青藏高原高寒草甸冻融退化过程中群落及化学计量特征的研究
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摘要
高寒草甸是青藏高原草地生态系统中的主体植被类型,不仅孕育着丰富的乡土生物资源,蕴藏着珍贵的高山植物抗逆基因,支撑着高原草地特色畜牧业,而且调蓄着江河水源,维系区域生态安全。然而高寒草甸的退化严重威胁着流域生态、区域生产和生计的安全。高寒草甸退化的致因主要有人为因素和自然因素,人为因素的研究主要集中于放牧和工程建设,自然因素的研究主要集中于全球变暖和鼠害。虽然冻融是高寒草甸退化的重要因素,但目前关于冻融导致高寒草甸退化的研究相对零星。本研究在青藏高原北部多年冻土区,基于冻融退化程度,选择了未退化、轻度、中度和重度退化四个阶段的样区,通过调查不同样区草地植物群落组成、总盖度、高度、多样性、地上生物量、分析优势种植物和土壤的碳氮磷含量,阐明青藏高原多年冻土区高寒草甸冻融退化过程中草地植物群落变化的特征,明晰不同退化阶段优势种植物和土壤的生态化学计量特征,从植物群落组分改良和养分限制的角度提出了退化高寒草甸的恢复策略。得到以下主要结果：
1明晰了高寒草甸冻融退化过程中群落组分的变化特征
高寒草甸植物群落组分伴随着冻融退化发生明显变化,表现为随着冻融退化程度加剧,莎草科和禾本科植物优势种地位逐渐削弱,而豆科和菊科的毒杂草组分却逐渐增加。
2查清了高寒草甸群落特征和生产力对冻融退化的响应
高寒草甸冻融退化过程中植物群落盖度和样方内丰富度先升后降,高度和地上生物量逐渐降低,而β多样性指数却升高,说明生境异质化增加。
3揭示了冻融退化对草甸优势种碳氮磷化学计量特征的影响
高寒草甸冻融退化过程中优势植物青藏苔草的C含量没有明显变化,N和P含量逐渐减小,而C：N、N：P和C：P比值却逐渐增加。轻度退化和未退化草地,优势种N：P介于14-16之间,其生长受NP共同限制,中度和重度退化阶段,优势种生长受P限制,需增施磷肥。
4明确了土壤碳氮磷化学计量特征对冻融退化的响应
高寒草甸冻融退化过程中土壤C、N和P含量逐渐降低,N：P和C：P也均逐渐减小,说明土壤养分逐渐流失,需要增施土壤养分,保证土壤养分的供给量。
5提出了退化高寒草甸恢复的策略
通过点播方式,补充扁蓿豆和垂穗披碱草,以及青海早熟禾；通过其他能源替代家畜粪便,将牛羊粪回归天然草甸,通过添施鸡粪,鸭粪和鸽粪,改善优势植物生长受P限制的现状。
Alpine meadow is the main vegetation type of grassland ecosystem in the Qinghai-Tibet Plateau, and it has rich native biological resources and precious alpine plant with anti-stresses genes, plays important roles in animal production with plateau characteristics, regulates river catchment capacity and maintains regional ecological security. However, the degeneration of alpine meadow seriously threatens ecological, productive and livelihood security throughout the rivers. The factors contributing to alpine meadow degradation mainly include artificial and natural factors, in which grazing and engineering construction are mainly artificial factors and global warming and rat trouble are mainly natural factors, neglecting freezing-thawing cycle, which is an important factor to cause alpine meadow degradation. On the basis of different degree of freezing-thawing degradation, This study investigated the composition of plant communities, cover, height, diversity and aboveground biomass and determined the content of C, N and P in dominant plants and soil at natural, light, moderate and severe degradation stages of alpine meadow caused freezing-thawing cycle for clarifying the features of alpine meadow and stoichiometry in the process of freezing-thawing cycle, which would benefit to propose the recovery strategies for the alpine meadow degradation from optimizing plant community composition and improving nutrient supply in the permafrost region of the northern Tibetan Plateau. The main results were as following,
1. The features of plant communities at natural, light, moderate and severe degradation stages was clarified in this study
This study showed that the plant community composition of alpine meadow at natural, light, moderate and severe degradation stages was different, indicating that the domination of sedge and gramineous plants gradually decreased, and poisonous and unpalatable plants in leguminosae and compositae gradually increased as the freezing-thawing degradation increased.
2. This study explained the response of community and productivity of alpine meadow to freezing-thawing degradation
Plant community cover and species richness index firstly increased and then decreased, and community height and aboveground biomass decreased gradually, while (3-diversity index increased in the process of freezing-thawing degradation, indicating that the freezing-thawing degradation increased the habitat heterogeneity in the study region.
3. This study disclosed the effect of the freezing-thawing degradation on C, N, P stoichiometry of dominant plants of alpine meadow
The results of this study showed that the freezing-thawing degradation did not affect the C content and decreased the N and P content of Carex moorcroftii, while C:N, N:P and C:P of Carex moorcroftii increased gradually, probably caused by a greater shortage of soil P in the study areas, which suggested that the application of phosphate fertilizer is beneficial to encourage the healthy growth of dominant species.
4. This study explained the respose of C, N, P stoichiometry of alpine meadow soil to freezing-thawing degradation
The C, N, P content and N:P and C:P in soil decreased with the increase of freezing-thawing degradation, indicating that the soil nutrient lost gradually in the process of freezing-thawing cycle, which suggested that the supply of soil nutrient is a basic way to restore alpine meadow degradation.
5. This study proposed the restorable strategies for alpine meadow degradation
This study proposed that the Medicago ruthenica from legumes and Elymus nutans and Qinghai bluegrass from gramineous were used to sow by mixture though bunch and reseeding pattern. The cattle and sheep manure had to return to natural meadow by finding other energy instead of animal feces. and poultry dung was used to improve the situation P limitation.
1明晰了高寒草甸冻融退化过程中群落组分的变化特征
高寒草甸植物群落组分伴随着冻融退化发生明显变化,表现为随着冻融退化程度加剧,莎草科和禾本科植物优势种地位逐渐削弱,而豆科和菊科的毒杂草组分却逐渐增加。
2查清了高寒草甸群落特征和生产力对冻融退化的响应
高寒草甸冻融退化过程中植物群落盖度和样方内丰富度先升后降,高度和地上生物量逐渐降低,而β多样性指数却升高,说明生境异质化增加。
3揭示了冻融退化对草甸优势种碳氮磷化学计量特征的影响
高寒草甸冻融退化过程中优势植物青藏苔草的C含量没有明显变化,N和P含量逐渐减小,而C：N、N：P和C：P比值却逐渐增加。轻度退化和未退化草地,优势种N：P介于14-16之间,其生长受NP共同限制,中度和重度退化阶段,优势种生长受P限制,需增施磷肥。
4明确了土壤碳氮磷化学计量特征对冻融退化的响应
高寒草甸冻融退化过程中土壤C、N和P含量逐渐降低,N：P和C：P也均逐渐减小,说明土壤养分逐渐流失,需要增施土壤养分,保证土壤养分的供给量。
5提出了退化高寒草甸恢复的策略
通过点播方式,补充扁蓿豆和垂穗披碱草,以及青海早熟禾；通过其他能源替代家畜粪便,将牛羊粪回归天然草甸,通过添施鸡粪,鸭粪和鸽粪,改善优势植物生长受P限制的现状。
Alpine meadow is the main vegetation type of grassland ecosystem in the Qinghai-Tibet Plateau, and it has rich native biological resources and precious alpine plant with anti-stresses genes, plays important roles in animal production with plateau characteristics, regulates river catchment capacity and maintains regional ecological security. However, the degeneration of alpine meadow seriously threatens ecological, productive and livelihood security throughout the rivers. The factors contributing to alpine meadow degradation mainly include artificial and natural factors, in which grazing and engineering construction are mainly artificial factors and global warming and rat trouble are mainly natural factors, neglecting freezing-thawing cycle, which is an important factor to cause alpine meadow degradation. On the basis of different degree of freezing-thawing degradation, This study investigated the composition of plant communities, cover, height, diversity and aboveground biomass and determined the content of C, N and P in dominant plants and soil at natural, light, moderate and severe degradation stages of alpine meadow caused freezing-thawing cycle for clarifying the features of alpine meadow and stoichiometry in the process of freezing-thawing cycle, which would benefit to propose the recovery strategies for the alpine meadow degradation from optimizing plant community composition and improving nutrient supply in the permafrost region of the northern Tibetan Plateau. The main results were as following,
1. The features of plant communities at natural, light, moderate and severe degradation stages was clarified in this study
This study showed that the plant community composition of alpine meadow at natural, light, moderate and severe degradation stages was different, indicating that the domination of sedge and gramineous plants gradually decreased, and poisonous and unpalatable plants in leguminosae and compositae gradually increased as the freezing-thawing degradation increased.
2. This study explained the response of community and productivity of alpine meadow to freezing-thawing degradation
Plant community cover and species richness index firstly increased and then decreased, and community height and aboveground biomass decreased gradually, while (3-diversity index increased in the process of freezing-thawing degradation, indicating that the freezing-thawing degradation increased the habitat heterogeneity in the study region.
3. This study disclosed the effect of the freezing-thawing degradation on C, N, P stoichiometry of dominant plants of alpine meadow
The results of this study showed that the freezing-thawing degradation did not affect the C content and decreased the N and P content of Carex moorcroftii, while C:N, N:P and C:P of Carex moorcroftii increased gradually, probably caused by a greater shortage of soil P in the study areas, which suggested that the application of phosphate fertilizer is beneficial to encourage the healthy growth of dominant species.
4. This study explained the respose of C, N, P stoichiometry of alpine meadow soil to freezing-thawing degradation
The C, N, P content and N:P and C:P in soil decreased with the increase of freezing-thawing degradation, indicating that the soil nutrient lost gradually in the process of freezing-thawing cycle, which suggested that the supply of soil nutrient is a basic way to restore alpine meadow degradation.
5. This study proposed the restorable strategies for alpine meadow degradation
This study proposed that the Medicago ruthenica from legumes and Elymus nutans and Qinghai bluegrass from gramineous were used to sow by mixture though bunch and reseeding pattern. The cattle and sheep manure had to return to natural meadow by finding other energy instead of animal feces. and poultry dung was used to improve the situation P limitation.
引文
1. Agren G I. Stoichiometry and nutrition of plant growth in natural communities. Annual review of ecology evolution and Systematics,2008,39:153-170.
2. Agren G I. The C:N:P stoichiometry of autotrop Hs-theory and observations. Ecology Letters,2004,7(3):185-191.
3. Bakker E, Ritchie M E, Olff H, et al.. Herbivore impact on grassland plant diversity depends on habitat productivity and herbivore size. Ecology Letters, 2006, (9):780-788.
4. Crick J C, Grime J P. Morphological plasticity andmineral nutrient capture in two herbaceous species of contrasted ecology. New Phytologist,1987,107: 403-414.
5. Chapin FSⅢ, Kedrowski R A. Seasonal changes in nitrogen and phosphorus fractions and autumn retranslocation in evergreen and deciduous taiga trees. Ecology,1983,64(2):376-391.
6. Elser JJ, Dobberful D, Mackay NA, et al.. Organism size, life history, and N:P stoichiometry:toward a unified view of cellular and ecosystem processes. BioScience,1996,46:674-684.
7. Elser JJ, Fagan W, Denno RF, et al.. Nutritional constraints in terrestrial and freshwater food webs. Nature,2000,408:578-580.
8. El ser JJ, Sterner RW, Gorokhova E, et al.. Biological stoiehiometry from genes to eeosystems. Ecology Letters,2000b, (3):540-550.
9. Gsewell S. N P ratios in terrestrial plants:variation and functional significance. New Phytologist,2004,164:243-266.
10. Guo Z G, Long R J, Niu F J, et al.. Effect of highway construction on plant diversity of grassland communities in the permafrost regions of the Qinghai-Tibet plateau. The Rangeland Journal,2007. (29):161-167.
11. Han W X, Fang J Y, Guo D L. Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China. New Phytologist,2005,168(2): 377-385.
12. Hessen DO. Stoichiometry in food webs-Lotka revisited. Oikos,1997,79: 195-200.
13. He J S, Fang J Y, Wang Z H. Stoichiometry and largescale patter ns of leaf carbon and itrogen in the grasslands of China. Oecologia,2006,149(1): 115-122.
14. Janssens F, Peeters A, Bakker J P, et al.. Relationship between soil chemical factors and grassland diversity. Plant and Soil,1998,202,69-78.
15. Michaels AF. The ratios of life. Seienee,2003,300:906-907.
16. Kooijman SALM. The stoichiometry of animal energetic. Journal of Theoretical Bilogy,1995,177:139-149.
17. Kooijman SALM. The Stoichiometry of animal energetics. Journal of Theoretical Biology,1995,177:139-14.
18. McGroddy M E, Daufresne T, Hedin L O. Scaling of C:N:P stoichiometry in forests worldwide:Implications of terrestrial Redfield type ratios. Ecology, 2004,85:2390-2401.
19. Reiners W A. Complementary models for ecosystems. American Naturalist, 1986,127:59-73.
20. Mayfield M M, Ackerly D, Daily G C. The diversity and conservation of plant reproductive and dispersal functional traits in human-dominated tropical landscapes. Ecology,2006,94:522-536.
21. Post W M, Pastor J, Zinke P J, et al.. Global patterns of soil nitrogen storage. Nature,1985,317:613-616.
22. Sterner R W, Hessen D O. Algal nutrient limitation and the nutrition of aquatic herbivores. Annual Review of Ecology and Systematic,1994,25:1-29.
23. Sterner R W, Elser JJ. Ecological Stoichiometry:The Biology of Elements from Moleculaes to the BiosPhere. Princeton University Press, Princeton.2002.
24. Tian H Q, Chen G S, Zhang C. Pattern and variation of C:N:P ratios in China'soils:a synthesis of observational data. Biogeochemistry,2010,98: 139-151.
25. Vanni M J, Flecker A S, Hood J M et al.. Stoichiometry of nutrient recycling by vertebrates in a tropical stream:linking biodiversity and ecosystem function. Ecology Letters,2002,5:285-293.
26. Zhang L X, Bai Y F, Han X G. Differential responses of N:P stoichiometry of Leymus chinensis and Carex korshinskyi to N additions in a steppe ecosystem in Nei Mongol. Acta Botanica Sinica,2004,46:259-270.
27.白永飞,许志信,李德新.内蒙古高原针茅草原群落土壤水分和碳、氮分布的小尺度空间异质性.生态学报,2002,22(8)：1215-1223.
28.包成兰,张世财.三种豆科牧草在高寒地区的生产特性比较研究.草业与畜牧,2008,(12)：13-15.
29.陈佐忠,汪诗平.中国典型草原生态系统.北京：科学出版社,2000,1-2.
30.陈全功,梁天刚,卫亚星.青海省达日县退化草地研究Ⅱ退化草地成因分与评估.草业科学,1998,7(4)：44-48.
31.陈拓,杨梅学,冯虎元,等.青藏高原北部植物叶片碳同位素组成的空间特征.冰川冻土,2003,25(1)：83-87.
32.陈生云,赵林,秦大河,等.青藏高原多年冻土区高寒草地生物量与环境因子关系的初步分析.2010,32(2)：405-413.
33. 陈荣毅,张元明,潘伯荣,等.古尔班通古特沙漠土壤养分空间分异与干扰的关系.中国沙漠,2007,27(2)：257-265.
34.程国栋,赵林.青藏高原开发中的冻土问题.第四纪研究,2000,20(6)：521-531.
35.程慧艳,王根绪,王一博.黄河源区不同植被类型覆盖下季节冻土冻融过程中的土壤温湿空间变化.兰州大学学报(自然科学版),2008,44(2)：15-21.
36.崔庆虎,蒋志刚,刘季科,等.青藏高原草地退化原因述评.草业科学,2007,24(5)：20-26.
37.单贵连,徐柱,宁发,等.围封年限对典型草原植被与土壤特征的影响.草业学报,2009,18(2)：3-10.
38.丁小慧,宫立,王东波,等.放牧对呼伦贝尔草地植物和土壤生态化学计量学特征的影响.生态学报,2012,32(15)：4722-4730.
39.丁小慧,罗淑政,刘金巍,等.呼伦贝尔草地植物群落与土壤化学计量学 特征沿经度梯度变化.生态学报,2012,32(11)：3467-3476.
40. 德科加,徐成体.高寒地区天然豆科牧草—青藏扁蓿豆的引种驯化.种子,2009,82(7)：73-75
41.伏洋,李凤霞,张国胜,等.青海省天然草地退化及其环境影响分析.冰川冻土,2007,29(4)：525-535.
42. 郭正刚,刘慧霞,王根绪,等.人类工程对青藏高原北部草地群落p多样性的影响.生态学报,2004,24(2)：384-388.
43.郭正刚,牛富俊,湛虎,等.青藏高原北部多年冻土退化过程中生态系统的变化特征.生态学报,2007,27(8)：3294-3301.
44. 郭正刚,张自和,侯扶江.河西走廊草地退化现状及其可控因素和分类综合治理.中国草地,2002,24(4)：53-58.
45.韩文轩,吴漪,汤璐瑛,等.北京及周边地区植物叶的碳氮磷元素计量特征.北京大学学报,2008,4：67-72.
46. 胡旭,王海涛,卢建国,等.干旱和半干旱区油蒿对土壤空间异质性的响应.中国沙漠,2007,27(4)：588-592.
47.侯扶江,南志标,肖金玉,等.重牧退化草地的植被、土壤及其耦合特征.应用生态学报,2002,13(8)：915-922.
48. 贾鸟敬,张映生,张自和,等.高寒地区优良豆科牧草阴山扁猎豆一些性状的研究.甘肃农业大学学报,1984,19(1)：63-69.
49.李博.中国北方草地退化及其防治对策.中国农业科学,1997,30(6)：1-9.
50. 李春吉.冻融交替作用与林木生态碳循环的关联探究.农业与技术,2012,(9)：111.
51. 李永宏.内蒙古草原操场退化模式研究及退化检测专家系统雏议.植物生态学报,1994,18(1)：68-79.
52.李午峰,唐祯敏.青藏铁路的环境与安全.安全与环境学报,2003,3(3)：65-67.
53. 李永宏.放牧空间梯度上和恢复时间演替时间梯度上羊草草原群落特征及其对应性.草原生态系统研究,1992,(4)：1-8.
54. 李世英,肖运峰.内蒙古呼盟末大木吉地区羊草草原放牧演替阶段的初步划 分.植物生态学和地植物学学报,1964,2：200-217.
55.李香真,陈位忠.不同放牧率对草原植物与土壤C、N、P含量的影响.草地学报,1998,6(2)：90-98.
56.李海英,彭红春,王启基.高寒矮嵩草草甸不同退化演替阶段植物群落地上生物量分析.草业学报,2004,13(5)：26-32.
57.李积兰,李希来,唐燕,等.三江源头“黑土滩”草地分布现状及退化特征研究.草业与畜牧,2009,(2)：6-13.
58.蔺照兰,王春梅,王汝南.冻融期温带森林土壤N20排放对模拟大气氮沉降的响应.生态环境学报,2012,21(11)：1804-1809.
59.林清,施苏华,彭平安,等.青藏高原多年冻土沉积物中埋藏植物DNA特性的初步分析.冰川冻,2003,25(1)：83-87.
60.林慧龙,任继周,傅华.草地农业生态系统中的能值分析方法评介.草业学报,2005,14(4)：1-7.
61.刘黎明,赵英伟,谢花林.我国草地退化的区域特征及其可持续利用管理.中国人口·资源与环境,2003,13(4)：46-50.
62.刘伟,王启基,王溪等.高寒草甸“黑土型”退化草地的成因及生态过程.草地学报,1999,7(4)：300-307.
63.刘钟龄,王炜,郝敦元,等.内蒙古草原退化与恢复演替机理的探讨.干旱区资源与环境,2002,(1)：84-91.
64.刘伟,王启基,王溪,等.高寒草甸“黑土型”退化草地的成因及生态过程.草地学报,1999,7(4)：300-307.
65. 罗亚勇,张宇,张静辉,等.不同退化阶段高寒草甸土壤化学计量特征.生态学杂志,2012,31(2)：254-260.
66. 李秋年.高寒草甸退化草地植物群落结构特征及物种多样性的初步分析.青海环境,2004,14(1)：30-33.
67. 马克平,刘玉明.生物群落多样性的测度方法Ⅰα多样性的测度方法(上).生物多样性,1994,2(3)：162-168.
68. 马克平,刘玉明.生物群落多样性的测度方法Ⅰα多样性的测度方法(下).生物多样性,1994,2(4)：231-239.
69.马克平,黄建辉,于顺利,等.北京东灵山地区植物群落多样性的研究Ⅱ：丰富度、均匀度和物种多样性指数.生态学报,1995,15(3)：268-277.
70. 马克平,刘灿然,刘玉明.生物群落多样性的测度方法Ⅱβ多样性的测度方法.生物多样性,1995,3(1)：38-43.
71.马玉寿,尚占环,施建军,等.黄河源区“黑土滩”退化草地群落类型多样性及其群落结研究.草业科学,2006,23(12)：6-11.
72.牛得草,董晓玉,傅华.长芒草不同季节碳氮磷生态化学计量特征.草业科学,2011,28(6)：915-920.
73. 牛亚菲.青藏高原生态环境问题研究.地理科学进展,1999,18(2)：163-171.
74.裴志勇,欧阳华,周才平.青藏高原高寒草原碳排放及其迁移过程研究.生态学报,2003,23(2)：231-236.
75. 戚秋慧.羊草草地的群落结构与生物量关系的初步研究.草地生态系统研究(第1集).北京：科学出版社,1985,38-47.
76.仁青吉,武高林,任国华.放牧强度对青藏高原东部高寒草甸植物群落特征的影响.草业学报,2009,18(5)：256-261.
77.孙海松.青海高寒草地资源的可持续利用.草业科学,1998,15(3)：1-3.
78. 尚占环,龙瑞军.青藏高原“黑土型”退化草地成因与恢复.生态学杂志,2005,24(6)：652-656.
79.王根绪,程国栋,沈永平,等.江河源区的生态环境变化及其综合保护研究.兰州：兰州大学出版社,2001,29-35.
80. 王根绪,李玉其,程国栋.40年来江河源区气候变化特征及其生态环境效应.冰川冻土,2001,23(4)：346-352.
81. 王德利,吕新民,罗卫生.不同放牧密度对草原退化群落恢复演替的研究.Ⅰ.退化草原的基本特征和恢复常规演替动力.植物生态学报,1996,20(5)：449-460.
82. 王仁忠,李建东.松嫩平原碱化羊草草地放牧空间演替规律的研究.应用生态学报,1995,6(3)：277-281.
83. 汪诗平.内蒙古典型草原糙隐子草种群补偿性生长机制的研究.植物学报, 2001,43(4)：413-418.
84.汪诗平,李永宏,王艳芬,等.不同放牧率对内蒙古冷蒿草原演替规律的影响.草地学报,1998,6(4)：299-305.
85.王长庭,龙瑞军,王启兰,等.江源区高寒草甸不同退化演替阶段土壤有机碳和微生物量碳的变化.2008,14(2)：225-230.
86.王文颖,王启基.高寒嵩草草甸退化生态系统植物群落结构特征及物种多样性分析.草业学报,2001,1(3)：8-14.
87.王秉玺,曹先.西宁曹家堡机场飞行区土壤养分特征的初步研究.中国沙漠,2008,28(1)：137-141.
88.汪建飞,董彩霞,谢越,等.铵硝比和磷素营养对菠菜生长、氮素吸收和相关酶活性的影响.土壤学报,2006,43(6)：954-960.
89.王会利,毕利东,张斌.退化红壤马尾松恢复林地土壤微生物生物量变化及其控制因素研究.土壤学报,2008,45(2)：313-320.
90.魏兴琥,杨萍,李森,等.超载放牧与那曲地区高山嵩草草甸植被退化及其退化指标的探讨.草业学报,2005,14(3)：41-49.
91. 王根绪,胡宏昌,王一博,等.青藏高原多年冻土区典型高寒草地生物量对气候变化的响应.冰川冻土,2007,29(5)：671-679.
92.汪诗平,李永宏,王艳芬,等.不同放牧率下冷蒿小禾草草原放牧演替规律与数量分析.草地学报,1998,6(4)：299-305.
93. 肖运峰,李世英.羊草草原退化演替及其退化原因.中国草地,1980,(3)：20-27.
94.杨建平,丁永建,陈仁升,等.长江黄河源区多年冻土变化及其生态环境效应.山地学报,2004,22(3)：278-285.
95.杨力军,李希来,石德军,等.青藏高原“黑土滩”退化草地植被演替规律的研究.青海草业,2005,14(1)：2-5.
96.杨惠敏,王冬梅.草-环境系统植物碳氮磷生态化学计量学及其对环境因子的响应研究进展.草业学报,2011,20(2)：244-252.
97.杨玉海,陈亚宁,李卫红,等.准噶尔盆地西北缘新垦绿洲土地利用对土壤养分变化的影响.2008,28(1)：94-100.
98. 银晓瑞,梁存柱,王立新,等.内蒙古典型草原不同恢复演替阶段植物养分化学计量学.植物生态学报,2010,34(1)：39-47.
99.于健龙,石红霄,李剑,等.不同退化程度高寒嵩草草甸基况的初步研究.草业科学,2010,27(9)：115-118.
100. 阎凯,付登高,何峰,等.滇池流域富磷区不同土壤磷水平下植物叶片的养分化学计量特征.植物生态学报,2011,35(4)：353-361.
101.张静,李希来,季刚.三江源地区不同退化草地群落特征分析.安徽农业科学,2008,36(18)：7738-7740.
102.张自和,郭正刚,吴素琴.西部高寒地区草业面临的问题与可持续发展.草业学报,2002,11(3)：29-33.
103.张金霞,曹广民,周党卫.高寒矮嵩草草甸大气-土壤-植被-动物系统碳素储量及碳素循环.生态学报,2003,3(4)：627-635.
104.张文彦,樊江文,钟华平,等.中国典型草原优势植物功能群氮磷化学计量学特征研究.草地学报,2010,18(4)：503-509.
105.张金屯,焦蓉.关帝山神尾沟森林群落木本植物种间联结性与相关性研究.植物研究,2003,23(4)：458-463.
106.赵玉红,魏学红,苗彦军,等.藏北高寒草甸不同退化阶段植物群落特征及其繁殖分配研究.草地学报,2012,20(2)：221-228.
107.赵新全,张耀生,周兴民.高寒草甸畜牧业可持续发展：理论与实践.资源科学,2000,22(4)：50-61.
108.赵雨霞,芮大虎,铃木辉之.寒区植被边坡冻融特性现场试验研究.低温建筑技术,2011,(7)：98-100.
109.周兴民.中国嵩草草甸.北京：科学出版社,2001：7-11.
110.周万海,冯瑞章,满元荣.黄河源区不同退化程度高寒草地土壤特征研究.2008,(4)：24-28.
111.昭和斯图,祁永.内蒙古短花针茅草原放牧退化序列的研究.中国草地,1987,(1)：29-35.
112.曾德慧,陈广生.生态化学计量学：复杂生命系统奥秘的探索.植物生态学报,2005,29(6)：1007-1019.
2. Agren G I. The C:N:P stoichiometry of autotrop Hs-theory and observations. Ecology Letters,2004,7(3):185-191.
3. Bakker E, Ritchie M E, Olff H, et al.. Herbivore impact on grassland plant diversity depends on habitat productivity and herbivore size. Ecology Letters, 2006, (9):780-788.
4. Crick J C, Grime J P. Morphological plasticity andmineral nutrient capture in two herbaceous species of contrasted ecology. New Phytologist,1987,107: 403-414.
5. Chapin FSⅢ, Kedrowski R A. Seasonal changes in nitrogen and phosphorus fractions and autumn retranslocation in evergreen and deciduous taiga trees. Ecology,1983,64(2):376-391.
6. Elser JJ, Dobberful D, Mackay NA, et al.. Organism size, life history, and N:P stoichiometry:toward a unified view of cellular and ecosystem processes. BioScience,1996,46:674-684.
7. Elser JJ, Fagan W, Denno RF, et al.. Nutritional constraints in terrestrial and freshwater food webs. Nature,2000,408:578-580.
8. El ser JJ, Sterner RW, Gorokhova E, et al.. Biological stoiehiometry from genes to eeosystems. Ecology Letters,2000b, (3):540-550.
9. Gsewell S. N P ratios in terrestrial plants:variation and functional significance. New Phytologist,2004,164:243-266.
10. Guo Z G, Long R J, Niu F J, et al.. Effect of highway construction on plant diversity of grassland communities in the permafrost regions of the Qinghai-Tibet plateau. The Rangeland Journal,2007. (29):161-167.
11. Han W X, Fang J Y, Guo D L. Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China. New Phytologist,2005,168(2): 377-385.
12. Hessen DO. Stoichiometry in food webs-Lotka revisited. Oikos,1997,79: 195-200.
13. He J S, Fang J Y, Wang Z H. Stoichiometry and largescale patter ns of leaf carbon and itrogen in the grasslands of China. Oecologia,2006,149(1): 115-122.
14. Janssens F, Peeters A, Bakker J P, et al.. Relationship between soil chemical factors and grassland diversity. Plant and Soil,1998,202,69-78.
15. Michaels AF. The ratios of life. Seienee,2003,300:906-907.
16. Kooijman SALM. The stoichiometry of animal energetic. Journal of Theoretical Bilogy,1995,177:139-149.
17. Kooijman SALM. The Stoichiometry of animal energetics. Journal of Theoretical Biology,1995,177:139-14.
18. McGroddy M E, Daufresne T, Hedin L O. Scaling of C:N:P stoichiometry in forests worldwide:Implications of terrestrial Redfield type ratios. Ecology, 2004,85:2390-2401.
19. Reiners W A. Complementary models for ecosystems. American Naturalist, 1986,127:59-73.
20. Mayfield M M, Ackerly D, Daily G C. The diversity and conservation of plant reproductive and dispersal functional traits in human-dominated tropical landscapes. Ecology,2006,94:522-536.
21. Post W M, Pastor J, Zinke P J, et al.. Global patterns of soil nitrogen storage. Nature,1985,317:613-616.
22. Sterner R W, Hessen D O. Algal nutrient limitation and the nutrition of aquatic herbivores. Annual Review of Ecology and Systematic,1994,25:1-29.
23. Sterner R W, Elser JJ. Ecological Stoichiometry:The Biology of Elements from Moleculaes to the BiosPhere. Princeton University Press, Princeton.2002.
24. Tian H Q, Chen G S, Zhang C. Pattern and variation of C:N:P ratios in China'soils:a synthesis of observational data. Biogeochemistry,2010,98: 139-151.
25. Vanni M J, Flecker A S, Hood J M et al.. Stoichiometry of nutrient recycling by vertebrates in a tropical stream:linking biodiversity and ecosystem function. Ecology Letters,2002,5:285-293.
26. Zhang L X, Bai Y F, Han X G. Differential responses of N:P stoichiometry of Leymus chinensis and Carex korshinskyi to N additions in a steppe ecosystem in Nei Mongol. Acta Botanica Sinica,2004,46:259-270.
27.白永飞,许志信,李德新.内蒙古高原针茅草原群落土壤水分和碳、氮分布的小尺度空间异质性.生态学报,2002,22(8)：1215-1223.
28.包成兰,张世财.三种豆科牧草在高寒地区的生产特性比较研究.草业与畜牧,2008,(12)：13-15.
29.陈佐忠,汪诗平.中国典型草原生态系统.北京：科学出版社,2000,1-2.
30.陈全功,梁天刚,卫亚星.青海省达日县退化草地研究Ⅱ退化草地成因分与评估.草业科学,1998,7(4)：44-48.
31.陈拓,杨梅学,冯虎元,等.青藏高原北部植物叶片碳同位素组成的空间特征.冰川冻土,2003,25(1)：83-87.
32.陈生云,赵林,秦大河,等.青藏高原多年冻土区高寒草地生物量与环境因子关系的初步分析.2010,32(2)：405-413.
33. 陈荣毅,张元明,潘伯荣,等.古尔班通古特沙漠土壤养分空间分异与干扰的关系.中国沙漠,2007,27(2)：257-265.
34.程国栋,赵林.青藏高原开发中的冻土问题.第四纪研究,2000,20(6)：521-531.
35.程慧艳,王根绪,王一博.黄河源区不同植被类型覆盖下季节冻土冻融过程中的土壤温湿空间变化.兰州大学学报(自然科学版),2008,44(2)：15-21.
36.崔庆虎,蒋志刚,刘季科,等.青藏高原草地退化原因述评.草业科学,2007,24(5)：20-26.
37.单贵连,徐柱,宁发,等.围封年限对典型草原植被与土壤特征的影响.草业学报,2009,18(2)：3-10.
38.丁小慧,宫立,王东波,等.放牧对呼伦贝尔草地植物和土壤生态化学计量学特征的影响.生态学报,2012,32(15)：4722-4730.
39.丁小慧,罗淑政,刘金巍,等.呼伦贝尔草地植物群落与土壤化学计量学 特征沿经度梯度变化.生态学报,2012,32(11)：3467-3476.
40. 德科加,徐成体.高寒地区天然豆科牧草—青藏扁蓿豆的引种驯化.种子,2009,82(7)：73-75
41.伏洋,李凤霞,张国胜,等.青海省天然草地退化及其环境影响分析.冰川冻土,2007,29(4)：525-535.
42. 郭正刚,刘慧霞,王根绪,等.人类工程对青藏高原北部草地群落p多样性的影响.生态学报,2004,24(2)：384-388.
43.郭正刚,牛富俊,湛虎,等.青藏高原北部多年冻土退化过程中生态系统的变化特征.生态学报,2007,27(8)：3294-3301.
44. 郭正刚,张自和,侯扶江.河西走廊草地退化现状及其可控因素和分类综合治理.中国草地,2002,24(4)：53-58.
45.韩文轩,吴漪,汤璐瑛,等.北京及周边地区植物叶的碳氮磷元素计量特征.北京大学学报,2008,4：67-72.
46. 胡旭,王海涛,卢建国,等.干旱和半干旱区油蒿对土壤空间异质性的响应.中国沙漠,2007,27(4)：588-592.
47.侯扶江,南志标,肖金玉,等.重牧退化草地的植被、土壤及其耦合特征.应用生态学报,2002,13(8)：915-922.
48. 贾鸟敬,张映生,张自和,等.高寒地区优良豆科牧草阴山扁猎豆一些性状的研究.甘肃农业大学学报,1984,19(1)：63-69.
49.李博.中国北方草地退化及其防治对策.中国农业科学,1997,30(6)：1-9.
50. 李春吉.冻融交替作用与林木生态碳循环的关联探究.农业与技术,2012,(9)：111.
51. 李永宏.内蒙古草原操场退化模式研究及退化检测专家系统雏议.植物生态学报,1994,18(1)：68-79.
52.李午峰,唐祯敏.青藏铁路的环境与安全.安全与环境学报,2003,3(3)：65-67.
53. 李永宏.放牧空间梯度上和恢复时间演替时间梯度上羊草草原群落特征及其对应性.草原生态系统研究,1992,(4)：1-8.
54. 李世英,肖运峰.内蒙古呼盟末大木吉地区羊草草原放牧演替阶段的初步划 分.植物生态学和地植物学学报,1964,2：200-217.
55.李香真,陈位忠.不同放牧率对草原植物与土壤C、N、P含量的影响.草地学报,1998,6(2)：90-98.
56.李海英,彭红春,王启基.高寒矮嵩草草甸不同退化演替阶段植物群落地上生物量分析.草业学报,2004,13(5)：26-32.
57.李积兰,李希来,唐燕,等.三江源头“黑土滩”草地分布现状及退化特征研究.草业与畜牧,2009,(2)：6-13.
58.蔺照兰,王春梅,王汝南.冻融期温带森林土壤N20排放对模拟大气氮沉降的响应.生态环境学报,2012,21(11)：1804-1809.
59.林清,施苏华,彭平安,等.青藏高原多年冻土沉积物中埋藏植物DNA特性的初步分析.冰川冻,2003,25(1)：83-87.
60.林慧龙,任继周,傅华.草地农业生态系统中的能值分析方法评介.草业学报,2005,14(4)：1-7.
61.刘黎明,赵英伟,谢花林.我国草地退化的区域特征及其可持续利用管理.中国人口·资源与环境,2003,13(4)：46-50.
62.刘伟,王启基,王溪等.高寒草甸“黑土型”退化草地的成因及生态过程.草地学报,1999,7(4)：300-307.
63.刘钟龄,王炜,郝敦元,等.内蒙古草原退化与恢复演替机理的探讨.干旱区资源与环境,2002,(1)：84-91.
64.刘伟,王启基,王溪,等.高寒草甸“黑土型”退化草地的成因及生态过程.草地学报,1999,7(4)：300-307.
65. 罗亚勇,张宇,张静辉,等.不同退化阶段高寒草甸土壤化学计量特征.生态学杂志,2012,31(2)：254-260.
66. 李秋年.高寒草甸退化草地植物群落结构特征及物种多样性的初步分析.青海环境,2004,14(1)：30-33.
67. 马克平,刘玉明.生物群落多样性的测度方法Ⅰα多样性的测度方法(上).生物多样性,1994,2(3)：162-168.
68. 马克平,刘玉明.生物群落多样性的测度方法Ⅰα多样性的测度方法(下).生物多样性,1994,2(4)：231-239.
69.马克平,黄建辉,于顺利,等.北京东灵山地区植物群落多样性的研究Ⅱ：丰富度、均匀度和物种多样性指数.生态学报,1995,15(3)：268-277.
70. 马克平,刘灿然,刘玉明.生物群落多样性的测度方法Ⅱβ多样性的测度方法.生物多样性,1995,3(1)：38-43.
71.马玉寿,尚占环,施建军,等.黄河源区“黑土滩”退化草地群落类型多样性及其群落结研究.草业科学,2006,23(12)：6-11.
72.牛得草,董晓玉,傅华.长芒草不同季节碳氮磷生态化学计量特征.草业科学,2011,28(6)：915-920.
73. 牛亚菲.青藏高原生态环境问题研究.地理科学进展,1999,18(2)：163-171.
74.裴志勇,欧阳华,周才平.青藏高原高寒草原碳排放及其迁移过程研究.生态学报,2003,23(2)：231-236.
75. 戚秋慧.羊草草地的群落结构与生物量关系的初步研究.草地生态系统研究(第1集).北京：科学出版社,1985,38-47.
76.仁青吉,武高林,任国华.放牧强度对青藏高原东部高寒草甸植物群落特征的影响.草业学报,2009,18(5)：256-261.
77.孙海松.青海高寒草地资源的可持续利用.草业科学,1998,15(3)：1-3.
78. 尚占环,龙瑞军.青藏高原“黑土型”退化草地成因与恢复.生态学杂志,2005,24(6)：652-656.
79.王根绪,程国栋,沈永平,等.江河源区的生态环境变化及其综合保护研究.兰州：兰州大学出版社,2001,29-35.
80. 王根绪,李玉其,程国栋.40年来江河源区气候变化特征及其生态环境效应.冰川冻土,2001,23(4)：346-352.
81. 王德利,吕新民,罗卫生.不同放牧密度对草原退化群落恢复演替的研究.Ⅰ.退化草原的基本特征和恢复常规演替动力.植物生态学报,1996,20(5)：449-460.
82. 王仁忠,李建东.松嫩平原碱化羊草草地放牧空间演替规律的研究.应用生态学报,1995,6(3)：277-281.
83. 汪诗平.内蒙古典型草原糙隐子草种群补偿性生长机制的研究.植物学报, 2001,43(4)：413-418.
84.汪诗平,李永宏,王艳芬,等.不同放牧率对内蒙古冷蒿草原演替规律的影响.草地学报,1998,6(4)：299-305.
85.王长庭,龙瑞军,王启兰,等.江源区高寒草甸不同退化演替阶段土壤有机碳和微生物量碳的变化.2008,14(2)：225-230.
86.王文颖,王启基.高寒嵩草草甸退化生态系统植物群落结构特征及物种多样性分析.草业学报,2001,1(3)：8-14.
87.王秉玺,曹先.西宁曹家堡机场飞行区土壤养分特征的初步研究.中国沙漠,2008,28(1)：137-141.
88.汪建飞,董彩霞,谢越,等.铵硝比和磷素营养对菠菜生长、氮素吸收和相关酶活性的影响.土壤学报,2006,43(6)：954-960.
89.王会利,毕利东,张斌.退化红壤马尾松恢复林地土壤微生物生物量变化及其控制因素研究.土壤学报,2008,45(2)：313-320.
90.魏兴琥,杨萍,李森,等.超载放牧与那曲地区高山嵩草草甸植被退化及其退化指标的探讨.草业学报,2005,14(3)：41-49.
91. 王根绪,胡宏昌,王一博,等.青藏高原多年冻土区典型高寒草地生物量对气候变化的响应.冰川冻土,2007,29(5)：671-679.
92.汪诗平,李永宏,王艳芬,等.不同放牧率下冷蒿小禾草草原放牧演替规律与数量分析.草地学报,1998,6(4)：299-305.
93. 肖运峰,李世英.羊草草原退化演替及其退化原因.中国草地,1980,(3)：20-27.
94.杨建平,丁永建,陈仁升,等.长江黄河源区多年冻土变化及其生态环境效应.山地学报,2004,22(3)：278-285.
95.杨力军,李希来,石德军,等.青藏高原“黑土滩”退化草地植被演替规律的研究.青海草业,2005,14(1)：2-5.
96.杨惠敏,王冬梅.草-环境系统植物碳氮磷生态化学计量学及其对环境因子的响应研究进展.草业学报,2011,20(2)：244-252.
97.杨玉海,陈亚宁,李卫红,等.准噶尔盆地西北缘新垦绿洲土地利用对土壤养分变化的影响.2008,28(1)：94-100.
98. 银晓瑞,梁存柱,王立新,等.内蒙古典型草原不同恢复演替阶段植物养分化学计量学.植物生态学报,2010,34(1)：39-47.
99.于健龙,石红霄,李剑,等.不同退化程度高寒嵩草草甸基况的初步研究.草业科学,2010,27(9)：115-118.
100. 阎凯,付登高,何峰,等.滇池流域富磷区不同土壤磷水平下植物叶片的养分化学计量特征.植物生态学报,2011,35(4)：353-361.
101.张静,李希来,季刚.三江源地区不同退化草地群落特征分析.安徽农业科学,2008,36(18)：7738-7740.
102.张自和,郭正刚,吴素琴.西部高寒地区草业面临的问题与可持续发展.草业学报,2002,11(3)：29-33.
103.张金霞,曹广民,周党卫.高寒矮嵩草草甸大气-土壤-植被-动物系统碳素储量及碳素循环.生态学报,2003,3(4)：627-635.
104.张文彦,樊江文,钟华平,等.中国典型草原优势植物功能群氮磷化学计量学特征研究.草地学报,2010,18(4)：503-509.
105.张金屯,焦蓉.关帝山神尾沟森林群落木本植物种间联结性与相关性研究.植物研究,2003,23(4)：458-463.
106.赵玉红,魏学红,苗彦军,等.藏北高寒草甸不同退化阶段植物群落特征及其繁殖分配研究.草地学报,2012,20(2)：221-228.
107.赵新全,张耀生,周兴民.高寒草甸畜牧业可持续发展：理论与实践.资源科学,2000,22(4)：50-61.
108.赵雨霞,芮大虎,铃木辉之.寒区植被边坡冻融特性现场试验研究.低温建筑技术,2011,(7)：98-100.
109.周兴民.中国嵩草草甸.北京：科学出版社,2001：7-11.
110.周万海,冯瑞章,满元荣.黄河源区不同退化程度高寒草地土壤特征研究.2008,(4)：24-28.
111.昭和斯图,祁永.内蒙古短花针茅草原放牧退化序列的研究.中国草地,1987,(1)：29-35.
112.曾德慧,陈广生.生态化学计量学：复杂生命系统奥秘的探索.植物生态学报,2005,29(6)：1007-1019.
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