延河流域植物功能性状对环境变化的响应和植物适应策略研究
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摘要
植物在长期适应环境的过程中，会通过内部不同功能性状之间的平衡变化来适应外部环境，以达到自身生存与繁衍的目的。因此，研究植物的功能性状及其关系沿环境梯度的变化规律，对于理解植物对环境的适应策略具有十分重要的意义。近年来，由于植物功能性状具有简单易于测定的特点，并且可以同时在不同植物的种类间进行比较研究，在植物与环境关系的研究中被广泛应用。黄土丘陵区是我国水土流失最严重的地区，植被恢复重建是改善该区环境的重要途径。如何针对不同的侵蚀环境，研究植物对环境的适应策略，从而构建适宜的植被体系是该区恢复生态学面临的重大科学问题之一。本文以延河流域为研究区，沿降雨与温度梯度，选择24个样地，共调查测定了149种植物(隶属于49科120属，其中草本植物102种，木本植物47种)的高度、比叶面积、叶面积、叶厚度、叶氮含量、叶组织密度、细根直径、比根长、细根组织密度、细根氮含量等功能性状特征，通过各个性状及其相互关系沿环境梯度的变化规律的分析，探究了植物对环境的适应机理和适应策略，取得的研究结果主要有：
1)延河流域植物功能性状的基本分布
延河流域各种植物的功能性状属性值变化范围很大。比叶面积均值为170.1mm~2·mg~(-1)，分布范围为5.26—9240.53mm~2·mg~(-1)；叶组织密度的均值为0.0989mg·mm~(-1)，分布在0.000789—1.39mg·mm~(-1)之间；叶氮含量均值为2.3%，分布在0.218—8.39%之间；植物的根氮含量、比根长、根组织密度分别在0.142—2.81%，0.158—66.77m·g~(-1)和0.0256—134.3mg·mm~(-1)之间波动，其平均值分别为：0.941%，6.67m·g~(-1)，26.82mg·mm-3。
2)延河流域植物功能性状的互补与替代
通过性状之间在功能上的平衡变化是植物适应环境的重要策略。本研究表明，延河流域叶性状之间的关系紧密，各叶性状均显著相关，其中比叶面积与叶氮含量正相关，但比叶面积与叶氮含量都与叶组织密度呈负相关；对根性状，除了比根长与根组织密度存在负相关关系，其余根性状间不存在相关关系。而不同器官间，根氮含量与比叶面积、叶氮含量正相关，与叶组织密度负相关；根组织密度与叶组织密度正相关，与比叶面积负相关，即叶片和细根对应的生理功能性状，如叶氮含量和根氮含量间存在显著关系，而结构性状，如比叶面积和比根长没有表现出显著性，即氮含量和组织密度在叶片和细根之间存在性状平行关系，其余性状则无平行性。
3)植物功能性状沿环境梯度的变化规律
植物功能性状沿环境梯度的变化规律反映了植物对延河流域环境变化的适应机理。研究表明，就植物功能性状而言，只有少数性状能够反映环境梯度变化信息，相比之下，性状间的关系更为紧密。比叶面积随着坡度的增加而增加；阳坡植物的叶片和细根氮含量大于阴坡；叶氮含量随着年均温度的升高而增加；随年均降雨量的增加，叶组织密度呈增加趋势，而细根氮含量则呈现出降低的趋势。同时，坡度较大地区(Slope>20°)的植物有较大的比叶面积，温暖或(和)隐域环境下的植物叶氮含量较高，而较干旱或(和)阳坡的植物叶片和细根氮含量均较高。
4)延河流域植物功能性状组合对环境梯度变化的响应
一些植物性状组合间存在较为稳定的关系，且不随环境因子的改变而改变，如根组织密度-比叶面积，根组织密度-叶组织密度和根组织密度-比根长，是延河流域不同种类的植物种为适应环境表现出的趋同进化。一些功能性状组合间关系则会随着环境因子的变化而变化，体现植物通过性状对动态变化在生长策略与防御策略间的平衡。在延河流域较温暖地区(MAT>9℃)，除了叶氮含量-叶组织密度组合的关系更显著外，其余性状组合间关系的斜率均不变，并呈现出一定比叶面积下，有更高的叶氮含量；一定叶组织密度时，根氮含量也更高的趋势。较干旱的地区(MAP<490mm)，根氮含量-叶氮含量间关系更为紧密，且当叶组织密度一定时，叶氮含量的增量较低。坡度并不影响延河流域植物性状组合间关系的标准化主轴斜率和截距。生长在阳坡的植物，叶氮含量-比叶面积和根氮含量-叶氮含量间关系更密切，且当比叶面积、叶组织密度一定时，相对阴坡的植物，具有更高的根氮含量。坡位也会影响性状组合间的关系，当比叶面积一定时，隐域环境下生长的植物具有较高的叶组织密度。总的来说，生长在温暖地区阳坡植物的叶氮含量-比叶面积，湿润地区阳坡植物的根氮含量-叶氮含量以及温暖地区植物的叶氮含量-叶组织密度间关系更为显著。
5）延河流域植物的CSR(Competitive-stress tolerant-ruderal scheme)策略
根据149种植物功能性状间的差异性，将物种划分为3大功能型，功能型I的植物具有最强的耐旱力和防御力，如代表性的地带性物种有辽东栎（Quercus liaotungensis），侧柏（Platycladus orientalis）、酸枣（Zizyphus jujuba）、荆条（Vitex negundo）等；功能型III的植物具有最强的养分维持能力用以对抗营养贫瘠的环境，有百里香（Thymusmongolicus）、草木樨状黄芪（Astragalus melilotoides）、灌木铁线莲（Clematis fruticosa），延安锦鸡儿（Caragana purdomii）等；功能型II植物居中，生长速率最高，具有较强的竞争力、分布最广，如代表性的物种有茭蒿（Artemisia giraldii）、蒙古蒿（Artemisiamongolica）、芦苇（Phragmites communi）、阿尔泰狗娃花（Heteropappus altaicus），土庄绣线菊（Spiraea pubescens）及软毛虫实（Corispermum puberullum）、小蓟（Circiumsegetum）、马齿苋（Portulaca oleracea）等分布广泛的杂草。其中，功能型I和III属于“胁迫忍耐型”策略，功能型II则属于“竞争型”和“干扰型”策略的综合。
6）延河流域植物的LHS(Leaf-height-seed strategy)策略
在原有植物叶片和细根性状的基础上加入2个植物种子性状（种子干重和种子体积），将延河流域30种植物划分为4个主要的功能型。功能型I的植物处于低胁迫的环境中，植株生长速率较快，易受到放牧干扰影响，属于干扰型和竞争，包括灰叶黄芪（Astragalus discolor）、砂珍棘豆（Oxytropis gracillima）、直立黄芪（Astragalusadsurgens）、南苜蓿（Medicago polymorpha）、虫实（Corispermum puberullum）、柴胡（Bupleurum scorzonerifolium）。功能型II的植物处于高胁迫的环境中，植物生长较慢，植株体将获得的营养大部分用来维持生命、抵抗外界压力，有臭草（Melica scabrosa）、白羊草（Bothriochloa ischaemum）、沙参（Adenophora elata）、铁杆蒿（Artemisia gmelinii）、阿尔泰狗娃花、蒙古蒿、委陵菜（Potentilla chinensis）。功能型III植物种子的扩散力和幼苗的成活率最强，能抵抗一定的胁迫压力，且最不易受外界放牧干扰影响，包括延安锦鸡儿、大针茅（Stipa grandis）、沙棘（Hippophae rhamnoidessi）、大蓟（Cirsiumjaponicum）、杠柳（Periploca sepium）、地梢瓜（Cynanchum thesioides）、灌木铁线莲、胡叶醉鱼草（Buddleya alternifolia）。功能型IV的植物，如虎榛子（Ostryopsis davidiana）、水栒子（Cotoneaster multiflorus）、黄刺玫（Rosa xanthina）、酸枣具有较强的繁殖力、对外界胁迫抵抗力较强具有较强的竞争力。
Plants use a variety of adaptive and functional strategies to adapt to environmentalchanges. Plants under specificenvironmental conditions usually share certain common traits,which allow plants to adapt to environment stress. The study of a plant’s functional traits iscurrently becoming a high priority area of plant ecology because these traits are both easy toquantify and are closely related to plant functioning. Consequently, variation in thesefunctional traits along environmental gradients reflects variation in plant adaptivemechanisms and each species’ interactions with climatic, edaphic or topographic drivers. Inhilly areas of the Loess Plateau, which is the focus area of national ecology restoration theefforts of revegetation has produced great achievements. However, serious soil erosion wasnot be kept within limits, the revegetation is still facingmany problems. Understanding planttraits in different habitats can provide scientific guidance for revegetation. The objective ofthis thesis was to assess the relationships and co-variation of plant functional traits alongenvironmental gradients, and to analyze the adaptive strategies of plants in semi-arid and aridenvironments. We measured plant height(PH), specific leaf area(SLA), leaf tissuedensity(LTD), leaf nitrogen concentrations(LNC), root nitrogen concentrations(RNC),specific root length(SRL) and fine root density(RTD) of149species at24sites in the YanheRiver watershed. By focusing on traits, our study uniquely demonstrates adaptivedifferentiation among species. These results will help the selection of species for restorationprograms in the hilly areas in Loess Plateau. The main results are as follows:
1) The general pattern of plant traits
Six plant attributes across all the149species of Yanhe River watershed showed largevariations, with the following range:5.26—9240.53mm~2·mg~(-1)for specific leaf area;0.000789—1.39mg·mm~(-1)for leaf tissue density;0.218—8.39%for leaf nitrogen concentrations;0.142—2.81%for root nitrogen concentrations;0.158—66.77m·g~(-1)for specific root length; and0.0256—134.3mg·mm~(-1)for root tissue density. The atithmetic means of these parameterswere170.1mm~2·mg~(-1),0.0989mg·mm~(-1),2.3%,0.941%,6.67m·g~(-1)and26.82mg·mm-3.
2) Summary of relationships among plant traits
We analyzed the patterns of correlations among six organ-level traits and how these traitsallow different species to deal with environmental conditions. The results showed that thesix plant functional traits of the149species were closely related. Leaf nitrogen concentrationswere positively related to specific leaf area and root nitrogen concentrations, and a negativelyrelated to leaf tissue density. Specific root length was negatively related to fine root tissuedensity, whereas no correlation existed between leaf and root traits except for root nitrogenconcentrations.
3) Relationships between plant traits and environmental variables
In general, the correlation between individual traits and environmental variables is nothigh. Specific leaf area increased with slope, and leaf nitrogen concentrations increased withincrease of mean annual temperature. As mean annual precipitation increased, leaf tissuedensity increased and root nitrogen concentrations decreased. Meanwhile, the mean specificleaf area was higher at steep gradient; leaf nitrogen concentrations was higher at hotter,hidden region; and mean leaf and root nitrogen concentrations were higher at drier, sunnyslope sites.
4) Variation of trait relationships along environmental gradient
The combinations of RTD-SLA，RTD-LTD and RTD-SRL recur in distantly related taxaacross Yanhe River watershed, which provide evidence for convergent evolution. However, athotter sites, average LNC was higher at a given SLA, RNC was higher at a given LTD. Atdrier sites, LNC was lower at a given LTD. RNC was higher at a given SLA or LTD at sunnyslope sites. And LTD was higher at a given SLA at hidden regions.
5) A multi-trait test of CSR(Competitive-stress tolerant-ruderal scheme) plant strategyscheme
Based on the dissimilarity of the six traits, the species were classified into three mainplant functional types (PFTs). The species of PFT I are best adapted to an arid environment,with the greatest resistance and resistance to herbivore and physical damage. The species ofPFT III may avoid nutrient stress by having the greatest nutrient conservation efficiency,which is believed to be important in minimizing nutrient losses in a nutrient-poorenvironment. The PFT II group is somewhat intermediate with a greater growth rate, highercompetitive ability and wider distribution in the study area. Based on CSR Triangle theoriesof Grime et al.(1979), both PFT I and PFT III plants adopt a stress-tolerance strategy to theenvironment (S strategy) while PFT II plants use a combination of competitiveness (C) andruderality (R) strategies.
6) A multi-trait test of LHS(Leaf-height-seed strategy) plant strategy scheme
Here we investigated30species and plus2seed traits(seed mass, seed area) to test Westoby’s LHS strategy scheme(1998) in Yanhe River watershed, and the species wereclassified into four main plant functional types. The species of PFT I are the combination ofrapid growth of grazing pressure individuals with low stress. The species of PFT II may avoidstress by having the greatest nutrient conservation efficiency, which is believed to beimportant in minimizing nutrient losses in a nutrient-poor environment. The species of PFTIII may avoid stress and grazing pressure, with high-effective colonizing abilities. The PFT IVgroup is higher competitive, effective colonizing ability and wider distribution in the studyarea.
1)延河流域植物功能性状的基本分布
延河流域各种植物的功能性状属性值变化范围很大。比叶面积均值为170.1mm~2·mg~(-1)，分布范围为5.26—9240.53mm~2·mg~(-1)；叶组织密度的均值为0.0989mg·mm~(-1)，分布在0.000789—1.39mg·mm~(-1)之间；叶氮含量均值为2.3%，分布在0.218—8.39%之间；植物的根氮含量、比根长、根组织密度分别在0.142—2.81%，0.158—66.77m·g~(-1)和0.0256—134.3mg·mm~(-1)之间波动，其平均值分别为：0.941%，6.67m·g~(-1)，26.82mg·mm-3。
2)延河流域植物功能性状的互补与替代
通过性状之间在功能上的平衡变化是植物适应环境的重要策略。本研究表明，延河流域叶性状之间的关系紧密，各叶性状均显著相关，其中比叶面积与叶氮含量正相关，但比叶面积与叶氮含量都与叶组织密度呈负相关；对根性状，除了比根长与根组织密度存在负相关关系，其余根性状间不存在相关关系。而不同器官间，根氮含量与比叶面积、叶氮含量正相关，与叶组织密度负相关；根组织密度与叶组织密度正相关，与比叶面积负相关，即叶片和细根对应的生理功能性状，如叶氮含量和根氮含量间存在显著关系，而结构性状，如比叶面积和比根长没有表现出显著性，即氮含量和组织密度在叶片和细根之间存在性状平行关系，其余性状则无平行性。
3)植物功能性状沿环境梯度的变化规律
植物功能性状沿环境梯度的变化规律反映了植物对延河流域环境变化的适应机理。研究表明，就植物功能性状而言，只有少数性状能够反映环境梯度变化信息，相比之下，性状间的关系更为紧密。比叶面积随着坡度的增加而增加；阳坡植物的叶片和细根氮含量大于阴坡；叶氮含量随着年均温度的升高而增加；随年均降雨量的增加，叶组织密度呈增加趋势，而细根氮含量则呈现出降低的趋势。同时，坡度较大地区(Slope>20°)的植物有较大的比叶面积，温暖或(和)隐域环境下的植物叶氮含量较高，而较干旱或(和)阳坡的植物叶片和细根氮含量均较高。
4)延河流域植物功能性状组合对环境梯度变化的响应
一些植物性状组合间存在较为稳定的关系，且不随环境因子的改变而改变，如根组织密度-比叶面积，根组织密度-叶组织密度和根组织密度-比根长，是延河流域不同种类的植物种为适应环境表现出的趋同进化。一些功能性状组合间关系则会随着环境因子的变化而变化，体现植物通过性状对动态变化在生长策略与防御策略间的平衡。在延河流域较温暖地区(MAT>9℃)，除了叶氮含量-叶组织密度组合的关系更显著外，其余性状组合间关系的斜率均不变，并呈现出一定比叶面积下，有更高的叶氮含量；一定叶组织密度时，根氮含量也更高的趋势。较干旱的地区(MAP<490mm)，根氮含量-叶氮含量间关系更为紧密，且当叶组织密度一定时，叶氮含量的增量较低。坡度并不影响延河流域植物性状组合间关系的标准化主轴斜率和截距。生长在阳坡的植物，叶氮含量-比叶面积和根氮含量-叶氮含量间关系更密切，且当比叶面积、叶组织密度一定时，相对阴坡的植物，具有更高的根氮含量。坡位也会影响性状组合间的关系，当比叶面积一定时，隐域环境下生长的植物具有较高的叶组织密度。总的来说，生长在温暖地区阳坡植物的叶氮含量-比叶面积，湿润地区阳坡植物的根氮含量-叶氮含量以及温暖地区植物的叶氮含量-叶组织密度间关系更为显著。
5）延河流域植物的CSR(Competitive-stress tolerant-ruderal scheme)策略
根据149种植物功能性状间的差异性，将物种划分为3大功能型，功能型I的植物具有最强的耐旱力和防御力，如代表性的地带性物种有辽东栎（Quercus liaotungensis），侧柏（Platycladus orientalis）、酸枣（Zizyphus jujuba）、荆条（Vitex negundo）等；功能型III的植物具有最强的养分维持能力用以对抗营养贫瘠的环境，有百里香（Thymusmongolicus）、草木樨状黄芪（Astragalus melilotoides）、灌木铁线莲（Clematis fruticosa），延安锦鸡儿（Caragana purdomii）等；功能型II植物居中，生长速率最高，具有较强的竞争力、分布最广，如代表性的物种有茭蒿（Artemisia giraldii）、蒙古蒿（Artemisiamongolica）、芦苇（Phragmites communi）、阿尔泰狗娃花（Heteropappus altaicus），土庄绣线菊（Spiraea pubescens）及软毛虫实（Corispermum puberullum）、小蓟（Circiumsegetum）、马齿苋（Portulaca oleracea）等分布广泛的杂草。其中，功能型I和III属于“胁迫忍耐型”策略，功能型II则属于“竞争型”和“干扰型”策略的综合。
6）延河流域植物的LHS(Leaf-height-seed strategy)策略
在原有植物叶片和细根性状的基础上加入2个植物种子性状（种子干重和种子体积），将延河流域30种植物划分为4个主要的功能型。功能型I的植物处于低胁迫的环境中，植株生长速率较快，易受到放牧干扰影响，属于干扰型和竞争，包括灰叶黄芪（Astragalus discolor）、砂珍棘豆（Oxytropis gracillima）、直立黄芪（Astragalusadsurgens）、南苜蓿（Medicago polymorpha）、虫实（Corispermum puberullum）、柴胡（Bupleurum scorzonerifolium）。功能型II的植物处于高胁迫的环境中，植物生长较慢，植株体将获得的营养大部分用来维持生命、抵抗外界压力，有臭草（Melica scabrosa）、白羊草（Bothriochloa ischaemum）、沙参（Adenophora elata）、铁杆蒿（Artemisia gmelinii）、阿尔泰狗娃花、蒙古蒿、委陵菜（Potentilla chinensis）。功能型III植物种子的扩散力和幼苗的成活率最强，能抵抗一定的胁迫压力，且最不易受外界放牧干扰影响，包括延安锦鸡儿、大针茅（Stipa grandis）、沙棘（Hippophae rhamnoidessi）、大蓟（Cirsiumjaponicum）、杠柳（Periploca sepium）、地梢瓜（Cynanchum thesioides）、灌木铁线莲、胡叶醉鱼草（Buddleya alternifolia）。功能型IV的植物，如虎榛子（Ostryopsis davidiana）、水栒子（Cotoneaster multiflorus）、黄刺玫（Rosa xanthina）、酸枣具有较强的繁殖力、对外界胁迫抵抗力较强具有较强的竞争力。
Plants use a variety of adaptive and functional strategies to adapt to environmentalchanges. Plants under specificenvironmental conditions usually share certain common traits,which allow plants to adapt to environment stress. The study of a plant’s functional traits iscurrently becoming a high priority area of plant ecology because these traits are both easy toquantify and are closely related to plant functioning. Consequently, variation in thesefunctional traits along environmental gradients reflects variation in plant adaptivemechanisms and each species’ interactions with climatic, edaphic or topographic drivers. Inhilly areas of the Loess Plateau, which is the focus area of national ecology restoration theefforts of revegetation has produced great achievements. However, serious soil erosion wasnot be kept within limits, the revegetation is still facingmany problems. Understanding planttraits in different habitats can provide scientific guidance for revegetation. The objective ofthis thesis was to assess the relationships and co-variation of plant functional traits alongenvironmental gradients, and to analyze the adaptive strategies of plants in semi-arid and aridenvironments. We measured plant height(PH), specific leaf area(SLA), leaf tissuedensity(LTD), leaf nitrogen concentrations(LNC), root nitrogen concentrations(RNC),specific root length(SRL) and fine root density(RTD) of149species at24sites in the YanheRiver watershed. By focusing on traits, our study uniquely demonstrates adaptivedifferentiation among species. These results will help the selection of species for restorationprograms in the hilly areas in Loess Plateau. The main results are as follows:
1) The general pattern of plant traits
Six plant attributes across all the149species of Yanhe River watershed showed largevariations, with the following range:5.26—9240.53mm~2·mg~(-1)for specific leaf area;0.000789—1.39mg·mm~(-1)for leaf tissue density;0.218—8.39%for leaf nitrogen concentrations;0.142—2.81%for root nitrogen concentrations;0.158—66.77m·g~(-1)for specific root length; and0.0256—134.3mg·mm~(-1)for root tissue density. The atithmetic means of these parameterswere170.1mm~2·mg~(-1),0.0989mg·mm~(-1),2.3%,0.941%,6.67m·g~(-1)and26.82mg·mm-3.
2) Summary of relationships among plant traits
We analyzed the patterns of correlations among six organ-level traits and how these traitsallow different species to deal with environmental conditions. The results showed that thesix plant functional traits of the149species were closely related. Leaf nitrogen concentrationswere positively related to specific leaf area and root nitrogen concentrations, and a negativelyrelated to leaf tissue density. Specific root length was negatively related to fine root tissuedensity, whereas no correlation existed between leaf and root traits except for root nitrogenconcentrations.
3) Relationships between plant traits and environmental variables
In general, the correlation between individual traits and environmental variables is nothigh. Specific leaf area increased with slope, and leaf nitrogen concentrations increased withincrease of mean annual temperature. As mean annual precipitation increased, leaf tissuedensity increased and root nitrogen concentrations decreased. Meanwhile, the mean specificleaf area was higher at steep gradient; leaf nitrogen concentrations was higher at hotter,hidden region; and mean leaf and root nitrogen concentrations were higher at drier, sunnyslope sites.
4) Variation of trait relationships along environmental gradient
The combinations of RTD-SLA，RTD-LTD and RTD-SRL recur in distantly related taxaacross Yanhe River watershed, which provide evidence for convergent evolution. However, athotter sites, average LNC was higher at a given SLA, RNC was higher at a given LTD. Atdrier sites, LNC was lower at a given LTD. RNC was higher at a given SLA or LTD at sunnyslope sites. And LTD was higher at a given SLA at hidden regions.
5) A multi-trait test of CSR(Competitive-stress tolerant-ruderal scheme) plant strategyscheme
Based on the dissimilarity of the six traits, the species were classified into three mainplant functional types (PFTs). The species of PFT I are best adapted to an arid environment,with the greatest resistance and resistance to herbivore and physical damage. The species ofPFT III may avoid nutrient stress by having the greatest nutrient conservation efficiency,which is believed to be important in minimizing nutrient losses in a nutrient-poorenvironment. The PFT II group is somewhat intermediate with a greater growth rate, highercompetitive ability and wider distribution in the study area. Based on CSR Triangle theoriesof Grime et al.(1979), both PFT I and PFT III plants adopt a stress-tolerance strategy to theenvironment (S strategy) while PFT II plants use a combination of competitiveness (C) andruderality (R) strategies.
6) A multi-trait test of LHS(Leaf-height-seed strategy) plant strategy scheme
Here we investigated30species and plus2seed traits(seed mass, seed area) to test Westoby’s LHS strategy scheme(1998) in Yanhe River watershed, and the species wereclassified into four main plant functional types. The species of PFT I are the combination ofrapid growth of grazing pressure individuals with low stress. The species of PFT II may avoidstress by having the greatest nutrient conservation efficiency, which is believed to beimportant in minimizing nutrient losses in a nutrient-poor environment. The species of PFTIII may avoid stress and grazing pressure, with high-effective colonizing abilities. The PFT IVgroup is higher competitive, effective colonizing ability and wider distribution in the studyarea.
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