松嫩草地植物功能生态学研究
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摘要
随着功能生态学的发展，植物功能生态学研究越来越受到重视，尤其是关于功能多样性对生态系统功能的作用。该论文以此为切入点，以松嫩草原为对象，开展了草地植物功能生态研究，首先论述了松嫩草地属性，然后从植物功能性状角度研究该区域植被状况及功能性状特征，最后通过控制实验研究刈割、施肥、植被低矮稀疏化等干扰条件下羊草性状和羊草群落特征的变化，验证在自然和干扰条件下物种丰富度、功能群丰富度、物种多样性指数、功能性状多样性与群落地上生物量的关系。主要结果如下：
松嫩平原未退化羊草草地的土壤pH、电导率、离子含量(8种)、碱化度分别为9.1-10.2，198.7-643.5μs/cm，734.9-9297.5mg/kg，4.8-28.4%，退化裸地的土壤pH、电导率、离子含量、碱化度分别为10.2-10.4，8240.3-18232.1mg/kg，459.3-1832.6μs/cm，43.0-54.0%。主要土壤类型各发生层的粘粒含量最高，盐土和草甸土淀积层粘粒含量(<0.002mm)超过50%，沼泽土和草甸土淀积层物理粘粒(<0.01mm)含量超过75%。未退化草地地上生物量干重超过250g/m~2，退化草地地上生物量干重超过300g/m~2。单优势种群落广布，羊草的地上生物量干重与群落地上生物量干重的比例最高达97.2%，物种丰富度不超过8种/m~2。退化草地由于高盐碱条件限制，单优势现象更为明显，有些群落仅有1个物种。盐分含量有微域差异地段形成不同的单种优势种群落，斑块景观明显。地表被干扰，原初盐渍化草地土壤的淋溶层消失(践踏风蚀等)，下层高含盐量的盐碱土裸露成新地表，新地表裸露或生长盐生植物群落。地表被干扰是松嫩草地盐碱化的主导原因。松嫩平原草地，不同于世界各地的地带性草原土壤，是土壤决定的草甸植被类型。
松嫩草地66种草本植物7个叶片性状特征表明：叶片厚度变异系数最大，比叶面积、叶片氮含量、叶片磷含量、叶绿素含量和类胡萝卜素含量之间存在显著的正相关关系；叶片干物质含量与叶片磷含量没有显著的相关关系，与其它叶片性状呈显著的负相关关系；叶片厚度与叶片干物质含量和比叶面积呈显著的负相关关系，与其它叶片性状不相关。方差分析表明，叶片干物质含量、比叶面积、叶片厚度、叶片氮、叶片磷含量在4个功能群(多年生根茎禾草、多年生丛生禾草、多年生杂类草、1年生或2年生植物)间差异显著，叶绿素含量和类胡萝卜素含量在各个功能群间差异不显著。多重比较结果显示多年生根茎禾草和多年生丛生禾草叶片的7个性状差异不显著；多年生根茎禾草和多年生丛生禾草的叶片干物质含量高于多年生杂类草和1年生或2年生草本，其它性状小于这两个功能群。松嫩草地草本植物叶片氮磷含量分别为22.7和1.8mg/g，氮磷比为13.0，氮相对缺乏。
松嫩草地生产力高，群落饱和度低，优势种在群落中占的比例大。多年生根茎禾草和多年生杂类草功能群物种丰富度最高，各功能群间功能群丰富度没有显著差异，1年生或2年生植物群落地上生物量最高。叶片干物质含量、比叶面积、叶片氮含量、叶片磷含量、株高、比根长等性状在各功能群间差异显著。1年生或2年生植物功能群叶片干物质含量高、比叶面积小、氮磷含量高。多年生杂类草群落土壤条件最好，1年生或2年生植物群落土壤条件最差，多年生根茎禾草和多年生丛生禾草没有显著差异。地上生物量与多样性关系表明，株高、叶面积和叶片氮含量等功能性状多样性比其它多样性指标能更好的预测生产力的变化，支持质量比假说，草地管理中应集中于优势种的管理。
秋季不同时间刈割，在第二年返青期和8月中旬地上生物量最大时期，多样性与生产力之间没有显著的关系。不同时间刈割对第二年草地的产量和质量没有显著影响，每年可以在8月中旬到9月上旬之间根据需要适时收获干草。春季5月下旬之前放牧或刈割不影响羊草种群和群落的产量和质量，后期放牧或刈割不利于羊草再生，群落中杂草比例增加。秋季或春季不同时间刈割，对群落植物物种丰富度和功能群丰富度都没有显著影响。秋季刈割或者春季刈割，群落高度性状与地上生物量有显著的正相关关系，物种丰富度、功能群丰富度和功能群多样性与地上生物量关系不显著。
氮添加对羊草的叶绿素a和b含量、类胡萝卜素含量、叶片氮含量、叶片干物质含量、比根长都有显著的作用；叶片的光合性状、比茎重和比根长之间有显著的相关性。羊草的叶绿素a和b含量、类胡萝卜素含量、叶片氮磷含量、叶片干物质含量、比茎重、株高与羊草的地上生物量都有显著的相关性。氮添加处理对羊草群落的物种丰富度和功能群丰富度都没有显著的影响，20g/m~2的氮添加处理羊草群落地上生物量最高。氮添加当年，地上生物量随着物种丰富度和功能群丰富度的增加有减少的趋势，但是不显著。
不同高度和密度立枯物处理间羊草的比叶面积、比茎重、植株高度、叶片数、叶绿素a和b含量、类胡萝卜素含量及叶片氮含量都有显著的差异。羊草各性状间，除了叶片厚度和比根长两个性状外，其它性状间都有较好的相关性。群落特征没有发生明显变化，随着植被低矮稀疏化，群落物种丰富度和地上生物量都有降低的趋势。土壤特征温度有显著的改变，但是其它土壤指标没有发生明显变化。植被高度变化对羊草性状、群落特征及土壤特征的影响要大于植被密度。
With the development of functional ecology, it is more and more important inresearch on functional plant ecology, especially with regard to the role of functionaldiversity on ecosystem functioning. This dissertation based on this point and studiedon functional plant ecology of grassland in Songnen Plain. We first discussed theproperty of Songnen grassland, and then, we stated functional traits properties and thevegetation conditions from plant functional traits perspective in Songnen grassland,third, we showed the variations of Leymus chinensis traits and L. chinensiscommunity characteristics after the control experiments by disturbance treatments ofdefoliation, nitrogen fertilization, vegetation shortness and sparseness, and last, wetested the relationships between aboveground biomass and species richness,functional group richness, species diversity and functional trait diversity at natural anddisturbed grassland. The main results as follows:
The soil pH, electrical conductivity, ion contents, exchangeable sodiumpercentage (ESP) of non-degraded grassland were9.1-10.2,198.7-643.5μs/cm,734.9-9297.5mg/kg,4.8-28.4%, respectively. The corresponding values were10.2-10.4,8240.3-18232.1mg/kg,459.3-1832.6μs/cm,43.0-54.0%for the degradedgrassland. The content of clay was the highest in every soil horigon. Soil clay (<0.002mm) of alkali and meadow soil was over50%in the illuvial horizon, soilphysical clay (<0.01mm) of swampy soil and meadow soil were over75%in theilluvial horizon. The aboveground biomass of the non-degraded grassland was morethan250g/m~2, and the aboveground biomass of degraded grassland was over300g/m~2. The monodominant L. chinensis communities were widely distributed, theaboveground biomass of L. chinensis made up97.2%of the community, the speciesrichness per m~2was no more than8species. The monodominant species was moreobvious in the degraded grassland, the richness was even only one per m~2for someannual plant communities. The soil salinity content was different at micro area wheregrew another mono-dominant community, therefore, the landscape of patches wasobvious in Songnen grassland. When the soil surface was disturbed and destroyed, theeluvial horizon of original salinization soil disappeared, then, the lower soil layerwhich contained higher salinity became bare surface or grew halophyte community.The disturbed land surface was the leading reason of soil salinization at Songnengrassland. The Songnen grassland were extremely different from the typical zonalsteppe soils all over the world, and it was a meadow which was determined by soilproperties.
The results of the66herbaceous plants leaf traits showed that leaf thickness (LT)variation (59.1%) was the highest among the7leaf traits measured. Pearsoncorrelation showed that specific leaf area (SLA), leaf nitrogen concentration (LNC), leaf phosphorus concentration (LPC), leaf chlorophyll (a+b) content (Chl) and leafcarotenoid content (Cd) were positively significantly correlated with each other. Therewas no significant correlation between LDMC and LPC, however, LDMC wasnegatively correlated with all other leaf traits significantly. LT was only negativelycorrelated to LDMC and SLA significantly, but not with others. LDMC, SLA, LT,LNC, LPC were significantly different among the4plant functional groups analyzedby one-way ANOVA (P <0.05). There was no difference between Chl and Cd.Perennial rhizome grass (PR) and perennial bunchgrasses (PB) had no differenceamong the7leaf traits. The LDMC of PR and PB were significantly higher thanperennial forbs (PF) and annuals/biannuals (AB). SLA, LNC, LPC of PR and PB werelower than PF and AB. Leaf N and P concentrations were22.7and1.8mg/g,respectively, N: P ratio was13.0. Plants growth were limited by nitrogen in Songnengrassland.
The plant community productivity and the proportion of the dominant specieswere high, and the community saturation was low in Songnen grassland. For the fourfunctional group communities, the species richness of PR and PF was highest, andthere was no significant difference of the functional group richness, and theaboveground biomass of AB was most. LDMC, SLA, LNC, LPC, plant height (PH),specific root length (SRL) were significantly different among the4plant functionalgroups. AB had high LDMC, LNC, LPC, and low SLA. The soil condition of PFcommunities were best and AB communities were worst, and there was no significantdifferent between PR and PB. The relationships between aboveground biomass anddiversity showed that the aggregated traits of PH, leaf area (LA), and LNC were betterthan other indices to predict the variation of the productivity. Our results were inagreement with the biomass ratio hypothesis. And the grassland management for themaintenance of ecosystem functioning should focus on the dominant species.
The relationship between diversity and aboveground biomass was not significantboth at the reviving stage and harvesting time after different autumn defoliation time.Defoliation treatments at autumn had no effect on grassland yield and quality in thesecond year. The harvest dates should be concentrated from the middle ten days ofAugust to the first ten days in September. Grazing or defoliation didn’t influence theyield and quality of L. chinensis population and community before late May. Grazingor defoliation limited L. chinensis regrowth, and the forbs proportion in thecommunity increased at later defoliation time. There were no significant difference ofspecies richness and functional group richness both after autumn or spring defoliation.However, there was significant positive relationship between aggregated PH andaboveground biomass, and the relationship between species richness, functional grouprichness, functional diversity and aboveground biomass were not significant.
Nitrogen addition had significant effected on Cha, Chb, Cd, LNC, LDMC andSRL of L. chinensis. There were significant correlations between leaf photosynthetic traits, stem specific density (SSD), and SRL. The relationships between Cha, Chb, Cd,LNC, LDMC, SSD, PH and aboveground biomass of L. chinensis were significant. Naddition had no significant effect on species richness, functional group richness, the20g/m~2treatment had highest aboveground biomass. The aboveground biomass haddecreased trend as the species richness and functional group richness increased, but itwas not significant.
The different height and density of reed stalks had significant effect on SLA,SSD, PH, leaf number (LN), Cha, Chb, Cd, LNC of L. chinensis. There weresignificant correlations between the traits of L. chinensis except LT and SRL. Therewas no substantial changes of the community characters. The species richness andfunctional group richness had shown decreased trends, and soil temperaturesincreased significantly as vegetation becoming shorter and sparser. However, therewere not obvious differences of other soil properties among the treatments. The effectof vegetation height on L. chinensis traits, community and soil properties was greaterthan that of vegetation density.
松嫩平原未退化羊草草地的土壤pH、电导率、离子含量(8种)、碱化度分别为9.1-10.2，198.7-643.5μs/cm，734.9-9297.5mg/kg，4.8-28.4%，退化裸地的土壤pH、电导率、离子含量、碱化度分别为10.2-10.4，8240.3-18232.1mg/kg，459.3-1832.6μs/cm，43.0-54.0%。主要土壤类型各发生层的粘粒含量最高，盐土和草甸土淀积层粘粒含量(<0.002mm)超过50%，沼泽土和草甸土淀积层物理粘粒(<0.01mm)含量超过75%。未退化草地地上生物量干重超过250g/m~2，退化草地地上生物量干重超过300g/m~2。单优势种群落广布，羊草的地上生物量干重与群落地上生物量干重的比例最高达97.2%，物种丰富度不超过8种/m~2。退化草地由于高盐碱条件限制，单优势现象更为明显，有些群落仅有1个物种。盐分含量有微域差异地段形成不同的单种优势种群落，斑块景观明显。地表被干扰，原初盐渍化草地土壤的淋溶层消失(践踏风蚀等)，下层高含盐量的盐碱土裸露成新地表，新地表裸露或生长盐生植物群落。地表被干扰是松嫩草地盐碱化的主导原因。松嫩平原草地，不同于世界各地的地带性草原土壤，是土壤决定的草甸植被类型。
松嫩草地66种草本植物7个叶片性状特征表明：叶片厚度变异系数最大，比叶面积、叶片氮含量、叶片磷含量、叶绿素含量和类胡萝卜素含量之间存在显著的正相关关系；叶片干物质含量与叶片磷含量没有显著的相关关系，与其它叶片性状呈显著的负相关关系；叶片厚度与叶片干物质含量和比叶面积呈显著的负相关关系，与其它叶片性状不相关。方差分析表明，叶片干物质含量、比叶面积、叶片厚度、叶片氮、叶片磷含量在4个功能群(多年生根茎禾草、多年生丛生禾草、多年生杂类草、1年生或2年生植物)间差异显著，叶绿素含量和类胡萝卜素含量在各个功能群间差异不显著。多重比较结果显示多年生根茎禾草和多年生丛生禾草叶片的7个性状差异不显著；多年生根茎禾草和多年生丛生禾草的叶片干物质含量高于多年生杂类草和1年生或2年生草本，其它性状小于这两个功能群。松嫩草地草本植物叶片氮磷含量分别为22.7和1.8mg/g，氮磷比为13.0，氮相对缺乏。
松嫩草地生产力高，群落饱和度低，优势种在群落中占的比例大。多年生根茎禾草和多年生杂类草功能群物种丰富度最高，各功能群间功能群丰富度没有显著差异，1年生或2年生植物群落地上生物量最高。叶片干物质含量、比叶面积、叶片氮含量、叶片磷含量、株高、比根长等性状在各功能群间差异显著。1年生或2年生植物功能群叶片干物质含量高、比叶面积小、氮磷含量高。多年生杂类草群落土壤条件最好，1年生或2年生植物群落土壤条件最差，多年生根茎禾草和多年生丛生禾草没有显著差异。地上生物量与多样性关系表明，株高、叶面积和叶片氮含量等功能性状多样性比其它多样性指标能更好的预测生产力的变化，支持质量比假说，草地管理中应集中于优势种的管理。
秋季不同时间刈割，在第二年返青期和8月中旬地上生物量最大时期，多样性与生产力之间没有显著的关系。不同时间刈割对第二年草地的产量和质量没有显著影响，每年可以在8月中旬到9月上旬之间根据需要适时收获干草。春季5月下旬之前放牧或刈割不影响羊草种群和群落的产量和质量，后期放牧或刈割不利于羊草再生，群落中杂草比例增加。秋季或春季不同时间刈割，对群落植物物种丰富度和功能群丰富度都没有显著影响。秋季刈割或者春季刈割，群落高度性状与地上生物量有显著的正相关关系，物种丰富度、功能群丰富度和功能群多样性与地上生物量关系不显著。
氮添加对羊草的叶绿素a和b含量、类胡萝卜素含量、叶片氮含量、叶片干物质含量、比根长都有显著的作用；叶片的光合性状、比茎重和比根长之间有显著的相关性。羊草的叶绿素a和b含量、类胡萝卜素含量、叶片氮磷含量、叶片干物质含量、比茎重、株高与羊草的地上生物量都有显著的相关性。氮添加处理对羊草群落的物种丰富度和功能群丰富度都没有显著的影响，20g/m~2的氮添加处理羊草群落地上生物量最高。氮添加当年，地上生物量随着物种丰富度和功能群丰富度的增加有减少的趋势，但是不显著。
不同高度和密度立枯物处理间羊草的比叶面积、比茎重、植株高度、叶片数、叶绿素a和b含量、类胡萝卜素含量及叶片氮含量都有显著的差异。羊草各性状间，除了叶片厚度和比根长两个性状外，其它性状间都有较好的相关性。群落特征没有发生明显变化，随着植被低矮稀疏化，群落物种丰富度和地上生物量都有降低的趋势。土壤特征温度有显著的改变，但是其它土壤指标没有发生明显变化。植被高度变化对羊草性状、群落特征及土壤特征的影响要大于植被密度。
With the development of functional ecology, it is more and more important inresearch on functional plant ecology, especially with regard to the role of functionaldiversity on ecosystem functioning. This dissertation based on this point and studiedon functional plant ecology of grassland in Songnen Plain. We first discussed theproperty of Songnen grassland, and then, we stated functional traits properties and thevegetation conditions from plant functional traits perspective in Songnen grassland,third, we showed the variations of Leymus chinensis traits and L. chinensiscommunity characteristics after the control experiments by disturbance treatments ofdefoliation, nitrogen fertilization, vegetation shortness and sparseness, and last, wetested the relationships between aboveground biomass and species richness,functional group richness, species diversity and functional trait diversity at natural anddisturbed grassland. The main results as follows:
The soil pH, electrical conductivity, ion contents, exchangeable sodiumpercentage (ESP) of non-degraded grassland were9.1-10.2,198.7-643.5μs/cm,734.9-9297.5mg/kg,4.8-28.4%, respectively. The corresponding values were10.2-10.4,8240.3-18232.1mg/kg,459.3-1832.6μs/cm,43.0-54.0%for the degradedgrassland. The content of clay was the highest in every soil horigon. Soil clay (<0.002mm) of alkali and meadow soil was over50%in the illuvial horizon, soilphysical clay (<0.01mm) of swampy soil and meadow soil were over75%in theilluvial horizon. The aboveground biomass of the non-degraded grassland was morethan250g/m~2, and the aboveground biomass of degraded grassland was over300g/m~2. The monodominant L. chinensis communities were widely distributed, theaboveground biomass of L. chinensis made up97.2%of the community, the speciesrichness per m~2was no more than8species. The monodominant species was moreobvious in the degraded grassland, the richness was even only one per m~2for someannual plant communities. The soil salinity content was different at micro area wheregrew another mono-dominant community, therefore, the landscape of patches wasobvious in Songnen grassland. When the soil surface was disturbed and destroyed, theeluvial horizon of original salinization soil disappeared, then, the lower soil layerwhich contained higher salinity became bare surface or grew halophyte community.The disturbed land surface was the leading reason of soil salinization at Songnengrassland. The Songnen grassland were extremely different from the typical zonalsteppe soils all over the world, and it was a meadow which was determined by soilproperties.
The results of the66herbaceous plants leaf traits showed that leaf thickness (LT)variation (59.1%) was the highest among the7leaf traits measured. Pearsoncorrelation showed that specific leaf area (SLA), leaf nitrogen concentration (LNC), leaf phosphorus concentration (LPC), leaf chlorophyll (a+b) content (Chl) and leafcarotenoid content (Cd) were positively significantly correlated with each other. Therewas no significant correlation between LDMC and LPC, however, LDMC wasnegatively correlated with all other leaf traits significantly. LT was only negativelycorrelated to LDMC and SLA significantly, but not with others. LDMC, SLA, LT,LNC, LPC were significantly different among the4plant functional groups analyzedby one-way ANOVA (P <0.05). There was no difference between Chl and Cd.Perennial rhizome grass (PR) and perennial bunchgrasses (PB) had no differenceamong the7leaf traits. The LDMC of PR and PB were significantly higher thanperennial forbs (PF) and annuals/biannuals (AB). SLA, LNC, LPC of PR and PB werelower than PF and AB. Leaf N and P concentrations were22.7and1.8mg/g,respectively, N: P ratio was13.0. Plants growth were limited by nitrogen in Songnengrassland.
The plant community productivity and the proportion of the dominant specieswere high, and the community saturation was low in Songnen grassland. For the fourfunctional group communities, the species richness of PR and PF was highest, andthere was no significant difference of the functional group richness, and theaboveground biomass of AB was most. LDMC, SLA, LNC, LPC, plant height (PH),specific root length (SRL) were significantly different among the4plant functionalgroups. AB had high LDMC, LNC, LPC, and low SLA. The soil condition of PFcommunities were best and AB communities were worst, and there was no significantdifferent between PR and PB. The relationships between aboveground biomass anddiversity showed that the aggregated traits of PH, leaf area (LA), and LNC were betterthan other indices to predict the variation of the productivity. Our results were inagreement with the biomass ratio hypothesis. And the grassland management for themaintenance of ecosystem functioning should focus on the dominant species.
The relationship between diversity and aboveground biomass was not significantboth at the reviving stage and harvesting time after different autumn defoliation time.Defoliation treatments at autumn had no effect on grassland yield and quality in thesecond year. The harvest dates should be concentrated from the middle ten days ofAugust to the first ten days in September. Grazing or defoliation didn’t influence theyield and quality of L. chinensis population and community before late May. Grazingor defoliation limited L. chinensis regrowth, and the forbs proportion in thecommunity increased at later defoliation time. There were no significant difference ofspecies richness and functional group richness both after autumn or spring defoliation.However, there was significant positive relationship between aggregated PH andaboveground biomass, and the relationship between species richness, functional grouprichness, functional diversity and aboveground biomass were not significant.
Nitrogen addition had significant effected on Cha, Chb, Cd, LNC, LDMC andSRL of L. chinensis. There were significant correlations between leaf photosynthetic traits, stem specific density (SSD), and SRL. The relationships between Cha, Chb, Cd,LNC, LDMC, SSD, PH and aboveground biomass of L. chinensis were significant. Naddition had no significant effect on species richness, functional group richness, the20g/m~2treatment had highest aboveground biomass. The aboveground biomass haddecreased trend as the species richness and functional group richness increased, but itwas not significant.
The different height and density of reed stalks had significant effect on SLA,SSD, PH, leaf number (LN), Cha, Chb, Cd, LNC of L. chinensis. There weresignificant correlations between the traits of L. chinensis except LT and SRL. Therewas no substantial changes of the community characters. The species richness andfunctional group richness had shown decreased trends, and soil temperaturesincreased significantly as vegetation becoming shorter and sparser. However, therewere not obvious differences of other soil properties among the treatments. The effectof vegetation height on L. chinensis traits, community and soil properties was greaterthan that of vegetation density.
引文
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