Chin J Plan Ecolo ›› 2018, Vol. 42 ›› Issue (1): 50-65.doi: 10.17521/cjpe.2017.0252

• Research Articles • Previous Articles     Next Articles

Nonlinear responses of productivity and diversity of alpine meadow communities to degradation

CHEN Ning1,2,ZHANG Yang-Jian1,3,4,ZHU Jun-Tao1,*(),LI Jun-Xiang5,LIU Yao-Jie1,2,ZU Jia-Xing1,2,CONG Nan1,HUANG Ke1,WANG Li5   

  1. 1 Lhasa Plateau Ecosystem Research Station, Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101 China

    2 University of Chinese Academy of Sciences, Beijing 100190, China

    3 CAS Center for Excellence in Tibetan Plateau Earth Sciences, Beijing 100101, China

    4 College of Resources and Environment, University of Chinese Academy of Sciences, Beijing 100190, China

    5 Peking University Shenzhen Graduate School, Shenzhen 518055, China
  • Online:2018-03-08 Published:2018-01-20
  • Contact: ZHU Jun-Tao ORCID:0000-0002-3506-1247
  • Supported by:
    Supported by the National Key Research and Development Project of China(2016YFC0501802);Supported by the National Key Research and Development Project of China(2017YFA0604802);the National Natural Science Foundation of China(41571195);the National Natural Science Foundation of China(41501103)


Aims The alpine meadow degradation could have profound effects on the grassland productivity. The aim of our study is to clarify the dynamic response of community productivity and species diversity in the process of alpine meadow degradation.

Methods In the Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Northern Tibetan Grassland Ecosystem Research Station (Nagqu station), we conducted stages experiments with multiple degradation levels: control, mild degraded meadow, moderate degraded meadow, severe degraded meadow and serious sandy meadow.

Important findings The response of aboveground biomass to alpine meadow degradation showed a linear or nonlinear increased response patterns, but the belowground biomass and total biomass decreased nonlinearly. As observed in measurement of aboveground biomass, Margalef index, Simpson index, Shannon-Wiener index and Pielou evenness index also exhibit a nonlinear increased response to degradation. The results of structural equation models showed that belowground biomass has a positive relationship with soil carbon content (p < 0.05) and volume water content (p < 0.1). However, soil nutrient and soil physical properties had no significant impact on aboveground biomass (p < 0.1). Compared with soil physical properties, soil nutrition is an important factor influencing the diversity index. In our study, the nonlinear responses of productivity and diversity of alpine meadow were described by using the multiple levels of degradation in space. The results suggested that aboveground productivity cannot interpret the degree of degradation of alpine meadow, and by contrast, alpine meadow degradation should be measured by the change of plant functional groups, such as edible grasses and poisonous forbs.

Key words: alpine meadow, degeneration gradients, species diversity, productivity, nonlinear response

Fig. 1

Plot layout. A, community survey; B, measure biomass; C, measure carbon flux; D, future related researches."

Fig. 2

The community coverage of alpine meadow in different degradation stages. Different letters in figures indicate significant difference (p < 0.1). The solid lines are nonlinear fitting results. Dotted lines are linear fitting results. Control, non-degraded meadow; D1, mild degraded meadow; D2, moderate degraded meadow; D3, severe degraded meadow; D4, serious sandy meadow."

Fig. 3

The aboveground biomass of alpine meadow in different degradation stages. Different letters in figures indicate significant difference (p < 0.1). The solid lines are nonlinear fitting results. Dotted lines are linear fitting results. Control, non-degraded meadow; D1, mild degraded meadow; D2, moderate degraded meadow; D3, severe degraded meadow; D4, serious sandy meadow."

Fig. 4

The total biomass and belowground biomass of alpine meadow in different degradation stages. Different letters in figures indicate significant difference (p < 0.1). The solid lines are nonlinear fitting results. Dotted lines are linear fitting results. Control, non-degraded meadow; D1, mild degraded meadow; D2, moderate degraded meadow; D3, severe degraded meadow; D4, serious sandy meadow."

Fig. 5

The species diversity of alpine meadow in different degradation stages. Different letters in figures indicate significant difference (p < 0.1). The solid lines are nonlinear fitting results. Dotted lines are linear fitting results. Control, non-degraded meadow; D1, mild degraded meadow; D2, moderate degraded meadow; D3, severe degraded meadow; D4, serious sandy meadow."

Table 1

The soil physicochemical properties of alpine meadow in different degradation stages"

Meadow type
Soil bulk density (g?cm-3)
Mass ratio of gravel (%)
Volume ratio of gravel (%)
Soil nitrogen content (%)
Soil carbon content (%)
对照 Non-degraded 1.23 ± 0.04a 29.77 ± 7.96a 16.80 ± 7.45a 0.45 ± 0.06a 5.29 ± 0.95a
轻度退化 Mild degraded 1.26 ± 0.04a 36.51 ± 1.34a 22.19 ± 4.00a 0.50 ± 0.04a 5.76 ± 0.59a
中度退化 Moderate degraded 1.50 ± 0.02b 37.43 ± 6.04a 28.24 ± 2.60ab 0.44 ± 0.11a 5.10 ± 1.30a
重度退化 Severe degraded 1.47 ± 0.02b 47.60 ± 4.52b 32.15 ± 2.41b 0.32 ± 0.05b 3.21 ± 0.69b
极度退化 Serious degraded 1.56 ± 0.11b 49.62 ± 4.94b 33.52 ± 0.68b 0.34 ± 0.02b 3.57 ± 0.23b

Table 2

Comparison between linear and nonlinear responses of soil physicochemical properties to alpine meadow degradation"

R2 拟合模型 Fitted model
土壤容重 Soil bulk density
线性模型 Linear model 0.66*** y = -0.006x + 1.695
非线性模型 Nonlinear model 0.76*** y = 0.0002x2 + 0.012x + 1.326
砾石质量比 Mass ratio of gravel
线性模型 Linear model 0.13 y = -0.155x + 47.310
非线性模型 Nonlinear model 0.53* y = 0.023x2-2.326x + 89.838
砾石体积比 Volume ratio of gravel
线性模型 Linear model 0.44** y = 0.240x + 37.607
非线性模型 Nonlinear model 0.58** y = 0.011x2-1.300x + 58.371
土壤氮含量 Soil nitrogen content
线性模型 Linear model 0.42*** y = 0.003x + 0.264
非线性模型 Nonlinear model 0.46*** y = -0.0001x2 + 0.011x + 0.115
土壤碳含量 Soil carbon content
线性模型 Linear model 0.50*** y = 0.049x + 2.35
非线性模型 Nonlinear model 0.55*** y = -0.001x2 + 0.159x + 0.183

Fig. 6

The soil water content of alpine meadow in different degradation stages (mean ± SE). Different letters indicate significant difference (p < 0.1). Control, non-degraded meadow; D1, mild degraded meadow; D2, moderate degraded meadow; D3, severe degraded meadow; D4, serious sandy meadow."

Table 3

Comparison between linear and nonlinear responses of soil water content to alpine meadow degradation"

R2 拟合模型 Fitted model
0-7.6 cm土层 0-7.6 cm soil layer
线性模型 Linear model 0.61*** y = 0.044x + 7.396
非线性模型 Nonlinear model 0.61*** y = 7.557e0.005x
0-12 cm土层 0-12 cm soil layer
线性模型 Linear model 0.39*** y = 0.066x + 10.148
非线性模型 Nonlinear model 0.41*** y = -0.001x2 + 0.169x + 8.136

Fig. 7

Correlation analysis among soil physico-chemical properties and species diversity of alpine meadow community. Black and from lower left to upper right of the slash said two variables are related, and black from the upper left to lower right slash said negative correlation between variables. The deeper the color, the higher the degree of saturation, explanatory variable correlation; *, **, *** represent statistically significant at the 10%, 5%, 1% confidence level, respectively."

Fig. 8

The results of structural equation models showed the direct and indirect impacts of various factors on biomass and species diversity. A dotted line relationship of the arrow was not significant (p > 0.1), the solid line shows the relationship between significantly (p < 0.1), the degree of thickness line to reflect the degree of a strong relation between variables, with the arrow line Numbers are standardized path coefficient (*, p < 0.1; **, p < 0.05; ***, p < 0.01). AIC, an information criterion; AICdm, default mode; AICim, independence model; AICsm, saturated model; DF, degree of freedom; NFI, normed fit index; RMSEA, root mean square error of approximation."

Appendix table 1

Land degradation index classification in China (Qiu, 2012)"

Index classification
Number of indexes
指标 Indexes
土壤 Soil 41 土壤质地、土层厚度、含水量、团粒结构稳定性等 Soil texture, soil thickness, water content, and stability of aggregates, etc.
植被 Vegetation 11 植被盖度、植被种类、植被均匀度指数、根系深度等 Vegetation cover, vegetation species, vegetation Pielou evenness index, and root depth, etc.
社会经济 Social economy 11 经济收入水平(农民收入、消费水平)、人口数量等 Economic income level (farmer's income, consumption level) and population, etc.
气候 Climate 7 降水量、蒸发量、风速、平均气温、湿润指数等 Precipitation, evaporation, wind speed, average temperature, and wetting index, etc.
生物 Biology 7 生物量、生物多样性指数、群落类型、种群优势度 Biomass, biodiversity index, community type, and population dominance
地形地貌 Topography 6 坡度、坡向、沟谷密度、地貌类型、海拔高度 Slope, slope direction, valley density, landform type, and altitude

Appendix table 2

Land degradation index of application frequency statistics (Qiu, 2012)"

Number of indexes
指标 Indexes
> 40 41 植被盖度、经济收入水平、坡度、有机质含量 Vegetation cover, economic income level, slope, organic matter content
30-39 11 土壤质地、氮、磷和钾元素含量、含水量、土地利用类型 Soil texture, nitrogen, phosphorus and potassium content, water content, land use type
20-29 11 生产力、降水量、生物多样性、生物量、地貌类型 Productivity, precipitation, biodiversity, biomass, landform type
10-19 7 土壤类型、覆沙盖度、土层厚度、风速、蒸发量 Soil type, cover sand cover, soil thickness, wind speed, evaporation
< 10 7 土壤容重、土壤侵蚀模数、电导率、径流量、结皮厚度 Soil bulk density, soil erosion modulus, conductivity, runoff, crust thickness
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