植物功能生物地理学的研究进展与展望

功能生物地理学研究性状及其多样性的时空分布变化、生态成因及其对生态系统功能的影响。近十来年,功能生物地理学领域发展迅速,性状数据呈指数增长,基于性状探索物种分布、群落结构和组成以及生态系统功能对环境变化响应的研究取得了重要进展。该文综述了植物功能生物地理学的核心内涵、发展历史、主要研究进展和未来展望。性状是功能生物地理学的研究核心,该文先总结了植物叶、茎、根、花、果实和种子等器官和整株关键性状的地理格局及其与环境间的关系,表明性状变异是植物适应进化和环境筛选的结果;概述了功能多样性的常用指标、地理分布与生态成因;介绍了性状数据的主要来源与性状缺失值的填充方法。随后,综述了植物性状间的关联与权衡,重点介绍了叶经济谱和植物经济谱的发展,指出其反映了植物对关键资源(如碳、养分和水分)的获取与分配策略;概述了基于性状预测物种分布的依据与进展,以及性状多样性与生态系统功能间的关系。在此基础上,提出了功能生物地理学研究所面临的挑战,强调未来研究要关注多性状在种内和种间的协同与权衡关系,将研究精度从物种水平推进到个体水平,采用性状网络等方法定量化性状间的关系及其对环境变化的响应,关注植物跨尺度的适应...     

植物功能性状、功能多样性与生态系统功能: 进展与展望

植物功能性状与生态系统功能是生态学研究的一个重要领域和热点问题。开展植物功能性状与生态系统功能的研究不仅有助于人类更好地应对全球变化情景下生物多样性丧失的生态学后果,而且能为生态恢复实践提供理论基础。近二十年来,该领域的研究迅速发展,并取得了一系列的重要研究成果,增强了人们对植物功能性状-生态系统功能关系的认识和理解。本文首先明确了植物功能性状的概念, 评述了近年来植物功能性状-生态系统功能关系领域的重要研究结果, 尤其是植物功能性状多样性-生态系统功能关系研究现状; 提出了未来植物功能性状与生态系统功能关系研究中应加强植物地上和地下性状之间关系及其与生态系统功能、植物功能性状与生态系统多功能性、不同时空尺度上植物功能性状与生态系统功能, 以及全球变化和消费者的影响等方面。     

木本植物茎叶功能性状及其关系随环境变化的研究进展

植物功能性状反映了植物对生长环境的响应和适应,是植物与环境相互作用的结果。植物功能性状之间的关系决定了植物的生活史对策,进而影响多物种的共存和生物多样性的维持。本文综述了叶大小、比叶面积、叶寿命和木质密度、叶大小-小枝大小、叶内生物量分配和叶大小-数量等随环境变化的研究进展,并简述其变化机理。植物功能性状及其关系的研究已经取得了很多成果,建立起了一套有效的研究方法和理论体系;但在今后的研究中,还需要对植物的地上部分和地下部分植物性状之间的协变关系、当年生末端小枝上功能性状间的关系的普遍性,以及叶大小-数量的等速关系在整个被子植物中的保守程度等几个方面进行深入研究。我国常绿阔叶林物种的植物功能性状及其关系研究较少,有待加强和系统化。     

叶脉网络功能性状及其生态学意义

叶脉网络结构是叶脉系统在叶片里的分布和排列样式。早期叶脉网络结构研究主要集中在其分类学意义上; 近年来叶脉网络功能性状及其在植物水分利用上的意义已成为植物生态学研究的热点。该文介绍了叶脉网络功能性状的指标体系(包括叶脉密度、叶脉直径、叶脉之间的距离、叶脉闭合度等), 综述了叶脉网络功能性状与叶脉系统功能(包括水分、养分和光合产物等物质运输、机械支撑和虫害防御等)的关系, 叶脉网络功能性状与叶片其他功能性状(包括比叶重、叶寿命、光合速率、叶片大小、气孔密度等)的协同变异和权衡关系, 以及叶脉网络功能性状随环境因子(包括水分、温度、光照等)的变化规律等方面的最新研究进展。此外, 叶脉网络功能性状的研究成果也被应用于古环境重建、城市交通规划、流域规划及全球变化研究中。由于叶脉网络功能性状是环境因子与系统发育共同作用的结果, 未来开展分子—叶片—植物—生态系统等多尺度的叶脉网络功能性状研究, 理清叶脉网络功能性状与气孔失水—茎干导水—根系吸水等植物水分利用的关系, 将为预测植物及生态系统对全球变化的响应提供新的启示。     

植物功能性状与湿地生态系统土壤碳汇功能

湿地生态系统碳平衡对气候变化极为敏感,是陆地生态系统碳循环响应全球变化的重要环节。然而,湿地生态系统碳汇调节机制仍不十分清楚,并且对影响因子的研究多集中在非生物因子上。综述了植物功能性状和功能性状多样性对湿地生态系统土壤碳汇功能的影响,阐明了生物因子对生态系统碳循环响应全球变化的重要性,介绍了植物功能性状对生态系统碳输入和输出过程的影响,简述了植物功能性状多样性的研究现状及其在指示生态系统碳汇功能现状和预测未来趋势等方面的应用。从优势植物、植物种间关系和植物-微生物种间关系3方面总结了植物功能性状多样性直接和间接影响生态系统碳循环的途径。展望了植物功能性状和功能性状多样性与湿地生态系统土壤碳汇功能的研究前景。     

植物群落构件的协同性研究进展

被子植物的营养构件如根、茎、叶以及繁殖构件如花、果实和种子等的结构及功能性状反映了物种对其生长环境的长期适应和响应。在植物种对环境的适应演化过程中,植物各构件在结构和功能上的变化不是孤立的、互不联系的,而是都具有内在的协调性、一致性,是协同进化的,各构件之间的关系决定了植物的生活史对策,进而影响群落(或生态系统)多物种的共存和生物多样性的维持。本文综述了群落水平上植物物种之间的各营养器官、各繁殖器官及二者之间结构性状及功能性状对环境适应协调性的生态学研究进展,不仅探究了植物构件间宏观性状的相关性,而且还涉及微观领域的研究成果及其相关机理,所涵盖的对象尺度有所扩大,是跨物种、跨群落或生态系统、甚至跨区域的。在形态学性状基础上,增加并拓展了其他学科的研究性状,例如解剖学性状、生理学性状、细胞学性状、遗传性状等。另外,也分析了相关研究存在的欠缺之处,并对今后的研究方向做出了展望。     

植物功能性状研究进展

植物功能性状是指植物体具有的与其定植、存活、生长和死亡紧密相关的一系列核心植物属性,且这些属性能够显著影响生态系统功能,并能够反映植被对环境变化的响应.越来越多的研究表明,相比大多数基于植物分类和数量的研究,植物功能性状在种群、群落和生态系统尺度上,都已成为解决重要生态学问题的可靠途径.本文回顾了植物功能性状研究的发展历程,总结了近10年来基于植物功能性状研究的前沿科学问题,包括功能性状的全球分布格局和内在关联,沿环境梯度的变化规律,功能多样性的定义及应用,与群落物种共存机制和群落动态变化的关系,与系统发育的关系,对生态系统功能的影响以及对各类干扰的影响和响应.尽管功能性状研究已经延伸到生态学领域的各个方面,有力推动了各个前沿科学问题的研究发展,仍然有很多值得关注和着重研究的方向.本文也对未来基于植物功能性状的研究,从性状测量和选取、研究方法以及研究方向上提出了展望,并指出,在当前全球气候变化背景下,功能性状也可应用于指导生物多样性保护和生态系统管理政策的制定.     

基于功能性状的毛乌素沙地不同演替阶段适生植物筛选

沙地生态系统修复是恢复生态学研究的热点问题,适生植物筛选是修复的关键。植物功能性状反映了植物在不同环境中的生存策略,探究沙地植物功能性状及其与环境之间的关系,有助于筛选用于植被恢复的物种,为保护沙地生态系统提供理论依据。以毛乌素沙地为研究区,分析了1983-2015年间沙地典型飞播样地群落演替特征及其对环境因子的响应,建立基于10个植物功能性状的毛乌素沙地潜在种库,进一步筛选飞播恢复下沙地不同演替阶段的适生植物。研究表明:(1)飞播恢复下的毛乌素沙地植物群落分为三个演替阶段:固沙先锋物种群落、沙生植物为主的杂类草群落、中生植物为主的杂类草群落。(2)土壤因子是群落演替的主要驱动力,其中土壤全氮、土壤总有机碳、土壤硝态氮是影响群落演替的关键因素。(3)基于功能性状筛选出29种适生物种用于植被恢复,演替第一阶段可用雾冰藜、猪毛菜等,演替第二阶段可用拂子茅、无芒隐子草等,演替第三阶段可用草地风毛菊、猪毛蒿等。通过物种功能性状特征可以快速选择适合沙地退化生态系统修复的候选物种,为植被恢复提供了一定的理论支持。     

新疆准噶尔荒漠植物叶片功能性状的进化和环境驱动机制初探

植物功能性状是由遗传因素和环境条件共同决定的。剖析各因素对植物性状变异的相对影响,对揭示植物对环境变化的响应和适应规律至关重要。作者以干旱区植物为研究对象,定量化分析了植物叶片功能性状变异及其与环境梯度的关系。研究区域位于中国新疆准噶尔盆地及其周边区域。在30个样地中,观测了110种植物的叶比重、叶片单位质量氮含量和单位面积氮含量以及叶片干物质含量,通过插值法获得每个样地的生物气候数据。结果表明:物种水平上叶片性状(性状值为每个物种的实际观测值)的变异在很大程度上由植物进化背景所决定,气候因子和功能群的作用次之;在群落尺度上(性状值为每个样地的权重和),叶比重与气候干旱程度呈正相关,单位质量氮含量在水热组合最优的区域出现最大值,而叶片干物质含量和单位面积氮含量与气候因子的相关性较小。叶比重是群落尺度上探讨叶片功能性状与环境梯度关系的一个合适的指标。此外,在研究植物性状-环境关系过程中,尽可能观测多个植物功能性状是必要的。但是,只有排除植物系统背景的影响,关于植物性状-环境关系的研究结论才可能接近真实情况。将来应该加强同一种内不同种群间的叶片性状的采样和分析工作。     

物种多样性和功能性状驱动高寒草原地上生物量对长期禁牧的响应

围封对草地生物多样性和初级生产力的影响是草地生态学研究的热点问题之一。基于2013—2021年在青藏高原东北部紫花针茅(Stipa purpurea)高寒草原围栏内外植物群落长期调查数据,从物种多样性、功能性状的角度解析了高寒草原地上生物量对长期围封的生态响应过程。结果表明：(1)围封处理对高寒草原物种多样性的负效应具有强烈的时间依赖性。围封处理显著提高地上生物量,但也显著降低了生物量稳定性和异步性,意味着高寒草原稳定的、可持续的生态系统服务功能被长期围封处理削弱。(2)植物功能性状对长期围封处理表现出差异性响应模式;与叶绿素性状相比,叶形态性状对长期围封处理表现出更强的敏感性。(3)物种多样性和功能性状与地上生物量之间均存在显著相关关系,并且物种多样性的影响被功能性状调控进而对地上生物量发挥间接效应,群落加权性状和功能分异度共同对草地生物量发挥直接的主导效应。研究结果证明了植物功能性状通过介导物种多样性与其共同驱动高寒草原地上生物量对长期围封的响应。因此,在未来草地管理过程中,同步研究植物物种和功能属性对于全面揭示生态系统的响应机制至关重要。     

Plant functional traits with particular reference to tropical deciduous forests: a review

Functional traits (FTs) integrate the ecological and evolutionary history of a species, and can potentially be used to predict its response as well as its influence on ecosystem functioning. Study of inter-specific variation in the FTs of plants aids in classifying species into plant functional types (PFTs) and provides insights into fundamental patterns and trade-offs in plant form and functioning and the effect of changing species composition on ecosystem functions. Specifically, this paper focuses on those FTs that make a species successful in the dry tropical environment. Following a brief overview, we discuss plant FTs that may be particularly relevant to tropical deciduous forests (TDFs). We consider the traits under the following categories: leaf traits, stem and root traits, reproductive traits, and traits particularly relevant to water availability. We compile quantitative information on functional traits of dry tropical forest species. We also discuss trait-based grouping of plants into PFTs. We recognize that there is incomplete knowledge about many FTs and their effects on TDFs and point out the need for further research on PFTs of TDF species, which can enable prediction of the dynamics of these forests in the face of disturbance and global climate change. Correlations between structural and ecophysiological traits and ecosystem functioning should also be established which could make it possible to generate predictions of changes in ecosystem services from changes in functional composition.     

Plant functional types and ecosystem function in relation to global change

Abstract. Plant functional types (PFTs) bridge the gap between plant physiology and community and ecosystem processes, thus providing a powerful tool in climate change research. We aimed at identifying PFTs within the flora of central-western Argentina, and to explore their possible consequences for ecosystem function. We analyzed 24 vegetative and regenerative traits of the 100 most abundant species along a steep climatic gradient. Based on plant traits and standard multivariate techniques, we identified eight PFTs. Our results confirmed, over a wide range of climatic conditions, the occurrence of broad recurrent patterns of association among plant traits reported for other floras; namely trade-offs between high investment in photosynthesis and growth on the one hand, and preferential allocation to storage and defence on the other. Regenerative traits were only partially coupled with vegetative traits. Using easily-measured plant traits and individual species cover in 63 sites, we predicted main community-ecosystem processes along the regional gradient. We hypothesized likely impacts of global climatic change on PFTs and ecosystems in situ, and analysed their probabilities of migrating in response to changing climatic conditions. Finally, we discuss the advantages and limitations of this kind of approach in predicting changes in plant distribution and in ecosystem processes over the next century.     

An Application of Plant Functional Types for Predicting Restoration Outcomes

We evaluated the restoration of native plant assemblages by topsoil translocation in the Hunter Valley, south‐east Australia. Species' responses were characterized by defining nine plant functional types (PFTs) based on combinations of four response mechanisms (seed bank persistence, germination cues, resprouting mechanisms, and longevity) through which species were predicted to persist or decline following translocation. The effects of community type and delay in topsoil restoration on restoration outcomes were tested in an orthogonal experiment. Changes in species' frequency were detected using Bayesian statistics with prior probabilities derived from pre‐clearing data. Few species failed to reestablish following translocation; these were offset by recruitment of other native species not detected prior to clearing. Compositional changes were more pronounced when topsoil was stockpiled (cf direct reinstatement), although there was no trend related to the period of stockpiling. The PFT response model correctly predicted the rank probability of decline in three of the nine PFTs, while a further three were correctly placed in the top ranks but in the incorrect order. Three PFTs were incorrectly ranked because the response model was incorrect. Resprouters declined more frequently than seeders; however, species with physical seed dormancy declined less frequently than those with either transient seed banks or physiological, morphological, or morpho‐physiological dormancy, irrespective of resprouting ability. Species with short juvenile periods were more likely to increase. We conclude that PFTs based on fire‐response traits represent a practical means of predicting species' responses to translocation and a basis for prioritizing species for supplementary planting.     

Plant Functional Types of Woody Species Related to Fire Disturbance in Forest–Grassland Ecotones

The study of plant functional types (PFTs) has been widely emphasized when analysing plant community changes in relation to variations in climate and disturbance regime. In this study, we search for PFTs of woody species near forest–grassland boundaries in South Brazil where, due to climate, forests tend to expand over grassland but are being restricted by frequent fires. We aimed at answering the questions: (i) which plant functional types of forest woody species can establish in adjacent grassland subject to fire disturbance and (ii) which plant functional types of forest and grassland woody species are related to short-term community dynamics in frequently burned grassland. Traits were assessed in woody plants in 156 plots (6.75 m²) arranged in 12 transects across forest–grassland boundaries with different fire history in their grassland part. The analysis used a recursive algorithm to search for traits and PFTs maximally associated to spatial distance from forest limit in one analysis, and elapsed time since last fire in another. As a result, nine PFTs of forest woody species were identified that best described community patterns associated to distance from forest. Resprouting ability characterized forest plants able to colonize grasslands. PFT diversity was higher in border plots than inside forest or grassland. Four PFTs of forest and grassland woody species best described woody species community patterns in the grassland associated to elapsed time since fire. Taller individuals of single-stemmed shrubs predominated in late post-fire recovery (3–4 years), while shorter multi-stemmed shrubs in recently burned areas (3 months to 1 year). PFTs of forest trees occurred in border plots or, as established adults, in grassland, remaining unaffected by fire. We conclude that easily measurable structural plant traits, such as those used in our study, are sufficient to evaluate post-fire community dynamics. Forest PFTs in burned grassland are restricted to those with resprouting ability to survive recurrent fire events. Establishment success is highest on protected sites with lesser or low-intensity fire.     

Using Plant Functional Traits and Phylogenies to Understand Patterns of Plant Community Assembly in a Seasonal Tropical Forest in Lao PDR

Plant functional traits reflect different evolutionary responses to environmental variation, and among extant species determine the outcomes of interactions between plants and their environment, including other plant species. Thus, combining phylogenetic and trait-based information can be a powerful approach for understanding community assembly processes across a range of spatial scales. We used this approach to investigate tree community composition at Phou Khao Khouay National Park (18°14’-18°32’N; 102°38’- 102°59’E), Laos, where several distinct forest types occur in close proximity. The aim of our study was to examine patterns of plant community assembly across the strong environmental gradients evident at our site. We hypothesized that differences in tree community composition were being driven by an underlying gradient in soil conditions. Thus, we predicted that environmental filtering would predominate at the site and that the filtering would be strongest on sandier soil with low pH, as these are the conditions least favorable to plant growth. We surveyed eleven 0.25 ha (50x50 m) plots for all trees above 10 cm dbh (1221 individual trees, including 47 families, 70 genera and 123 species) and sampled soils in each plot. For each species in the community, we measured 11 commonly studied plant functional traits covering both the leaf and wood economic spectrum traits and we reconstructed a phylogenetic tree for 115 of the species in the community using rbcL and matK sequences downloaded from Genebank (other species were not available). Finally we compared the distribution of trait values and species at two scales (among plots and 10x10m subplots) to examine trait and phylogenetic community structures. Although there was strong evidence that an underlying soil gradient was determining patterns of species composition at the site, our results did not support the hypothesis that the environmental filtering dominated community assembly processes. For the measured plant functional traits there was no consistent pattern of trait dispersion across the site, either when traits were considered individually or when combined in a multivariate analysis. However, there was a significant correlation between the degree of phylogenetic dispersion and the first principle component axis (PCA1) for the soil parameters. Moreover, the more phylogenetically clustered plots were on sandier soils with lower pH. Hence, we suggest that the community assembly processes across our site may reflect the influence of more conserved traits that we did not measure. Nevertheless, our results are equivocal and other interpretations are possible. Our study illustrates some difficulties in combining trait and phylogenetic approaches that may result from the complexities of integrating spatial and evolutionary processes that vary at different scales.     

Woody Plant Encroachment into Grasslands: Spatial Patterns of Functional Group Distribution and Community Development

Woody plant encroachment into grasslands has been globally widespread. The woody species invading grasslands represent a variety of contrasting plant functional groups and growth forms. Are some woody plant functional types (PFTs) better suited to invade grasslands than others? To what extent do local patterns of distribution and abundance of woody PFTs invading grasslands reflect intrinsic topoedaphic properties versus plant-induced changes in soil properties? We addressed these questions in the Southern Great Plains, United States at a subtropical grassland known to have been encroached upon by woody species over the past 50-100 years. A total of 20 woody species (9 tree-statured; 11 shrub-statured) were encountered along a transect extending from an upland into a playa basin. About half of the encroaching woody plants were potential N₂-fixers (55% of species), but they contributed only 7% to 16 % of the total basal area. Most species and the PFTs they represent were ubiquitously distributed along the topoedaphic gradient, but with varying abundances. Overstory-understory comparisons suggest that while future species composition of these woody communities is likely to change, PFT composition is not. Canonical correspondence analysis (CCA) ordination and variance partitioning (Partial CCA) indicated that woody species and PFT composition in developing woody communities was primarily influenced by intrinsic landscape location variables (e.g., soil texture) and secondarily by plant-induced changes in soil organic carbon and total nitrogen content. The ubiquitous distribution of species and PFTs suggests that woody plants are generally well-suited to a broad range of grassland topoedaphic settings. However, here we only examined categorical and non-quantitative functional traits. Although intrinsic soil properties exerted more control over the floristics of grassland-to-woodland succession did plant modifications of soil carbon and nitrogen concentrations, the latter are likely to influence productivity and nutrient cycling and may, over longer time-frames, feed back to influence PFT distributions.     

Functional traits and root morphology of alpine plants

Background and Aims

Vegetation has long been recognized to protect the soil from erosion. Understanding species differences in root morphology and functional traits is an important step to assess which species and species mixtures may provide erosion control. Furthermore, extending classification of plant functional types towards root traits may be a useful procedure in understanding important root functions.

Methods

In this study, pioneer data on traits of alpine plant species, i.e. plant height and shoot biomass, root depth, horizontal root spreading, root length, diameter, tensile strength, plant age and root biomass, from a disturbed site in the Swiss Alps are presented. The applicability of three classifications of plant functional types (PFTs), i.e. life form, growth form and root type, was examined for above- and below-ground plant traits.

Key Results

Plant traits differed considerably among species even of the same life form, e.g. in the case of total root length by more than two orders of magnitude. Within the same root diameter, species differed significantly in tensile strength: some species (Geum reptans and Luzula spicata) had roots more than twice as strong as those of other species. Species of different life forms provided different root functions (e.g. root depth and horizontal root spreading) that may be important for soil physical processes. All classifications of PFTs were helpful to categorize plant traits; however, the PFTs according to root type explained total root length far better than the other PFTs.

Conclusions

The results of the study illustrate the remarkable differences between root traits of alpine plants, some of which cannot be assessed from simple morphological inspection, e.g. tensile strength. PFT classification based on root traits seems useful to categorize plant traits, even though some patterns are better explained at the individual species level.     

Divergence of functional traits at early stages of development in Stipa tenacissima populations distributed along an environmental gradient of the Mediterranean

Assessing differences in plant functional traits (PFTs) along climatic gradients is potentially useful for understanding variation within and across populations, and for predicting their responses to climate change. This study investigates the intraspecific variability of several PFTs in Stipa tenacissima (Alpha grass) seedlings from different populations distributed across a climatic gradient. Seven populations from Tunisia to Spain within a 100–600 mm/year rainfall range were selected. Seedlings from each population were grown in a common garden. We expected the functional characteristics to differ among seedling populations according to their climatic gradient. The response patterns were helpful to predict acclimation and fitness under future climatic conditions in these populations. The seedling development analysis showed differences in PFTs among S. tenacissima populations. The biomass traits analysis revealed that higher above-ground biomass was related to higher below-ground development. The leaf traits proved that seedlings with longer leaf length would have less sclerophyllous leaves, a trade-off between productivity and drought resistance. The root traits analysis reflects seedling strategies to maximize resource uptake efficiency. PFTs showed several significant relationships with climatic conditions. The less rainfall, the higher plant allocation to root systems exploring soil. Higher mean temperatures were related to reduced root/plant development. The PFT analysis proves that species followed the ‘optimal partitioning theory’, in that plants preferentially allocate biomass to acquire the resource that most limits their development. However, both the environmental conditions and genetic diversity in S. tenacissima populations influenced seedling growth and behaviour to face ongoing climate change.

     

Plant functional types and climate at the global scale

Abstract. Globally applicable sets of terrestrial plant functional types (PFTs) have been identified as a major need in the development of dynamic global vegetation models for use with global atmospheric models. Global sets of PFTs should represent the world's most important plant types; characterize them through their functional behavior; and provide complete, geographically representative coverage of the world's land areas. Three main schools of thought on PFTs have emerged: (1) a physiological focus on internal function, especially at the level of basic metabolism; (2) an ecological focus on function in relation to plant form and environmental conditions; and (3) a geophysical focus on how plant functions affect the adjacent atmosphere. A structural approach based on pheno-physiognomy permits ready identification of relatively familiar, recognizable plant types. Many of the criteria cited by other approaches also are intimately related to structure and its seasonal changes. An earlier global system of structural-functional PFTs and their climatic relations has been improved, including addition of less well-known plant types, and is briefly described. A more strictly ‘functional’ approach is proposed, in which major aspects of plant function, initially metabolism and water balance, are used to classify functional types and suggest how these are constrained by climate. Such functional considerations, however, are closely linked to structural manifestations - but also require other functional criteria for more completely functional classifications. A recent global model of potential natural vegetation types suggested ca. 15 major plant types as necessary to cover the world's main terrestrial vegetation patterns. These essential types correspond well with a first-cut set of structural types implied by metabolic considerations.