Chin J Plan Ecolo ›› 2015, Vol. 39 ›› Issue (4): 343-351.doi: 10.17521/cjpe.2015.0033

• Orginal Article • Previous Articles     Next Articles

Community succession and photosynthetic physiological characteristics of pasture plants in a sub-alpine meadow in Gannan, China

CHEN Shi-Wei, LIU Min-Xia*(), JIA Yun, AN Qi, AN Yan-Fei   

  1. College of Geography and Environmental Science, Northwest Normal University, Lanzhou 730070, China
  • Received:2014-07-07 Accepted:2015-01-28 Online:2015-04-21 Published:2015-04-01
  • Contact: Min-Xia LIU
  • About author:

    # Co-first authors

Abstract: <i>Aims</i>

A plant’s photosynthetic characteristics reflect its adaptive strategies to a given environment. Using pasture plants within enclosures representing communities at different stages of habitat restoration, our objective was to determine how photosynthetic characteristics vary between these different communities and what causes these differences in order to find the theoretical basis to foster rehabilitation of degraded grassland in sub-alpine meadows.


We predicated a succession sequence according to the species richness and the Shannon-Wiener diversity indices, the important values of the main species, and the biotype of five different communities. We measured several photosynthetic parameters including area-based leaf CO2 assimilation rate (Aarea), special leaf area (SLA), foliar nitrogen content based on mass (Nmass), photosynthetic nitrogen-use efficiency (PNUE), water-use efficiency (WUE) and chlorophyll content (SPAD) of dominant species and three common species in each succession stage. Soil water content and total nitrogen of surface soil (0-20 cm) for each community were measured as well. One-way ANOVA was used to find the differences between dominant species, while principal components analysis (PCA) was used to reveal the variation in different communities for each measured parameter.

<i>Important findings</i>

Photosynthetic traits were different among dominant species and different succession communities. The Aarea, WUE and SPAD of the dominant species decreased as succession progressed, but the Nmass, PNUE and SLA showed no consistent patterns related to succession; they varied between different functional groups. For each of the non-dominant species, the Aarea and SPAD gradually decreased as succession proceeded from initial stage to climax stage. With succession, WUE and PNUE of the non-leguminous plants (Elymus dahuricus and Geranium wilfordii) decreased while SLA and Nmass increased. However, there were no obvious changes in these parameters for the leguminous plant (Medicago sativa). Soil water content and total nitrogen increased with succession, suggesting that water content and nitrogen are two important factors affecting variation of community photosynthetic characteristics in different stages of restoration succession.

Key words: functional group, photosynthetic physiological characteristics, restoration, succession, resource-use efficiency

Table 1

Characters of five different community plots"

Important value of dominant and main species
Shannon indices
A 刺儿菜 Cirsium setosum (0.251) 20 3.55
紫苜蓿 Medicago sativa (0.093)
披碱草 Elymus dahuricus (0.054)
B 披碱草 Elymus dahuricus (0.429) 18 2.79
刺儿菜 Cirsium setosum (0.069)
二裂委陵菜 Potentilla bifurca (0.060)
紫苜蓿 Medicago sativa (0.055)
C 紫苜蓿 Medicago sativa (0.148) 32 4.00
甘青蒿 Artemisia tangutica (0.068)
老鹳草 Geranium wilfordii (0.047)
披碱草 Elymus dahuricus (0.060)
少花米口袋 Gueldenstaedtia verna (0.017)
D 沙蒿 Artemisia desertorum (0.140) 36 4.01
羊茅 Festuca ovina (0.109)
圆穗蓼 Polygonum macrophyllum (0.083)
紫苜蓿 Medicago sativa (0.071)
披碱草 Elymus dahuricus (0.056)
E 金露梅 Potentilla fruticosa (0.433) 40 4.15
蕨麻 Potentilla anserina (0.060)
披碱草 Elymus dahuricus (0.047)
紫苜蓿 Medicago sativa (0.011)

Table 2

Area-based leaf CO2 assimilation rate (Aarea), chlorophyll content (SPAD), foliar nitrogen content based on mass (Nmass) and specific leaf area (SLA) of dominant species at different succession stages (mean ± SD)"

(μmol CO2 m-1·s-1)
SPAD Nmass
刺儿菜 Cirsium setosum 15.25 ± 2.71c 47.99 ± 2.36e 18.14 ± 1.97bcd 150.19 ± 5.48a
披碱草 Elymus dahuricus 17.41 ± 0.64c 39.69 ± 0.20cd 11.52 ± 3.18a 180.40 ± 4.88ab
紫苜蓿 Medicago sativa 16.51 ± 2.95c 50.96 ± 1.99e 23.04 ± 1.77de 241.35 ± 35.07c
少花米口袋 Gueldenstaedtia verna 14.20 ± 1.48bc 43.02 ± 1.97d 26.23 ± 3.62e 200.12 ± 10.21bc
甘青蒿 Artemisia tangutica 13.52 ± 0.72bc 30.75 ± 0.49b 19.48 ± 2.66cd 242.24 ± 15.87c
老鹳草 Geranium wilfordii 16.06 ± 2.52c 24.75 ± 1.72a 15.43 ± 0.42abc 165.20 ± 17.33ab
圆穗蓼 Polygonum macrophyllum 10.40 ± 4.21b 35.90 ± 5.62c 13.19 ± 2.77ab 194.10 ± 27.70ab
沙蒿 Artemisia desertorum 13.30 ± 1.55bc 35.48 ± 2.65c 16.61 ± 3.09abc 207.30 ± 31.33bc
金露梅 Potentilla fruticosa 4.29 ± 0.92a 27.10 ± 1.25ab 18.82 ± 1.72cd 203.66 ± 37.09bc

Fig. 1

Resource-use efficiency of different dominant species (mean ± SD). CE, Cirsium setosum; GQ, Artemisia tangutica; JL, Potentilla fruticosa; LG, Geranium wilfordii; MD, Gueldenstaedtia verna; MX, Medicago sativa; PJ, Elymus dahuricus; QH, Polygonum macrophyllum; SH, Artemisia desertorum. PNUE, photosynthetic nitrogen-use efficiency; WUE, water-use efficiency. Different letters in the same column are significantly different (p < 0.05)."

Table 3

The lowerpart of coefficient matrix among photosynthetic characteristics"

Photosynthetic physiological characteristics
基于面积的CO2同化速率Aarea 1
基于质量的叶片氮含量 Nmass 0.151 1
光合氮利用效率 PNUE 0.725*** -0.436** 1
叶绿素含量 SPAD 0.469** 0.311 0.103 1
气孔导度 Gs 0.319* 0.060 0.398* 0.140 1
水分利用效率 WUE 0.712*** -0.136 0.369* 0.293 -0.314* 1
比叶面积 SLA -0.215 0.511** -0.105 0.133 0.092 -0.353* 1

Fig. 2

Principal components (PC) analysis of photosynthetic physiological characteristics of nine different species. CE, Cirsium setosum; GQ, Artemisia tangutica; JL, Potentilla fruticosa; LG, Geranium wilfordii; MD, Gueldenstaedtia verna; MX, Medicago sativa; PJ, Elymus dahuricus; QH, Polygonum macrophyllum; SH, Artemisia desertorum."

Fig. 3

Principal components (PC) analysis of photosynthetic physiological characteristics of common species in five communities. ■, Geranium wilfordii; ○, Medicago sativa; △, Elymus dahuricus; A, Cirsium setosum-Artemisia tangutica community; B, Elymus dahuricus-Cirsium setosum community; C, Medicago sativa-Artemisia tangutica community; D, Artemisia desertorum-Festuca ovina community; E, Potentilla fruticosa community."

Fig. 4

Soil water content and total nitrogen content (TN) of five different community plots (mean ± SD). A, Cirsium setosum-Artemisia tangutica community; B, Elymus dahuricus- Cirsium setosum community; C, Medicago sativa-Artemisia tangutica community; D, Artemisia desertorum-Festuca ovina community; E, Potentilla fruticosa community. Different capital letters indicate significant differences among soil water contents; different small letters indicate significant differences among soil total nitrogen contents (p < 0.05)."

[1] An H, Shangguan ZP (2007). Photosynthetic characteristics of dominant plant species at different succession stages of vegetation on Loess Plateau.Chinese Journal of Applied Ecology, 18, 1175-1180.(in Chinese with English abstract)
[安慧, 上官周平 (2007). 黄土高原植被不同演替阶段优势种的光合生理特性. 应用生态学报, 18, 1175-1180.]
[2] An H, Shangguan ZP (2008). Specific leaf area, leaf nitrogen content, and photosynthetic acclimation of Trifolium repens L. seedlings grown at different irradiances and nitrogen concentrations.Photosynthetica, 46, 143-147.
[3] Anten NPR, Miyazawa K, Hikosaka K, Nagashima H, Hirose T (1998a). Leaf nitrogen distribution in relation to leaf age and photon flux density in dominant and subordinate plants in dense stands of a dicotyledonous herb.Oecologia, 113, 314-324.
[4] Anten NPR, Werger MJA, Medina E (1998b). Nitrogen distribution and leaf area indices in relation to photosynthetic nitrogen use efficiency in savanna grasses.Plant Ecology, 138, 63-75.
[5] Bassow SL, Bazzaz FA (1997). Intra-and inter-specific variation in canopy photosynthesis in a mixed deciduous forest.Oecologia, 109, 507-515.
[6] Bazzaz FA (1979). The physiological ecology of plant succession.Annual Review of Ecology and Systematics, 10, 351-371.
[7] Castellanos AE, Martinez MJ, Llano JM, Halvorson WL, Espiricueta M, Espejel I (2005). Successional trends in Sonoran Desert abandoned agricultural fields in northern Mexico.Journal of Arid Environments, 60, 437-455.
[8] Cui XY, Du ZC, Wang YF (2000). Photosynthetic characteristics of a semi-arid sandy grassland community in inner mongolia.Acta Phytoecologica Sinica, 24, 541-546.(in Chinese with English abstract)
[崔骁勇, 杜占池, 王艳芬 (2000). 内蒙古半干旱草原区沙地植物群落光合特征的动态研究. 植物生态学报, 24, 541-546.]
[9] Ding SY, Song YC (1999). The comparation of photosynthesis physi-ecology of evergreen broad-leaved forest of Tiantong national forest park in Zhejiang Province, China.Acta Ecologica Sinica, 19, 318-323.(in Chinese with English abstract)
[丁圣彦, 宋永昌 (1999). 浙江天童常绿阔叶林演替系列优势种光合生理生态的比较. 生态学报, 19, 318-323.]
[10] Ellsworth DS, Reich PB (1996). Photosynthesis and leaf nitrogen in five Amazonian tree species during early secondary succession.Ecology, 77, 581-594.
[11] Evans JR (1989). Photosynthesis and nitrogen relationships in leaves of C3 plants.Oecologia, 78, 9-19.
[12] Funk JL, Vitousek PM (2007). Resource-use efficiency and plant invasion in low-resource systems.Nature, 446, 1079-1081.
[13] Givnish TJ (1988). Adaptation to sun and shade: A whole-plant perspective.Australian Journal of Plant Physiology, 15, 63-92.
[14] Hikosaka K, Hirose T (2000). Photosynthetic nitrogen-use efficiency in evergreen broad-leaved woody species coexisting in a warm-temperate forest.Tree Physiology, 20, 1249-1254.
[15] Huang ZQ, Xu ZH, Blumfield TJ, Bubb K (2008). Effects of mulching on growth, foliar photosynthetic nitrogen and water use efficiency of hardwood plantations in subtropical Australia.Forest Ecology and Management, 255, 3447-3454.
[16] Jiang GM, He WM (1999). Species-and habitat-variability of photosynthesis, transpiration and water use efficiency of different plant species in Maowusu Sand Area.Acta Botanica Sinica, 41, 114-1124.
[17] Lambers H, Chapin III FS, Pons TL (2008). Plant Physiological Ecology. 2nd edn. Springer, New York.
[18] Li QK, Ma KP (2002). Advances in plant succession ecophysiology.Acta Phytoecologica Sinica, 26(S1), 9-19.(in Chinese with English abstract)
[李庆康, 马克平 (2002). 植物群落演替过程中植物生理生态学特性及其主要环境因子的变化. 植物生态学报, 26(S1), 9-19.]
[19] Li YL, Cui JY, Su YZ (2005). Specific leaf area and leaf dry matter content of some plants in different dune habitats.Acta Ecologica Sinica, 25, 304-311.(in Chinese with English abstract)
[李玉霖, 崔建垣, 苏永中 (2005). 不同沙丘生境主要植物比叶面积和叶干物质含量的比较. 生态学报, 25, 304-311.]
[20] Liu FD, Yang WJ, Wang ZS, Xu Z, Liu H, Zhang M, Liu YH, An SQ, Sun SC (2010). Plant size effects on the relationships among specific leaf area, leaf nutrient content, and photosynthetic capacity in tropical woody species.Acta Oecologica, 36, 149-159.
[21] Mahajan S, Narendra T (2005). Cold, salinity and drought stresses: An overview.Archives of Biochemistry and Biophysis, 444, 139-158.
[22] Manetas Y, Grammatikopoulos G, Kyparissis A (1998). The use of the portable, non-destructive, SPAD-502 (Minolta) chlorophyll meter with leaves of varying trichome density and anthocyanin content.Journal of Plant Physiology, 153, 513-516.
[23] Niu SL, Jiang GM, Gao LM, Li YG, Liu MZ (2003). Comparison of gas exchange traits of different plant species in Hunshandak sand area.Acta Phytoecologica Sinica, 27, 318-324.
[24] Novriyanti E, Watanabe M, Makoto K, Takeda T, Hashidoko Y, Koike T (2012). Photosynthetic nitrogen and water use efficiency of acacia and eucalypt seedlings as afforestation species.Photosynthetica, 50, 273-281.
[25] Ran F, Zhang XL, Zhang YB, Korpelainen H, Li CY (2013). Altitudinal variation in growth, photosynthetic capacity and water use efficiency of Abies faxoniana Rehd. et Wils. seedlings as revealed by reciprocal transplantations.Trees, 27, 1405-1416.
[26] Shangguan ZP, Shao MG, Dyckmans J (2000). Effects of nitrogen nutrition and water deficit on net photosynthetic rate and chlorophyll fluorescence in winter wheat.Journal of Plant Physiology, 156, 46-51.
[27] Sun HQ, Zhou H, Wang P (1999). Progress on grassland degenerated succession.Grassland of China, 21(1), 51-56.(in Chinese with English abstract)
[孙海群, 周禾, 王培 (1999). 草地退化演替研究进展. 中国草地, 21(1), 51-56.]
[28] Wang GH (2002). Plant traits and soil chemical variables during a secondary vegetation succession in abandoned fields on the loess plateau.Acta Botanica Sinica, 44, 990-998.
[29] Wright LJ, Reich PB, Cornelissen JHC, Falster DS, Garnier E, Hikosaka K, Lamont BB, Lee W, Oleksyn J, Osada N, Poorter H, Villar R, Warton DI, Westoby M (2005). Assessing the generality of global leaf trait relationships.New Phytologist, 166, 485-496.
[30] Wu GL, Du GZ (2007). Discussion on ecological construction and sustainable development of degraded alpine grassland ecosystem of the Qinghai-Tibetan Plateau.Chinese Journal of Nature, 29, 159-164.(in Chinese)
[武高林, 杜国祯 (2007). 青藏高原退化高寒草地生态系统恢复和可持续发展探讨. 自然杂志, 29, 159-164.]
[31] Yan CR, Han XG, Chen LZ (2001). Water use efficiency of six woody species in relation to micro-environmental factors of different habitats.Acta Ecologica Sinica, 21, 1952-1956.(in Chinese with English abstract)
[严昌荣, 韩兴国, 陈灵芝 (2001). 六种木本植物水分利用效率和其小生境关系研究. 生态学报, 21, 1952-1956.]
[32] Yu GR, Wang QF (2010). Ecophysiology of Plant Photosynthesis, Transpiration, and Water Use. Science Press, Beijing. 152.(in Chinese)
[于贵瑞, 王秋凤 (2010). 植物光合、蒸腾与水分利用的生理生态学. 科学出版社, 北京. 152.]
[33] Zhang DY, Wang G, Zhao SL (1988). A quantitative study of the vegetation succession on the abandoned arable lands of the subalpine meadows in Gannan Prefecture of Gansu Province I. Analysis of community comosition.Acta Phytoecological et Geobotanica Sinica, 12, 283-291.(in Chinese with English abstract)
[张大勇, 王刚, 赵松岭 (1988). 亚高山草甸弃耕地植物群落演替的数量研究I. 群落组成分析. 植物生态学与地植物学学报, 12, 283-291.]
[34] Zhang WY, Fan JW, Zhong HP, Hu ZM, Song LL, Wang N (2010). The nitrogen: phosphorus stoichiometry of different plant functional groups for dominant species of typical steppes in China.Acta Agrestia Sinica, 18, 503-509.(in Chinese with English abstract)
[张文彦, 樊江文, 钟华平, 胡中民, 宋璐璐, 王宁 (2010). 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, 18, 503-509.]
[35] Zhou HK, Zhao XQ, Zhao L, Li YN, Wang SP, Xu SX, Zhou L (2008). Restoration capability of alpine meadow ecosystem on Qinghai-Tibetan Plateau.Chinese Journal of Ecology, 27, 697-704.(in Chinese with English abstract)
[周华坤, 赵新全, 赵亮, 李英年, 汪诗平, 徐世晓, 周立 (2008). 青藏高原高寒草甸生态系统的恢复能力. 生态学杂志, 27, 697-704.]
[1] DING Wei,WANG Yu-Bing,XIANG Guan-Hai,CHI Yong-Gang,LU Shun-Bao,ZHENG Shu-Xia. Effects of Caragana microphylla encroachment on community structure and ecosystem function of a typical steppe [J]. Chin J Plant Ecol, 2020, 44(1): 33-43.
[2] WANG Yu-Bing,SUN Yi-Han,DING Wei,ZHANG En-Tao,LI Wen-Huai,CHI Yong-Gang,ZHENG Shu-Xia. Effects and pathways of long-term nitrogen addition on plant diversity and primary productivity in a typical steppe [J]. Chin J Plant Ecol, 2020, 44(1): 22-32.
[3] CHEN Chan,ZHANG Shi-Ji,LI Lei-Da,LIU Zhao-Dan,CHEN Jin-Lei,GU Xiang,WANG Liu-Fang,FANG Xi. Carbon, nitrogen and phosphorus stoichiometry in leaf, litter and soil at different vegetation restoration stages in the mid-subtropical region of China [J]. Chin J Plant Ecol, 2019, 43(8): 658-671.
[4] WANG Ming-Ming,LIU Xin-Ping,HE Yu-Hui,ZHANG Tong-Hui,WEI Jing,Chelmge ,SUN Shan-Shan. How enclosure influences restored plant community changes of different initial types in Horqin Sandy Land [J]. Chin J Plant Ecol, 2019, 43(8): 672-684.
[5] FU Yi-Wen, TIAN Da-Shuan, WANG Jin-Song, NIU Shu-Li, ZHAO Ken-Tian. Patterns and affecting factors of nitrogen use efficiency of plant leaves and roots in Nei Mongol and Qinghai-Xizang Plateau grasslands [J]. Chin J Plant Ecol, 2019, 43(7): 566-575.
[6] MIAO Bai-Ling, LIANG Cun-Zhu, SHI Ya-Bo, LIANG Mao-Wei, LIU Zhong-Ling. Temporal changes in precipitation altered aboveground biomass in a typical steppe in Nei Mongol, China [J]. Chin J Plant Ecol, 2019, 43(7): 557-565.
[7] Mo Zhangqin. Re-legalizing China’s ecological conservation redline: The position, dilemma and path [J]. Biodiv Sci, 2019, 27(3): 347-352.
[8] Rijin Jiang,Linlin Zhang,Kaida Xu,Pengfei Li,Yi Xiao,Ziwei Fan. Characteristics and diversity of nekton functional groups in the coastal waters of south-central Zhejiang Province [J]. Biodiv Sci, 2019, 27(12): 1330-1338.
[9] SUN Hui-Min, JIANG Jiang, CUI Li-Na, ZHANG Shui-Feng, ZHANG Jin-Chi. Effects of Spartina alterniflora invasion on soil organic carbon composition of mangrove wetland in Zhangjiang River Estuary [J]. Chin J Plan Ecolo, 2018, 42(7): 774-784.
[10] LIU Yuan-Yuan, MA Jin-Ze, BU Zhao-Jun, WANG Sheng-Zhong, ZHANG Xue-Bing, ZHANG Ting-Yu, LIU Sha-Sha, FU Biao, KANG Yuan. Effect of geographical sources and biochemical traits on plant litter decomposition in a peatland [J]. Chin J Plan Ecolo, 2018, 42(7): 713-722.
[11] Xiaorong Huang. Relationship between plant functional diversity and productivity of Pinus massoniana plantations in Guangxi [J]. Biodiv Sci, 2018, 26(7): 690-700.
[12] YU Xiao-Ya, LI Yu-Hui, YANG Guang-Rong. Fruit types and seed dispersal modes of plants in different communities in Shilin Geopark, Yunnan, China [J]. Chin J Plan Ecolo, 2018, 42(6): 663-671.
[13] SONG Xiao-Yan,WANG Gen-Xu,RAN Fei,YANG Yan,ZHANG Li,XIAO Yao. Flowering phenology and growth of typical shrub grass plants in response to simulated warmer and drier climate in early succession Taiga forests in the Da Hinggan Ling of northeast China [J]. Chin J Plan Ecolo, 2018, 42(5): 539-549.
[14] GU Xiang,ZHANG Shi-Ji,LIU Zhao-Dan,LI Lei-Da,CHEN Jin-Lei,WANG Liu-Fang,FANG Xi. Effects of vegetation restoration on soil organic carbon concentration and density in the mid-subtropical region of China [J]. Chin J Plan Ecolo, 2018, 42(5): 595-608.
[15] Qian YANG, Wei WANG, Hui ZENG. Effects of nitrogen addition on the plant diversity and biomass of degraded grasslands of Nei Mongol, China [J]. Chin J Plan Ecolo, 2018, 42(4): 430-441.
Full text



[1] Zhang Zhen-jue. Some Principles Governing Shedding of Flowers and Fruits in Vanilla fragrans[J]. Chin Bull Bot, 1985, 3(05): 36 -37 .
[2] Qian Gao;Yuying Liu;Yinan Fei;Dapeng Li;Xianglin Liu* . Research Advances into the Root Radial Patterning Gene SHORT-ROOT[J]. Chin Bull Bot, 2008, 25(03): 363 -372 .
[3] Wang Bao-shan;Zou Qi and Zhao Ke-fu. Advances in Mechanism of Crop Salt Tolerance and Strategies for Raising Crop Salt Tolerance[J]. Chin Bull Bot, 1997, 14(增刊): 25 -30 .
[4] HE Feng WU Zhen-Bin. Application of Aquatic Plants in Sewage Treatment and Water Quality Improvement[J]. Chin Bull Bot, 2003, 20(06): 641 -647 .
[5] ZHANG Yan FANG Li LI Tian-Fei YAO Zhao-BingJIANG Jin-Hui. Effect of Calcium on the Heat Tolerance and Active Oxygen Metabolism of Tobacco Leaves[J]. Chin Bull Bot, 2002, 19(06): 721 -726 .
[6] JIA Hu-Sen LI De-QuanHAN Ya-Qin. Cytochrome b-559 in Chloroplasts[J]. Chin Bull Bot, 2001, 18(02): 158 -162 .
[7] Wei Sun;Chonghui Li;Liangsheng Wang;Silan Dai*. Analysis of Anthocyanins and Flavones in Different-colored Flowers of Chrysanthemum[J]. Chin Bull Bot, 2010, 45(03): 327 -336 .
[8] . Phosphate_Stress Protein and Iron_Stress Protein in Plants[J]. Chin Bull Bot, 2001, 18(05): 571 -576 .
[9] ZHANG Da-Yong, JIANG Xin-Hua. An Ecological Perspective on Crop Prduction[J]. Chin J Plan Ecolo, 2000, 24(3): 383 -384 .
[10] Gui Ji-xun, Zhu Ting-cheng. Study of Energy Flow Between Litter and Decomposers in Aneurolepidium chinese Grassland[J]. Chin J Plan Ecolo, 1992, 16(2): 143 -148 .