氮添加对中国陆地植被地上-地下生物量分配的影响

大气氮沉降水平持续升高导致的外源氮输入增加,强烈影响了陆地生态系统的碳循环。目前,已有大量报道证实了氮沉降升高对全球陆地植被固碳的积极影响。虽然之前大部分研究将这一结果归因于光合作用增强导致的地上生物量增加,但最近的研究发现长期氮添加对植物地下根系的影响也同样重要。归纳整理了181篇公开发表的我国野外模拟氮沉降试验结果,采用整合分析(Meta-analysis)方法,定量评估了氮添加对我国陆地植被地上-地下生物量分配的影响特征和不同生态系统类型及施氮方式之间的影响差异。通过分析地上-地下生物量分配对氮添加的响应差异来探究植被碳增益对长期大气氮沉降增加的潜在响应机制。结果表明,氮添加显著增强了我国陆地植被的光合作用及碳固存,且植物碳增益在不同生态系统类型及施氮制度间有所差异。植物叶片的氮含量显著增加,使得叶片碳氮比及凋落物碳氮比显著降低,但并未显著影响细根的碳氮比。氮添加总体上显著提高了植物的净光合速率,但降低了光合利用效率。地上生物量,凋落物产量和根生物量平均分别显著增加了38%,17%和18%,总体上植物地上部分对氮添加的响应程度比地下部分更高。然而,不同生态系统类型的地上-地下生物...     

陆生植物生物量分配对模拟氮沉降响应的Meta分析

分析了陆生植物地上、地下各组织中生物量分配对氮沉降的响应,为研究大气氮沉降背景下陆地生态系统的碳、氮循环过程及植物生物量分配、立木收获、定向培育等相关研究和实践提供参考依据。共收集整理了国内外63篇论文的原始数据资料进行Meta分析(Meta-analysis),用以定量评估氮沉降对植物生物量分配的影响,并通过亚组分析进一步探讨了不同生态系统类型、植物种类、氮肥形式、施氮水平和持续时间对生物量分配的影响。结果表明,总体来看施氮会显著促进植物地上部分生物量分配,植物叶生物量和茎生物量在施氮条件下均显著增加;然而地下生物量所受促进作用要低于地上部分,表现为植物细根生物量和粗根生物量在氮输入下并没有显著变化;植物根冠比在氮沉降下显著降低;叶重比、茎重比和根重比在氮沉降下没有显著变化。此外,亚组分析结果表明生态系统类型和植物类型会显著影响植物总生物量和根冠比对氮沉降的响应,草本植物在氮沉降下的生物量累积明显优于木本,这说明短期氮沉降可能会增加草本的覆盖面积;施肥形式对根冠比的影响存在明显差异,相比于尿素,硝酸铵对植物根冠比的作用更显著;不同施氮水平显著影响地上生物量分配,中氮水平(本研究为60—120 kg hm-2a-1)促进作用最大,高氮水平(本研究为≥120 kg hm-2a-1)促进作用明显减弱,这与总生物量的变化一致,表明过高的氮沉降量将抑制植物生长;氮沉降处理时间长短对植物地上生物量的影响也存在显著差异,当施氮时间高于3年,氮沉降对地上生物量的促进作用几乎消失。总之,短期氮沉降会使植物分配更多生物量给地上部分,且氮沉降对草本植物生物量的累积作用明显优于木本,这些发现可为未来大气氮沉降背景下植物地上、地下部分碳存储、植物群落结构、植被动态等相关研究提供科学依据。     

Nitrogen addition affects plant biomass allocation but not allometric relationships among different organs across the globe

全球尺度上氮添加影响植物生物量分配但不影响不同器官间的异速生长关系 生物量在不同器官间的分配是植物对环境变化响应的一个关键生态生理学过程。然而,在全球尺度上有关不同陆地生态系统植物生物量分配对氮沉降响应的认识还比较欠缺。本文通过整合分析333篇已发表文章的5474个观测值,基于“最优分配假说”和“异速分配假说”,评估了全球尺度上氮添加对植物生物量及其在不同器官间分配的影响。结果表明：(1)氮添加显著增加了整株植物或不同器官的生物量,降低了根冠比和根质量分数,但对叶质量分数和茎质量分数无显著影响;(2)氮添加对不同器官质量分数的影响受实验条件、植物功能性状、纬度、氮添加率等因子单独或交互作用的调控;(3)氮添加对生物量在不同器官间的异速分配率无显著影响,表明氮添加导致的根冠比和根质量分数减少是在异速分配模式下由整株植物生物量增加而致。虽然氮添加改变了地上和地下部分的质量分数,但不同器官生物量间异速分配模式的稳定性表明“异速分配假说”能更好地描述植物生物量分配对氮添加的响应规律。该研究结果将有助于深入认识氮沉降环境下植物生物量的分配规律,同时表明将生物量分配纳入有关氮沉降对陆地生态系统影响的预测模型中的重要性。     

陆地生态系统植物的氮源及氮素吸收

氮是植物生长发育所必需的营养元素,也是其主要的限制因子之一.陆地生态系统植物所需氮的来源及植物对氮素的吸收利用均受控于其种类和生长环境.环境条件的改变,一方面可能改变植物生长区原有氮的形态、浓度、赋存方式等,从而改变氮对植物的供给状况;另一方面可能引起植物生长区土壤质量、水分利用状况、光照等的改变,从而产生耦合现象,直接影响植物的生理生态特性,使植物对氮素的吸收利用发生改变,导致植物生长区的种群类型及物种多样性发生改变,并直接影响到生态系统的功能及演替.本文主要对陆地生态系统中高等植物生长发育所需氮素的来源及植物对氮素吸收利用过程中的影响因素进行了综述和讨论,并结合国内外在该领域的研究现状对其研究前景进行了展望.     

Biochar amendment boosts photosynthesis and biomass in C3 but not C4 plants: A global synthesis

Biochar is a carbon (C)‐rich solid produced from the thermochemical pyrolysis of biomass. Its amendment to soils has been proposed as a promising mean to mitigate greenhouse gas emissions and simultaneously benefit agricultural crops. However, how biochar amendment affects plant photosynthesis and growth remains unclear, especially on a global scale. In this study, we conducted a global synthesis of 74 publications with 347 paired comparisons to acquire an overall tendency of plant photosynthesis and growth following biochar amendment. Overall, we found that biochar amendment significantly increased photosynthetic rate by 27.1%, and improved stomatal conductance, transpiration rate, water use efficiency, and chlorophyll concentration by 19.6%, 26.9%, 26.8%, and 16.1%, respectively. Meanwhile, plant total biomass, shoot biomass, and root biomass increased by 25.4%, 22.1%, and 34.4%, respectively. Interestingly, plant types (C₃ and C₄ plants) showed greater control over plant photosynthesis and biomass than a broad suite of soil and biochar factors. Biochar amendment largely boosted photosynthesis and biomass on C₃ plants, but had a limited effect on C₄ plants. Our results highlight the importance of the differential response of plant types to biochar amendment with respect to plant growth and photosynthesis, providing a scientific foundation for making reasonable strategies towards an extensive application of biochar for agricultural production management.     

Aggravated phosphorus limitation on biomass production under increasing nitrogen loading: a meta‐analysis

Nitrogen (N) and phosphorus (P), either individually or in combination, have been demonstrated to limit biomass production in terrestrial ecosystems. Field studies have been extensively synthesized to assess global patterns of N impacts on terrestrial ecosystem processes. However, to our knowledge, no synthesis has been done so far to reveal global patterns of P impacts on terrestrial ecosystems, especially under different nitrogen (N) levels. Here, we conducted a meta‐analysis of impacts of P addition, either alone or with N addition, on aboveground (AGB) and belowground biomass production (BGB), plant and soil P concentrations, and N : P ratio in terrestrial ecosystems. Overall, our meta‐analysis quantitatively confirmed existing notions: (i) colimitation of N and P on biomass production and (ii) more P limitation in tropical forest than other ecosystems. More importantly, our analysis revealed new findings: (i) P limitation on biomass production was aggravated by N enrichment and (ii) plant P concentration was a better indicator of P limitation than soil P availability. Specifically, P addition increased AGB and BGB by 34% and 13%, respectively. The effect size of P addition on biomass production was larger in tropical forest than grassland, wetland, and tundra and varied with P fertilizer forms, P addition rates, or experimental durations. The P‐induced increase in biomass production and plant P concentration was larger under elevated than ambient N. Our findings suggest that the global limitation of P on biomass production will become severer under increasing N fertilizer and deposition in the future.     

A new method to estimate branch biomass from terrestrial laser scanning data by bridging tree structure models

Background and AimsBranch biomass and other attributes are important for estimating the carbon budget of forest stands and characterizing crown structure. As destructive measuring is time-consuming and labour-intensive, terrestrial laser scanning (TLS) as a solution has been used to estimate branch biomass quickly and non-destructively. However, branch information extraction from TLS data alone is challenging due to occlusion and other defects, especially for estimating individual branch attributes in coniferous trees.MethodsThis study presents a method, entitled TSM_tls, to estimate individual branch biomass non-destructively and accurately by combining tree structure models and TLS data. The TSM_tls method constructs the stem-taper curve from TLS data, then uses tree structure models to determine the number, basal area and biomass of individual branches at whorl level. We estimated the tree structural model parameters from 122 destructively measured Scots pine (Pinus sylvestris) trees and tested the method on six Scots pine trees that were first TLS-scanned and later destructively measured. Additionally, we estimated the branch biomass using other TLS-based approaches for comparison.Key ResultsTree-level branch biomass estimates derived from TSM_tls showed the best agreement with the destructive measurements [coefficient of variation of root mean square error (CV-RMSE) = 9.66 % and concordance correlation coefficient (CCC) = 0.99], outperforming the other TLS-based approaches (CV-RMSE 12.97–57.45 % and CCC 0.43–0.98 ). Whorl-level individual branch attributes estimates produced from TSM_tls showed more accurate results than those produced from TLS data directly.ConclusionsThe results showed that the TSM_tls method proposed in this study holds promise for extension to more species and larger areas.     

Global response patterns of plant photosynthesis to nitrogen addition: A meta‐analysis

A mechanistic understanding of plant photosynthetic response is needed to reliably predict changes in terrestrial carbon (C) gain under conditions of chronically elevated atmospheric nitrogen (N) deposition. Here, using 2,683 observations from 240 journal articles, we conducted a global meta‐analysis to reveal effects of N addition on 14 photosynthesis‐related traits and affecting moderators. We found that across 320 terrestrial plant species, leaf N was enhanced comparably on mass basis (N_mass, +18.4%) and area basis (N_area, +14.3%), with no changes in specific leaf area or leaf mass per area. Total leaf area (TLA) was increased significantly, as indicated by the increases in total leaf biomass (+46.5%), leaf area per plant (+29.7%), and leaf area index (LAI, +24.4%). To a lesser extent than for TLA, N addition significantly enhanced leaf photosynthetic rate per area (A_area, +12.6%), stomatal conductance (g_s, +7.5%), and transpiration rate (E, +10.5%). The responses of A_area were positively related with that of g_s, with no changes in instantaneous water‐use efficiency and only slight increases in long‐term water‐use efficiency (+2.5%) inferred from ¹³C composition. The responses of traits depended on biological, experimental, and environmental moderators. As experimental duration and N load increased, the responses of LAI and A_area diminished while that of E increased significantly. The observed patterns of increases in both TLA and E indicate that N deposition will increase the amount of water used by plants. Taken together, N deposition will enhance gross photosynthetic C gain of the terrestrial plants while increasing their water loss to the atmosphere, but the effects on C gain might diminish over time and that on plant water use would be amplified if N deposition persists.     

The effects of urbanization on pollinators and pollination: A meta-analysis

Urbanization is increasing worldwide, with major impacts on biodiversity, species interactions and ecosystem functioning. Pollination is an ecosystem function vital for terrestrial ecosystems and food security; however, the processes underlying the patterns of pollinator diversity and the ecosystem services they provide in cities have seldom been quantified. Here, we perform a comprehensive meta-analysis of 133 studies examining the effects of urbanization on pollinators and pollination. Our results confirm the widespread negative impacts of urbanization on pollinator richness and abundance, with Lepidoptera being the most affected group. Furthermore, pollinator responses were found to be trait-specific, with below-ground nesting and solitary Hymenoptera, and spring flyers more severely affected by urbanization. Meanwhile, cities promote non-native pollinators, which may exacerbate conservation risks to native species. Surprisingly, despite the negative effects of urbanization on pollinator diversity, pollination service measured as seed set is enhanced in non-tropical cities likely due to abundant generalists and managed pollinators therein. We emphasize that the richness of local flowering plants could mitigate the negative impacts of urbanization on pollinator diversity. Overall, the results demonstrate the varying magnitudes of multiple moderators on urban pollinators and pollination services and could help guide conservation actions for biodiversity and ecosystem function for a sustainable future.     

Responses of terrestrial ecosystems to temperature and precipitation change: a meta‐analysis of experimental manipulation

Global mean temperature is predicted to increase by 2–7 °C and precipitation to change across the globe by the end of this century. To quantify climate effects on ecosystem processes, a number of climate change experiments have been established around the world in various ecosystems. Despite these efforts, general responses of terrestrial ecosystems to changes in temperature and precipitation, and especially to their combined effects, remain unclear. We used meta‐analysis to synthesize ecosystem‐level responses to warming, altered precipitation, and their combination. We focused on plant growth and ecosystem carbon (C) balance, including biomass, net primary production (NPP), respiration, net ecosystem exchange (NEE), and ecosystem photosynthesis, synthesizing results from 85 studies. We found that experimental warming and increased precipitation generally stimulated plant growth and ecosystem C fluxes, whereas decreased precipitation had the opposite effects. For example, warming significantly stimulated total NPP, increased ecosystem photosynthesis, and ecosystem respiration. Experimentally reduced precipitation suppressed aboveground NPP (ANPP) and NEE, whereas supplemental precipitation enhanced ANPP and NEE. Plant productivity and ecosystem C fluxes generally showed higher sensitivities to increased precipitation than to decreased precipitation. Interactive effects of warming and altered precipitation tended to be smaller than expected from additive, single‐factor effects, though low statistical power limits the strength of these conclusions. New experiments with combined temperature and precipitation manipulations are needed to conclusively determine the importance of temperature–precipitation interactions on the C balance of terrestrial ecosystems under future climate conditions.     

Plant secondary metabolic responses to global climate change: A meta-analysis in medicinal and aromatic plants

Plant secondary metabolites (SMs) play crucial roles in plant-environment interactions and contribute greatly to human health. Global climate changes are expected to dramatically affect plant secondary metabolism, yet a systematic understanding of such influences is still lacking. Here, we employed medicinal and aromatic plants (MAAPs) as model plant taxa and performed a meta-analysis from 360 publications using 1828 paired observations to assess the responses of different SMs levels and the accompanying plant traits to elevated carbon dioxide (eCO₂), elevated temperature (eT), elevated nitrogen deposition (eN) and decreased precipitation (dP). The overall results showed that phenolic and terpenoid levels generally respond positively to eCO₂ but negatively to eN, while the total alkaloid concentration was increased remarkably by eN. By contrast, dP promotes the levels of all SMs, while eT exclusively exerts a positive influence on the levels of phenolic compounds. Further analysis highlighted the dependence of SM responses on different moderators such as plant functional types, climate change levels or exposure durations, mean annual temperature and mean annual precipitation. Moreover, plant phenolic and terpenoid responses to climate changes could be attributed to the variations of C/N ratio and total soluble sugar levels, while the trade-off supposition contributed to SM responses to climate changes other than eCO₂. Taken together, our results predicted the distinctive SM responses to diverse climate changes in MAAPs and allowed us to define potential moderators responsible for these variations. Further, linking SM responses to C-N metabolism and growth-defence balance provided biological understandings in terms of plant secondary metabolic regulation.     

Nitrogen availability mediates soil carbon cycling response to climate warming: A meta-analysis

Global climate warming may induce a positive feedback through increasing soil carbon (C) release to the atmosphere. Although warming can affect both C input to and output from soil, direct and convincing evidence illustrating that warming induces a net change in soil C is still lacking. We synthesized the results from field warming experiments at 165 sites across the globe and found that climate warming had no significant effect on soil C stock. On average, warming significantly increased root biomass and soil respiration, but warming effects on root biomass and soil respiration strongly depended on soil nitrogen (N) availability. Under high N availability (soil C:N ratio < 15), warming had no significant effect on root biomass, but promoted the coupling between effect sizes of root biomass and soil C stock. Under relative N limitation (soil C:N ratio > 15), warming significantly enhanced root biomass. However, the enhancement of root biomass did not induce a corresponding C accumulation in soil, possibly because warming promoted microbial CO₂ release that offset the increased root C input. Also, reactive N input alleviated warming-induced C loss from soil, but elevated atmospheric CO₂ or precipitation increase/reduction did not. Together, our findings indicate that the relative availability of soil C to N (i.e., soil C:N ratio) critically mediates warming effects on soil C dynamics, suggesting that its incorporation into C-climate models may improve the prediction of soil C cycling under future global warming scenarios.     

Biomass partitioning in response to resources availability: A comparison between native and invaded ranges in the clonal invader Carpobrotus edulis

Identifying the mechanism underlying plant invasiveness is a fast-moving research topic in current ecology. Phenotypic plasticity has been pointed out as a trait that can contribute to plant invasiveness. This experiment examines the presence of rapid adaptive evolution favoring plastic biomass partitioning during the invasion process. With that aim, we tested differences in patterns of biomass allocation between populations of Carpobrotus edulis from South Africa (native area) and the Iberian Peninsula (invaded area) growing under different nutrient, water and light availabilities in a common garden experiment. Here we demonstrate that biomass partitioning in response to nutrient availability in C. edulis differs between populations from native and invaded ranges, indicating that this trait could be under selection during the invasion process. Thus, nutrient shortage significantly increased the proportional production of roots in populations from the invaded range, but not in populations from the native area. This plastic root-foraging response may contribute to the optimization of nutrient uptake by plants, and therefore could be considered as an adaptive strategy. Understanding the ecological implications of rapid evolution for plastic biomass partitioning is important in determining processes of plant adaptation to new environments, and contributes to disentangling the mechanisms underlying plant invasiveness.     

中国亚热带不同菌根树种的根叶形态学性状特征与生长差异: 以江西新岗山为例

探究植物功能性状的种内和种间变异不仅有助于揭示植物对环境的适应, 也能够反映植物的生态策略, 但不同菌根类型树木生长过程中根叶形态学功能性状的适应策略仍有待探究。本研究依托中国亚热带森林生物多样性与生态系统功能实验研究平台(BEF-China)选取7种丛枝菌根(AM)树木和7种外生菌根(EM)树木的纯林, 测定各个树种的比叶面积、叶干物质含量、比根长、根系直径、树高生长速率、地径生长速率及细根生物量等根叶形态学功能性状和生长指标, 探讨了两种菌根类型树种间的根叶形态学特征的差异。结果表明: 与AM树种相比, EM树种具有较小的比叶面积、吸收根平均直径和生长速率, 但具有更大的叶干物质含量; 两种菌根树种之间的比根长和细根生物量无显著差异。比叶面积、叶干物质含量、树高生长速率、地径生长速率和细根生物量等功能性状及生长指标在不同菌根类型、树种及二者的交互作用中均存在显著差异; 且树种、根功能型、菌根类型及三者之间的交互作用均对根功能性状有显著影响。EM树种地上指标的种内变异均大于种间变异, 而AM树种地上指标的种内和种间变异程度类似; 但两种菌根树种细根生物量的种间变异均大于种内变异。尽管两种菌根树种地上部分生长速率较快通常表现为较低的叶干物质含量, 但AM树种通常拥有较高的吸收根比根长, 而EM树种拥有较粗的运输根平均直径。吸收根比根长越低, 两类菌根树种的细根生物量就越多。由此可见, 根叶功能性状对植物地上部分的生长具有一定的协同效应, 其中运输根主要在EM树种地上生长过程中发挥重要作用, 吸收根主要与AM树种的地上部分生长有关; 但两类菌根树种的地下细根生物量均与吸收根有关。     

Effects of plant traits on ecosystem and regional processes: a conceptual framework for predicting the consequences of global change

Human activities are causing widespread changes in the species composition of natural and managed ecosystems, but the consequences of these changes are poorly understood. This paper presents a conceptual framework for predicting the ecosystem and regional consequences of changes in plant species composition. Changes in species composition have greatest ecological effects when they modify the ecological factors that directly control (and respond to) ecosystem processes. These interactive controls include: functional types of organisms present in the ecosystem; soil resources used by organisms to grow and reproduce; modulators such as microclimate that influence the activity of organisms; disturbance regime; and human activities. Plant traits related to size and growth rate are particularly important because they determine the productive capacity of vegetation and the rates of decomposition and nitrogen mineralization. Because the same plant traits affect most key processes in the cycling of carbon and nutrients, changes in plant traits tend to affect most biogeochemical cycling processes in parallel. Plant traits also have landscape and regional effects through their effects on water and energy exchange and disturbance regime.     

PhenoSpace: A Shiny application to visualize trait data in the phenotypic space of the global spectrum of plant form and function

1. A recent analysis of variation in six major traits conducted on a large worldwide sample of vascular plant species showed that three‐quarters of trait variation was captured by a two‐dimensional global spectrum of plant form and function (“global spectrum” hereafter). We developed the PhenoSpace application, whose aim is to visualize and export the position of any individual/population/species in the phenotypic space of the global spectrum.
2. PhenoSpace is a Shiny application that helps users to manipulate and visualize data pertaining to the global spectrum of plant form and function. It is freely accessible at the following URL: https://shiny.cefe.cnrs.fr/PhenoSpace/.
3. PhenoSpace has three main functionalities. First, it allows users to visualize the phenotypic space of the global spectrum using different combinations of traits and growth forms. Second, trait data from any new user‐defined dataset can be projected onto the phenotypic space of the global spectrum, provided that at least two of the six traits are available. Finally, figures produced and loadings of the imported data on the PCA axes can be downloaded, allowing users to conduct further analyses.
4. PhenoSpace fulfills the practical goal of positioning plants in the phenotypic space of the global spectrum, making it possible to compare trait variation at any level of organization against the worldwide background. This serves a major aim of comparative plant ecology, which is to put specific sets of individuals, populations or species into a broader context, facilitating comparison and synthesis of results across different continents and environments using relevant indicators of plant design and function.

     

Seed size,number and strategies in annual plants: a comparative functional analysis and synthesis

Background and AimsPlants depend fundamentally on establishment from seed. However, protocols in trait-based ecology currently estimate seed size but not seed number. This can be rectified. For annuals, seed number should simply be a positive function of vegetative biomass and a negative function of seed size.MethodsUsing published values of comparative seed number as the ‘gold standard’ and a large functional database, comparative seed yield and number per plant and per m² were predicted by multiple regression. Subsequently, ecological variation in each was explored for English and Spanish habitats, newly calculated C-S-R strategies and changed abundance in the British flora.Key ResultsAs predicted, comparative seed mass yield per plant was consistently a positive function of plant size and competitive ability, and largely independent of seed size. Regressions estimating comparative seed number included, additionally, seed size as a negative function. Relationships differed numerically between regions, habitats and C-S-R strategies. Moreover, some species differed in life history over their geographical range. Comparative seed yield per m² was positively correlated with FAO crop yield, and increasing British annuals produced numerous seeds. Nevertheless, predicted values must be viewed as comparative rather than absolute: they varied according to the ‘gold standard’ predictor used. Moreover, regressions estimating comparative seed yield per m² achieved low precision.ConclusionsFor the first time, estimates of comparative seed yield and number for >800 annuals and their predictor equations have been produced and the ecological importance of these regenerative traits has been illustrated. ‘Regenerative trait-based ecology’ remains in its infancy, with work needed on determinate vs. indeterminate flowering (‘bet-hedging’), C-S-R methodologies, phylogeny, comparative seed yield per m² and changing life history. Nevertheless, this has been a positive start and readers are invited to use estimates for >800 annuals, in the Supplementary data, to help advance ‘regenerative trait-based ecology’ to the next level.