高寒草甸不同演替阶段优势种叶片光合性状研究
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摘要
围封年限不同导致形成以不同优势种植物建立的群落样带。按照植被的演替序列(围封1、3、5、15、30 a),在甘南高寒草甸选择不同演替阶段的优势植物:刺儿菜(Cirsium setosum)、披碱草(Elymus nutans)、紫花苜蓿(Medicago sativa)、沙蒿(Artemisia desertorum)和金露梅(Potentilla fruticosa),对其叶片光合性状进行了测定,探究植物叶片光合性状与环境因子之间的关系,旨在确定演替过程中控制植物光合性状变化的主要环境因子,为进一步促进高寒草甸草地恢复年限的设定及草地资源的可持续利用提供科学依据。结果表明,随着植被演替的进行,群落形成了以草本→半灌木→灌木为主的演替序列。表层(0～20 cm)土壤含水量(SWC)随演替进行呈增大趋势,变化范围为0.323～0.390 g·g~(-1);土壤温度(t_s)和光照强度(I_l)在前期最大(23.974℃;90.382 lx),后期最小(20.310℃;65.220 lx)。各演替阶段优势种光合特征值在时间尺度上除相对叶绿素外均存在明显差异,草本群落优势种(刺儿菜、披碱草、紫花苜蓿)净光合速率(Pn)高于灌丛群落优势种(金露梅),瞬时水分利用效率(WUEi)与Pn变化规律相似。蒸腾速率(T_r)与稳定碳同位素(δ13C)呈缓慢降低趋势,分别降低2.128 mmol·m~(-2)·s~(-1)、4.359‰。各演替群落优势种叶片气孔导度(Gs)随演替的进行逐步升高,最大值为0.155mol·m~(-2)·s~(-1);叶片含水量呈先升高后降低趋势,演替15年最大(5.207g·g~(-1));而相对叶绿素(SPAD)呈先降低后稳定趋势。在植被演替进程中,LWC、G_s与SWC呈正相关,δ~(13)C、T_r、P_n、WUE_i、SPAD与SWC呈负相关,t_s、I_l与SPAD、T_r、P_n、WUE_i、δ~(13)C呈正相关,与LWC、G_s呈负相关。环境因子对植物叶光合性状的影响大小为I_l>SWC>t_s,这表明光照度是该区演替发生及影响光合性状的主要因子。综合分析表明,群落演替是物种扩散和环境综合作用的过程,该围封地由物质获得能力强的群落向物质保持能力强的群落过渡,围封5～15 a群落环境最优。
Different enclosure years leads to the formation of community transect_s with different dominant species. According to the succession sequence of vegetation(1, 3, 5, 15, 30 a), the dominant plant_s at different successional stages were selected respectively in alpine meadow of Gannan: Cirsium setosum, Elymus nutans, Medicago sativa, Artemisia desertorum, Potentilla fruticosa in different successional stages. The photosynthetic characteristics of dominant plant leaves were measured, according to the succession sequence in the alpine meadow of Gannan. The relationship between plant photosynthetic characteristics and environmental factors was explored to determine the main environmental factors during succession. The result_s showed that successional sequences(herbaceous→semi-shrubs→shrubs) were formed with the community change, soil water content(SWC) at 0-20 cm increased, the range of variation was 0.323-0.390 g·g~(-1), and soil temperature(t_s) and light intensity(I_l) were the highest at the early stage(23.974 ℃; 90.382 l_x), and the lowest at the later stage(20.310 ℃; 65.220 l_x). The photosynthetic trait_s of the dominant species were significantly different on the time scale. The net photosynthetic rate(Pn) of the dominant species(Cirsium setosum, Elmus dahuricus, Medicago sativa) was higher than scrubs community(Potentilla fruticosa), and instantaneous water use efficiency(WUEi) were similar to the Pn. The transpiration rate(Tr) and the stable carbon isotope(δ~(13) C) showed a decreasing trend, the reduction were 2.128 mmol·m~(-2)·s~(-1) and 4.359‰ respectively. The leaf stomatal conductance(Gs) was lower in the early stage, and gradually increased in the late stage, with the maximum of 0.155 mol·m~(-2)·s~(-1). The water content of leaves increased first and then decreased, with the maximum at 15 years(5.207 g·g~(-1)). The relative chlorophyll(SPAD) decreased first, followed by a steady trend. LWC, Gs and SWC were positively correlated in the course of succession, and δ~(13) C, Tr, Pn, WUEi, SPAD were negatively correlated with SWC. t_s, I_l was positively correlated with SPAD, Tr, Pn, WUEi and δ~(13) C, but negatively correlated with LWC and Gs. The influence of environmental factors on leaf photosynthetic characteristics were as follows: I_l>SWC>t_s, these characteristics indicated that light intensity was the main factor affecting photosynthetic trait_s in succession stage. The comprehensive analysis showed that the community succession was a process from species dispersal and environment effect. The meadow enclosure was transformed from a community with the strong ability of material acquisition to the strong ability of material conservation. The optimal enclosure environment would be in 5-15 years.
Different enclosure years leads to the formation of community transect_s with different dominant species. According to the succession sequence of vegetation(1, 3, 5, 15, 30 a), the dominant plant_s at different successional stages were selected respectively in alpine meadow of Gannan: Cirsium setosum, Elymus nutans, Medicago sativa, Artemisia desertorum, Potentilla fruticosa in different successional stages. The photosynthetic characteristics of dominant plant leaves were measured, according to the succession sequence in the alpine meadow of Gannan. The relationship between plant photosynthetic characteristics and environmental factors was explored to determine the main environmental factors during succession. The result_s showed that successional sequences(herbaceous→semi-shrubs→shrubs) were formed with the community change, soil water content(SWC) at 0-20 cm increased, the range of variation was 0.323-0.390 g·g~(-1), and soil temperature(t_s) and light intensity(I_l) were the highest at the early stage(23.974 ℃; 90.382 l_x), and the lowest at the later stage(20.310 ℃; 65.220 l_x). The photosynthetic trait_s of the dominant species were significantly different on the time scale. The net photosynthetic rate(Pn) of the dominant species(Cirsium setosum, Elmus dahuricus, Medicago sativa) was higher than scrubs community(Potentilla fruticosa), and instantaneous water use efficiency(WUEi) were similar to the Pn. The transpiration rate(Tr) and the stable carbon isotope(δ~(13) C) showed a decreasing trend, the reduction were 2.128 mmol·m~(-2)·s~(-1) and 4.359‰ respectively. The leaf stomatal conductance(Gs) was lower in the early stage, and gradually increased in the late stage, with the maximum of 0.155 mol·m~(-2)·s~(-1). The water content of leaves increased first and then decreased, with the maximum at 15 years(5.207 g·g~(-1)). The relative chlorophyll(SPAD) decreased first, followed by a steady trend. LWC, Gs and SWC were positively correlated in the course of succession, and δ~(13) C, Tr, Pn, WUEi, SPAD were negatively correlated with SWC. t_s, I_l was positively correlated with SPAD, Tr, Pn, WUEi and δ~(13) C, but negatively correlated with LWC and Gs. The influence of environmental factors on leaf photosynthetic characteristics were as follows: I_l>SWC>t_s, these characteristics indicated that light intensity was the main factor affecting photosynthetic trait_s in succession stage. The comprehensive analysis showed that the community succession was a process from species dispersal and environment effect. The meadow enclosure was transformed from a community with the strong ability of material acquisition to the strong ability of material conservation. The optimal enclosure environment would be in 5-15 years.
引文
BANIYA C B,SOLH?Y T,VETAAS O R,2009.Torstein Solhy,Ole R.Vetaas.2009.Temporal changes in species diversity and composition in abandoned fields in a trans-himalayan landscape Nepal[J].Plant Ecology,201(2):383-399.
BAZZAZ F A,1979.The physiological ecology of plant succession[J].Annual Review of Ecology and Systematics,10:351-371.
BAZZAZ F A,1996.Plants in changing environments:linking physiological,population,and community ecology[M].New York:Cambridge University Press:264-280.
ClEMENT F E,1916.Plant succession:an analysis of the development of vegetation[J].Carnegie Institution of Washington Publ.242(1162):339-341.
CRAIG H,1957.Isotope standards for carbon and oxygen and correlation factors for mass spectrometric analysis of carbon dioxide[J].Geochimica Cosmochimica Acta,12(1-2):133-149.
FARQUHAR G D,O′LEARY M H,1982.On the relationship between carbon isotope discrimination and the intercellular carbon dioxide concentration of leaves[J].Australian Jo urnal of Plant Physiology,9(2):121-137.
HORN H,1974.The ecology of secondary succession[J].Annual Review of Ecology and Systematics,5:25-37.
KAZAKOU E,VILE D,SHIPLEY B,et al.,2006.Co-variations in litter decomposition,leaf traits and plant growth in species from a Mediterranean old-field succession[J].Functional Ecology,20(1):21-30.
LAWLOR D W,1995.Photosynthesis,productivity and environment[J].Journal of Experimental Botany,46:1449-1461.
LLAMBI L D,FONTAINE M,RADA F,et al.,2003.Ecophysiology of dominant plant spMecies during old-field succession in a high tropical Andean ecosystem[J].Arctic,Antarctic,and Alpine Research,35(4):447-453.
MA K P,HUANG J H,YU S L,et al.,1995.Plant community diversity in Dongling Mountain[J].Acta Ecologica Sinica,15(3):268-277.
MARTIN M A,GUILLERMO B F,JUAN A M S,et al.,2009.Gustavo AS.Radiation interception and use efficiency as affected by breeding in Mediterranean wheat[J].Field Crops Res,110(2):91-97.
VIOLLE C,NAVAS M L,VILE D,2007.Let the concept of trait be functional[J].Oikos,116(5):882-892.
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[J].应用生态学报,18(6):1175-1180.
白炜,王根绪,刘光生,2011.青藏高原高寒草甸生长期CO2排放对气温升高的响应[J].生态学杂志,30(6):1045-1051.
曹广民,吴琴,李东,等,2004.土壤-牧草氮素供需状况变化对高寒草甸植被演替与草地退化的影响[J].生态学杂志,23(6):25-28.
陈世伟,2015.甘南亚高寒草甸恢复演替动态及其光合特性研究[D].兰州:西北师范大学.
杜国祯,王刚,1995.甘南亚高山草甸人工草地的演替和质量变化[J].植物学报,37(4):306-313.
冯晓钰,周广胜,2018.夏玉米叶片水分变化与光合作用和土壤水分的关系[J].生态学报,38(1):1-9.
管铭,金则新,王强,等,2014.千岛湖次生林优势种植物光合特性对不同光环境的响应[J].应用生态学报,25(6):1615-1622.
李宏林,徐当会,杜国祯,2012.青藏高原高寒沼泽湿地在退化梯度上植物群落组成的改变对湿地水分状况的影响[J].植物生态学报,36(5):403-410.
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[J].植物生态学报,26(S1):9-19.
李善家,苏培玺,张海娜,等,2013.荒漠植物叶片水分和功能性状特征及其相互关系[J].植物生理学报,49(2):153-160.
李征珍,杨琼,石莎,等,2017.蒙古沙冬青光合作用特征及其影响因素[J].生态学杂志,36(9):2481-2488.
刘旻霞,刘洋洋,陈世伟,等,2016.高寒草甸不同坡向条件下植物叶片δ13C及水分利用效率的变化[J].应用生态学报,27(12):3816-3822.
刘旻霞,2017.亚高寒草甸不同坡向植物光合生理和叶片形态差异[J].生态学报,37(24):1-11.
马剑英,陈发虎,夏敦胜,等,2008.荒漠植物红砂(Reaumuria soongorica)叶片元素和水分含量与土壤因子的关系[J].生态学报,28(3):983-992.
任青吉,李宏林,卜海燕,2015.玛曲高寒沼泽化草甸51种植物光合生理和叶片形态特征的比较[J].植物生态学报,39(6):593-603.
王世雄,王孝安,李国庆,等,2010.陕西子午岭植物群落演替过程中物种多样性变化与环境解释[J].生态学报,30(6):1638-1647.
王亚婷,王玺,赵天启,等,2017.不同放牧强度上内蒙古短花针茅草原植物功能群水分和氮素利用效率相关分析[J].生态环境学报,26(6):964-970.
王长庭,龙瑞军,王启兰,等,2008.放牧扰动下高寒草甸植物多样性、生产力对土壤养分条件变化的响应[J].生态学报,28(9):4144-4152.
韦兰英,上官周平,2006.黄土高原不同演替阶段草地植被细根垂直分布特征与土壤环境的关系[J].生态学报,26(11):3740-3748.
吴统贵,周和锋,吴明,等,2008.旱柳光合作用动态及其与环境因子的关系[J].生态学杂志,27(12):2056-2061.
吴统贵,2009.杭州湾滨海湿地植被群落演替及优势物种生理生态学特征[D].北京:中国林业科学研究院.
夏艳菊,张静,邹顺,等,2018.南亚热带森林群落演替过程中结构多样性与碳储量的变化[J].生态环境学报,27(3):424-431.
岳明,周虹霞,1997.太白山北坡落叶阔叶林物种多样性特征[J].云南植物研究,19(2):171-176.
张金屯,2004.数量生态学[M].北京:科学出版社.
张梦弢,张青,亢新刚,等,2015.长白山云冷杉林不同演替阶段群落稳定性[J].应用生态学报,26(6):1609-1616.
BAZZAZ F A,1979.The physiological ecology of plant succession[J].Annual Review of Ecology and Systematics,10:351-371.
BAZZAZ F A,1996.Plants in changing environments:linking physiological,population,and community ecology[M].New York:Cambridge University Press:264-280.
ClEMENT F E,1916.Plant succession:an analysis of the development of vegetation[J].Carnegie Institution of Washington Publ.242(1162):339-341.
CRAIG H,1957.Isotope standards for carbon and oxygen and correlation factors for mass spectrometric analysis of carbon dioxide[J].Geochimica Cosmochimica Acta,12(1-2):133-149.
FARQUHAR G D,O′LEARY M H,1982.On the relationship between carbon isotope discrimination and the intercellular carbon dioxide concentration of leaves[J].Australian Jo urnal of Plant Physiology,9(2):121-137.
HORN H,1974.The ecology of secondary succession[J].Annual Review of Ecology and Systematics,5:25-37.
KAZAKOU E,VILE D,SHIPLEY B,et al.,2006.Co-variations in litter decomposition,leaf traits and plant growth in species from a Mediterranean old-field succession[J].Functional Ecology,20(1):21-30.
LAWLOR D W,1995.Photosynthesis,productivity and environment[J].Journal of Experimental Botany,46:1449-1461.
LLAMBI L D,FONTAINE M,RADA F,et al.,2003.Ecophysiology of dominant plant spMecies during old-field succession in a high tropical Andean ecosystem[J].Arctic,Antarctic,and Alpine Research,35(4):447-453.
MA K P,HUANG J H,YU S L,et al.,1995.Plant community diversity in Dongling Mountain[J].Acta Ecologica Sinica,15(3):268-277.
MARTIN M A,GUILLERMO B F,JUAN A M S,et al.,2009.Gustavo AS.Radiation interception and use efficiency as affected by breeding in Mediterranean wheat[J].Field Crops Res,110(2):91-97.
VIOLLE C,NAVAS M L,VILE D,2007.Let the concept of trait be functional[J].Oikos,116(5):882-892.
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[J].应用生态学报,18(6):1175-1180.
白炜,王根绪,刘光生,2011.青藏高原高寒草甸生长期CO2排放对气温升高的响应[J].生态学杂志,30(6):1045-1051.
曹广民,吴琴,李东,等,2004.土壤-牧草氮素供需状况变化对高寒草甸植被演替与草地退化的影响[J].生态学杂志,23(6):25-28.
陈世伟,2015.甘南亚高寒草甸恢复演替动态及其光合特性研究[D].兰州:西北师范大学.
杜国祯,王刚,1995.甘南亚高山草甸人工草地的演替和质量变化[J].植物学报,37(4):306-313.
冯晓钰,周广胜,2018.夏玉米叶片水分变化与光合作用和土壤水分的关系[J].生态学报,38(1):1-9.
管铭,金则新,王强,等,2014.千岛湖次生林优势种植物光合特性对不同光环境的响应[J].应用生态学报,25(6):1615-1622.
李宏林,徐当会,杜国祯,2012.青藏高原高寒沼泽湿地在退化梯度上植物群落组成的改变对湿地水分状况的影响[J].植物生态学报,36(5):403-410.
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[J].植物生态学报,26(S1):9-19.
李善家,苏培玺,张海娜,等,2013.荒漠植物叶片水分和功能性状特征及其相互关系[J].植物生理学报,49(2):153-160.
李征珍,杨琼,石莎,等,2017.蒙古沙冬青光合作用特征及其影响因素[J].生态学杂志,36(9):2481-2488.
刘旻霞,刘洋洋,陈世伟,等,2016.高寒草甸不同坡向条件下植物叶片δ13C及水分利用效率的变化[J].应用生态学报,27(12):3816-3822.
刘旻霞,2017.亚高寒草甸不同坡向植物光合生理和叶片形态差异[J].生态学报,37(24):1-11.
马剑英,陈发虎,夏敦胜,等,2008.荒漠植物红砂(Reaumuria soongorica)叶片元素和水分含量与土壤因子的关系[J].生态学报,28(3):983-992.
任青吉,李宏林,卜海燕,2015.玛曲高寒沼泽化草甸51种植物光合生理和叶片形态特征的比较[J].植物生态学报,39(6):593-603.
王世雄,王孝安,李国庆,等,2010.陕西子午岭植物群落演替过程中物种多样性变化与环境解释[J].生态学报,30(6):1638-1647.
王亚婷,王玺,赵天启,等,2017.不同放牧强度上内蒙古短花针茅草原植物功能群水分和氮素利用效率相关分析[J].生态环境学报,26(6):964-970.
王长庭,龙瑞军,王启兰,等,2008.放牧扰动下高寒草甸植物多样性、生产力对土壤养分条件变化的响应[J].生态学报,28(9):4144-4152.
韦兰英,上官周平,2006.黄土高原不同演替阶段草地植被细根垂直分布特征与土壤环境的关系[J].生态学报,26(11):3740-3748.
吴统贵,周和锋,吴明,等,2008.旱柳光合作用动态及其与环境因子的关系[J].生态学杂志,27(12):2056-2061.
吴统贵,2009.杭州湾滨海湿地植被群落演替及优势物种生理生态学特征[D].北京:中国林业科学研究院.
夏艳菊,张静,邹顺,等,2018.南亚热带森林群落演替过程中结构多样性与碳储量的变化[J].生态环境学报,27(3):424-431.
岳明,周虹霞,1997.太白山北坡落叶阔叶林物种多样性特征[J].云南植物研究,19(2):171-176.
张金屯,2004.数量生态学[M].北京:科学出版社.
张梦弢,张青,亢新刚,等,2015.长白山云冷杉林不同演替阶段群落稳定性[J].应用生态学报,26(6):1609-1616.