例，增加大种子的比例，并使不同大小种子的发芽率、发芽速率以及所产生幼苗的生长均受到促进。这暗示着氮素添加对羊草的有性生殖过程有促进作用，有利于羊草占据更多的生长空间。
综上所述，我们发现增温尽管维持了较高的生物多样性，但由于增温造成的干旱效应，生产力明显降低；氮素添加造成一些对氮素敏感物种的大量繁殖促进了净初级生产力，但导致群落的物种数量减少。即羊草群落的生长和结构对模拟增温和氮素添加的响应是敏感而迅速的，结果为解释和预测草原生态系统在全球变化背景下的变化趋势提供了依据。
Over the coming decades the signal of anthropogenic climate change is expectedto become increasingly apparent, with accelerated production of greenhouse gasesfurther warming the earth’s surface and modifying nitrogen deposition patterns.Global warming and nitrogen deposition may interact in their effects on the diversityand composition of natural communities. The two related goals of studying bioticresponses to climate change are to understand changes in species distributions andabundances and to understand ecosystem feedbacks to climate change. Songnenmeadow steppe is the largest, most representative grassland in China, which plays animportant role on response and regulation to global climate change. This studyevaluated Leymus chinensis plant communities, a dominant perennial grass widelydistributed in the Songnen plain, as a model to investigate the effect of increasingambient temperature on community composition, diversity, biomass, dominantspecies phenology, seed production from2006to2009. Our study showed potentialimportance of understanding ecosystem responses to climate change and nitrogendeposition in terms of species responses. The main results can be summarized asfollows:
(1) Warming and nitrogen addition may indirectly affect species compositionthrough species numbers variation. Warming increased species numbers, whereasnitrogen addition decreased it. At the same time, community characteristics changedobviously. Warming reduced important value of grass, while enhanced the importantvalue of herbs, the opposite result occurred by nitrogen addition. Both speciesrichness and evenness increased by warming, in contrast, the effect of nitrogenaddition decreased them. Finally, diversity index showed positive dependence onwarming but negative dependence on nitrogen addition.
(2) We found significant positive responses of warming to the undergroundbiomass but negative responses to the aboveground biomass. In particular, for theaboveground biomass, a negative impact in dry years and little affects in moreprecipitation years were found. Moreover, nitrogen addition significantly enhancedboth aboveground biomass and underground biomass regardless of rainfall. Theinteraction between them is the same result.
(3) Warming accelerated Leymus chinensis spiking stage, flowering stage andseed-setting stage, but had no effect on seed maturity stage. Nitrogen addition hadlittle effect on phenology score of spiking stage, but obviously delayed the floweringstage and setting stage, and seed maturing stage.
(4) Warming reduced the mean seed number per square meter and potentialgerminating seeds, and resulted in decreased proportion of light weight seeds butincreased the proportion of heavy weight seeds. The seed germination, germinationrate, and seedling shoot of light weight seeds were reduced by warming but the sameeffects were not observed in heavy weight seeds. Despite of increased proportion ofheavy weight seeds compensated decreased plant number because of reduced seednumber per square meter; a reduction in the potential germination seed number byincreased temperature may imply that climate warming will constrain populationexpansion depending on the viability implications for seeds under further globalclimate warming.
Nitrogen deposition cause a decreases in proportion of light weight seeds butresult an increases in heavy weight seeds, which lead to an improving of seedproduction, seed germination success, germination rate and growth of correspondingshoots in Leymus chinensis. These results implied that continued nitrogen depositionwill promote the sexual reproduction process and improve the growth of Leymuschinensis by occupy more space.
Overall, warming leaded a decrease in grassland productivity due to the soildrought from warming depspite warming maintained a high biodiversity. Nitrogendeposition leaded an increase in primary productivity, but a decrease of speciesnumber. And it was therefore that the growth and structure of the Leymus chinensiscommunity responded to global warming and nitrogen deposition were sensitive,which might provide a theoretical basis to explain and predict the trends of grasslandecosystems faced with further global changing.
引文
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