Coordination between leaf and root traits in Mediterranean coastal dune plants

• Plant trait-based functional spectra are crucial to assess ecosystem functions and services. Whilst most research has focused on aboveground vegetative traits (leaf economic spectrum, LES), contrasting evidence on any coordination between the LES and root economic spectrum (RES) has been reported. Studying spectra variation along environmental gradients and accounting for species' phylogenetic relatedness may help to elucidate the strength of coordination between above- and belowground trait variation.
• We focused on leaf and root traits of 39 species sampled in three distinct habitats (front, back and slack) along a shoreline–inland gradient on coastal dunes. We tested, within a phylogenetic comparative framework, for the presence of the LES and RES, for any coordination between these spectra, and explored their relation to variation in ecological strategies along this gradient.
• In each habitat, three-quarters of trait variation is captured in two-dimensional spectra, with species' phylogenetic relatedness moderately influencing coordination and trade-off between traits. Along the shoreline–inland gradient, aboveground traits support the LES in all habitats. Belowground traits are consistent with the RES in the back-habitat only, where the environmental constraints are weaker, and a coordination between leaf and root traits was also found, supporting the whole-plant spectrum (PES).
• This study confirms the complexity when seeking any correlation between the LES and RES in ecosystems characterized by multiple environmental pressures, such as those investigated here. Changes in traits adopted to resist environmental constraints are similar among species, independent of their evolutionary relatedness, thus explaining the low phylogenetic contribution in support of our results.

     

人工控制条件下9种园林植物叶功能性状对短期NO2污染的响应

叶功能性状能反映植物对环境变化的适应策略,利用开顶式熏气法,对9种常见园林植物进行为期20天的NO₂熏气实验,分析不同NO₂浓度熏气下叶片形态结构指标（单叶干重、单叶面积、比叶面积）、光合生理指标（光合速率、荧光参数）及化学性状指标（叶N含量、叶P含量、N：P比值）的差异,从而探讨以上植物叶性状对NO₂污染的短期响应。结果表明,不同植物叶性状对不同浓度NO₂污染的响应存在显著差异,即不同植物应对NO₂污染的适应策略不同,同种植物在不同浓度NO₂熏气下叶性状指标变化趋势相同,但变化幅度不同,可见同种植物对不同NO₂浓度的响应策略也存在差异;大部分叶性状指标间表现出显著的相关性,但叶N含量与比叶面积间未发现显著相关性,表明叶经济谱性状间权衡机制的稳定性在个体尺度上可能会发生改变。研究结果揭示了不同植物对NO₂污染的适应与响应差异,对预测城市NO₂污染可能带来的植物功能性状的协同进化以及植物生态策略的改变具有重要意义。     

内蒙古荒漠草原植物叶片功能性状关系及其经济谱分析

气候变化和人为干扰导致草原荒漠化加剧, 引发了严重的环境问题。因此, 对荒漠草原植物与环境变化关系的研究愈加迫切, 分析比较荒漠草原不同功能型物种叶片经济谱具有重要意义。该研究通过测定内蒙古荒漠草原生态系统不同功能型植物叶片的光合及叶绿素荧光参数、比叶面积和叶片氮素含量, 验证了荒漠草原植物叶片经济谱的存在, 明确了各功能型植物叶片性状间的关系及其在叶片经济谱中的位置。荒漠草原不同功能型植物叶片性状差异明显, 草本植物的比叶面积(SLA)、单位质量叶氮含量(N_mass)分别是灌木的2.39倍和1.20倍; 一年生植物单位面积最大净光合速率(A_area)、SLA、光合氮利用效率(PNUE)分别是多年生植物的1.93倍、2.13倍和4.24倍; C₄植物的A_area、SLA、PNUE分别是C₃植物的2.25倍、1.73倍和3.61倍。除A_area与单位面积叶氮含量(N_area)、PSII的实际光化学效率(Φ_PSII)与SLA之间不存在显著相关关系外, 叶片性状间存在广泛的相关关系, 且均达到极显著水平。这验证了叶片经济谱在内蒙古荒漠草原植物中也同样存在。进一步分析表明, 一年生植物、草本植物、C₄植物叶片在叶片经济谱中位于靠近薄叶、光合能力强、寿命短的一端; 而多年生植物、灌木、C₃植物叶片靠近厚叶、光合能力弱、寿命长的一端。这说明荒漠草原中不同功能型植物可通过权衡其经济性状间的关系而采取不同的适应策略, 对于荒漠草原生态系统管理具有重要的理论指导意义。     

Effects of soil organisms on aboveground multitrophic interactions are consistent between plant genotypes mediating the interaction

Belowground communities can affect interactions between plants and aboveground insect communities. Such belowground–aboveground interactions are known to depend on the composition of belowground communities, as well as on the plant species that mediates these interactions. However, it is largely unknown whether the effect of belowground communities on aboveground plant–insect interactions also depends on genotypic variation within the plant species that mediates the interaction. To assess whether the outcome of belowground–aboveground interactions can be affected by plant genotype, we selected two white cabbage cultivars [Brassica oleracea L. var. capitata (Brassicaceae)]. From previous studies, it is known that these cultivars differ in their chemistry and belowground and aboveground multitrophic interactions. Belowground, we inoculated soils of the cultivars with either nematodes or microorganisms and included a sterilized soil as a control treatment. Aboveground, we quantified aphid [Brevicoryne brassicae (L.) (Hemiptera: Aphididae)] population development and parasitoid [Diaeretiella rapae (McIntosh) (Hymenoptera: Braconidae)] fitness parameters. The cultivar that sustained highest aphid numbers also had the best parasitoid performance. Soil treatment affected aphid population sizes: microorganisms increased aphid population growth. Soil treatments did not affect parasitoid performance. Cultivars differed in their amino acid concentration, leaf relative growth rate, and root, shoot, and phloem glucosinolate composition but showed similar responses of these traits to soil treatments. Consistent with this observation, no interactions were found between cultivar and soil treatment for aphid population growth or parasitoid performance. Overall, the aboveground community was more affected by cultivar, which was associated with glucosinolate profiles, than by soil community.     

The effect of altered sink-source relations on photosynthetic traits and matter transport during the phase of reproductive growth in the annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis>

Annual plants transport a large portion of carbohydrates and nitrogenous compounds from leaves to seeds during the phase of reproductive growth. This study aimed to clarify how reproductive growth affects photosynthetic traits in leaves and matter transport within the plant in the annual herb Chenopodium album L. Plants were grown in pots and either reproductive tissues or axillary leaves were removed at anthesis. Matter transport was evaluated as temporal changes in dry mass (as a substitute of carbohydrates) and nitrogen content of aboveground organs: leaves, axillary leaves, stems and reproductive tissues. Photosynthetic capacity (light-saturated photosynthetic rate under ambient CO₂ concentration), nitrogen, chlorophyll and soluble protein content were followed in the 20^th leaf that was mature at the start of the experiment. Removal of reproductive tissues resulted in accumulation of dry mass in leaves and axillary leaves, and accumulation of nitrogen in stem as nitrogen resorption from leaves and axillary leaves proceeded with time. Removal of axillary leaves proportionally reduced dry mass and nitrogen allocation to reproductive tissues, thus affecting the quantity but not quality of seeds. Removal treatments did not alter the time course of photosynthetic capacity, nitrogen, chlorophyll or soluble protein content during senescence in the 20^th leaf, but changed the photosynthetic capacity per unit of leaf nitrogen according to demand from reproductive tissues. Together, the results indicate that reproductive tissues affected carbon and nitrogen economy separately. The amount of carbon was adjusted in leaves through photosynthetic capacity and carbohydrate export from them, and the amount of nitrogen was adjusted by transport from stem to reproductive tissues. The plant’s ability to independently regulate carbon and nitrogen economy should be important in natural habitats where the plant carbon-nitrogen balance can easily be disturbed by external factors.     

Aboveground vs. Belowground Carbon Stocks in African Tropical Lowland Rainforest: Drivers and Implications

BackgroundAfrican tropical rainforests are one of the most important hotspots to look for changes in the upcoming decades when it comes to C storage and release. The focus of studying C dynamics in these systems lies traditionally on living aboveground biomass. Belowground soil organic carbon stocks have received little attention and estimates of the size, controls and distribution of soil organic carbon stocks are highly uncertain. In our study on lowland rainforest in the central Congo basin, we combine both an assessment of the aboveground C stock with an assessment of the belowground C stock and analyze the latter in terms of functional pools and controlling factors.Conclusions/SignificanceWe suggest nutrient limitation, especially potassium, as the driver for aboveground versus belowground C allocation. However, other drivers such as C turnover, tree functional traits or demographic considerations cannot be excluded. We argue that large and unaccounted variability in C stocks is to be expected in African tropical rain-forests. Currently, these differences in aboveground and belowground C stocks are not adequately verified and implemented mechanistically into Earth System Models. This will, hence, introduce additional uncertainty to models and predictions of the response of C storage of the Congo basin forest to climate change and its contribution to the terrestrial C budget.     

内蒙古高原湖滨湿地优势植物功能性状特征及其适应性

受人类活动和气候变化的影响,湖泊湖滨带退化速度显著加快。植物功能性状的方法可以量化植物特征,预测植物对外界环境干扰的反应,有助于理解退化湖滨带湿地植物应对环境变化所表现出的适应机制,对湖泊湖滨湿地生态系统植被的恢复与重建具有重要意义。在内蒙古高原典型湖泊湖滨湿地选取芦苇（Phragmites australis）、赖草（Leymus secalinus）、毛茛（Ranunculus japonicus）、鹅绒委陵菜（Potentilla anserina）、碱蓬（Suaeda glauca）、盐角草（Salicornia europaea）和拂子茅（Calamagrostis epigeios）7种优势植物的叶片和根系作为研究对象,对不同湿地植物的11种功能性状变化规律及其与环境因子的关系进行研究。旨在探究环境变化影响下湖滨带湿地植物的物种分布和功能性状的差异,以及湿地植物在不同湖滨带湿地生境下的适应策略。在评估植物功能性状差异基础上,采用环境矩阵连接性状矩阵（RLQ）结合第四角分析（Fourth-Corner）的方法分析环境因子对植物功能性状的影响。结果表明,内蒙古湖滨带湿地中7种优势植物为了适应不同的环境的影响,植物的功能性状均产生不同程度的种间与种内变异,在湖滨带湿地中植物的植株高度、叶片碳含量、叶片氮含量、叶片碳氮比、比根长、根组织密度、根氮含量对环境变化的响应比较敏感,土壤pH与叶片干物质含量呈显著负相关;土壤盐分与植株高度、叶片碳含量和叶碳氮比显著负相关,与叶片氮含量、根组织密度显著正相关;土壤的总氮含量与植株高度显著正相关,与比根长显著负相关;土壤碳氮比与植株高度和叶片碳含量显著负相关,与植物比根长显著正相关;土壤容重与根氮含量显著负相关。研究表明内蒙古高原湖滨带湿地植物的功能性状受环境的作用强烈,植物采取了不同的性状策略来适应环境。     

Trade-offs between plant leaf hydraulic and economic traits

Leaf is the most important organ for carbon-water coupling of a plant because it is the primary medium for photosynthesis. It also acts as the hydraulic bottleneck and safety valve against hydraulic catastrophic dysfunctions. The leaf economics spectrum, which reflects the balance between investments and returns of leaf economic traits, provides a useful framework for examining species strategies as shaped by their evolutionary history. Changes in leaf hydraulic traits will influence leaf economic traits as well as plant survival and growth. Exploring trade-offs between leaf hydraulic and economic traits is thus of significance for modeling carbon-water relations, understanding the mechanisms of water/carbon investments, and extending the leaf economic spectrum. In this review, we first examined the trade-offs between leaf hydraulic and economic traits. Specially, we analyzed the relationships between leaf hydraulic conductivity and hydraulic vulnerability, water potential at the turgor loss point, water capacitance, safety margin, and leaf morphological, structural and functional traits. We then discussed potential mechanisms regulating leaf hydraulic and economic traits from leaf morphology, anatomy, venation, and stomatal functions. Finally, we proposed future research to: (1) develop an integrated whole-plant economics spectrum, including carbon-nitrogen-water resources and root-stem-leaf hydraulic transport system that will help revealing ecophysiological mechanisms of plant structure-functional coupling, carbon sequestration and water use; (2) explore a generalized trade-offs among leaf hydraulic safety, hydraulic efficiency and carbon fixation efficiency to advance our understanding of the relationships between biophysical structure and physiological metabolism in plant leaf construction under drought stress; and (3) explore the carbon-water metabolic relationship and coupling of water transport and growth rate for the metabolic theory and predictions at community scale.     

植物叶片水力与经济性状权衡关系的研究进展

叶片既是植物光合产物形成的主要场所, 又是整株植物的水力瓶颈、应对灾难性水力失调的安全阀门, 是植物碳水耦合权衡的重要器官。叶经济型谱反映了叶片经济性状“投资-收益”的权衡, 为验证植物进化过程中形成的物种对策提供了适用的理论框架。叶片水力性状变化会影响叶片经济性状及植物存活和生长。因此, 探索植物叶片水力与经济性状的权衡关系, 对建立植物碳-水耦合模型、揭示植物水-碳投资机理、扩展植物性状型谱等均有重要意义。该文首先综述了叶片水力性状、经济性状及两者之间的权衡关系, 分析了叶片导水率与水力脆弱性、失膨点水势、水容、安全阈值等水力性状以及与叶片的形态、结构和气体交换功能性状之间的关系。然后, 从叶片形态、解剖和叶脉网络结构以及气孔功能方面探讨了叶片水力性状与经济性状的调节机制。最后, 提出今后应加强三方面的研究: (1)探索建立植物根-茎-叶水力输导系统的碳-氮-水资源的整株经济型谱, 以揭示植物功能结构耦合、高效固碳用水的生理生态学机制; (2)探索叶片水力安全、水力效率和固碳效率之间的普适性权衡关系, 以深入理解抗旱植物叶片构建的生物物理结构与生理代谢的关系; (3)探索个体水平碳水代谢关系、水分运输与生长速率的耦合, 为代谢推演理论和植物群落尺度预测提供基础。     

Leaf traits and performance vary with plant age and water availability in Artemisia californica

Background and AimsLeaf functional traits are strongly tied to growth strategies and ecological processes across species, but few efforts have linked intraspecific trait variation to performance across ontogenetic and environmental gradients. Plants are believed to shift towards more resource-conservative traits in stressful environments and as they age. However, uncertainty as to how intraspecific trait variation aligns with plant age and performance in the context of environmental variation may limit our ability to use traits to infer ecological processes at larger scales.MethodsWe measured leaf physiological and morphological traits, canopy volume and flowering effort for Artemisia californica (California sagebrush), a dominant shrub species in the coastal sage scrub community, under conditions of 50, 100 and 150 % ambient precipitation for 3 years.Key ResultsPlant age was a stronger driver of variation in traits and performance than water availability. Older plants demonstrated trait values consistent with a more conservative resource-use strategy, and trait values were less sensitive to drought. Several trait correlations were consistent across years and treatments; for example, plants with high photosynthetic rates tended to have high stomatal conductance, leaf nitrogen concentration and light-use efficiency. However, the trade-off between leaf construction and leaf nitrogen evident in older plants was absent for first-year plants. While few traits correlated with plant growth and flowering effort, we observed a positive correlation between leaf mass per area and performance in some groups of older plants.ConclusionsOverall, our results suggest that trait sensitivity to the environment is most visible during earlier stages of development, after which intraspecific trait variation and relationships may stabilize. While plant age plays a major role in intraspecific trait variation and sensitivity (and thus trait-based inferences), the direct influence of environment on growth and fecundity is just as critical to predicting plant performance in a changing environment.     

Deciduous and evergreen trees differ in juvenile biomass allometries because of differences in allocation to root storage

Background and Aims

Biomass partitioning for resource conservation might affect plant allometry, accounting for a substantial amount of unexplained variation in existing plant allometry models. One means of resource conservation is through direct allocation to storage in particular organs. In this study, storage allocation and biomass allometry of deciduous and evergreen tree species from seasonal environments were considered. It was expected that deciduous species would have greater allocation to storage in roots to support leaf regrowth in subsequent growing seasons, and consequently have lower scaling exponents for leaf to root and stem to root partitioning, than evergreen species. It was further expected that changes to root carbohydrate storage and biomass allometry under different soil nutrient supply conditions would be greater for deciduous species than for evergreen species.

Methods

Root carbohydrate storage and organ biomass allometries were compared for juveniles of 20 savanna tree species of different leaf habit (nine evergreen, 11 deciduous) grown in two nutrient treatments for periods of 5 and 20 weeks (total dry mass of individual plants ranged from 0·003 to 258·724 g).

Key Results

Deciduous species had greater root non-structural carbohydrate than evergreen species, and lower scaling exponents for leaf to root and stem to root partitioning than evergreen species. Across species, leaf to stem scaling was positively related, and stem to root scaling was negatively related to root carbohydrate concentration. Under lower nutrient supply, trees displayed increased partitioning to non-structural carbohydrate, and to roots and leaves over stems with increasing plant size, but this change did not differ between leaf habits.

Conclusions

Substantial unexplained variation in biomass allometry of woody species may be related to selection for resource conservation against environmental stresses, such as resource seasonality. Further differences in plant allometry could arise due to selection for different types of biomass allocation in response to different environmental stressors (e.g. fire vs. herbivory).     

Changes in plant taxonomic and functional diversity patterns following treeline advances in the South Urals

Background: Treeline ecotones represent environmental boundaries that fluctuate in space and time and thus induce changes in plant taxonomic and functional diversity.

Aims: To study changes through time in taxonomic and functional plant diversity patterns along the treeline ecotone.

Methods: In 2002, vegetation was sampled along a gradient from upper montane forest to the treeline–alpine transition in the South Ural Mountains, Russia. In 2014, vegetation was resampled and plant functional traits were collected. We studied spatial and temporal changes in plant species composition, functional composition and functional diversity.

Results: Species composition and diversity changed along the elevational gradient. The functional composition in height, leaf area, specific leaf area and leaf nitrogen content decreased with elevation, whereas functional composition of leaf carbon content increased. We found a temporal shift towards shorter plants with smaller leaves in treeline sites. Functional richness varied in several traits along the elevational gradient, while functional dispersion showed a trend towards increased functional dispersion in height, specific leaf area and leaf nitrogen in the treeline–tundra transition.

Conclusions: Tree encroachment across the treeline ecotone has resulted in a shift in plant species relative abundances and functional diversity, possibly affecting plant community assembly patterns.     

Shifts in fine root traits within and among species along a fine-scale hydrological gradient

Background and AimsLessons from above-ground trait ecology and resource economics theory may not be directly translatable to below-ground traits due to differences in function, trade-offs and environmental constraints. Here we examine root functional traits within and across species along a fine-scale hydrological gradient. We ask two related questions: (1) What is the relative magnitude of trait variation across the gradient for within- versus among-species variation? (2) Do correlations among below-ground plant traits conform with predictions from resource-economic spectrum theory?MethodsWe sampled four below-ground fine-root traits (specific root length, branching intensity, root tissue density and root dry matter content) and four above-ground traits (specific leaf area, leaf size, plant height and leaf dry matter content) in vascular plants along a fine-scale hydrological gradient within a wet heathland community in south-eastern Australia. Below-ground and above-ground traits were sampled both within and among species.Key ResultsRoot traits shifted both within and among species across the hydrological gradient. Within- and among-species patterns for root tissue density showed similar declines towards the wetter end of the gradient. Other root traits showed a variety of patterns with respect to within- and among-species variation. Filtering of species has a stronger effect compared with the average within-species shift: the slopes of the relationships between soil moisture and traits were steeper across species than slopes of within species. Between species, below-ground traits were only weakly linked to each other and to above-ground traits, but these weak links did in some cases correspond with predictions from economic theory.ConclusionsOne of the challenges of research on root traits has been considerable intraspecific variation. Here we show that part of intraspecific root trait variation is structured by a fine-scale hydrological gradient, and that the variation aligns with among-species trends in some cases. Patterns in root tissue density are especially intriguing and may play an important role in species and individual response to moisture conditions. Given the importance of roots in the uptake of resources, and in carbon and nutrient turnover, it is vital that we establish patterns of root trait variation across environmental gradients.     

典型草原围封后羊草地上功能性状对枯落物累积的响应

植物功能性状反映植物适应环境变化过程中在不同器官形态与功能间的资源权衡与分配策略。典型草原围封后枯落物累积导致群落光照、热量和水分的重新分配并改变微环境特征。在此过程中植物地上功能性状将通过怎样的变化来适应新的环境,目前尚不清楚。2015—2017年每年8月对内蒙古地区3种典型草原共有物种羊草（Leymus chinensis）的植株、叶片和茎干功能性状进行了测量与分析。结果表明：枯落物累积显著增加了羊草的植株高度、单株重量、茎叶比和总叶面积;枯落物累积显著增加了羊草的叶片长度、叶片重量、单叶面积、节间长度和茎干重量,这些性状属于敏感性状;枯落物累积对羊草的叶片数量和节间数量无显著影响,相对而言,这些性状属于惰性性状;羊草的单株重量与植株高度、叶片重量呈极显著的正相关关系（P<0.0001）;羊草的植株高度与节间数量、节间长度呈极显著的正相关关系（P<0.0001）。本研究结果从植物地上功能性状的角度阐明了典型草原植物对环境变化的适应方式,可为围封草原的合理管理提供基础数据与理论依据。     

寄主植物对桑寄生植物叶片功能性状的影响

为了解寄生植物叶片功能性状的差异及其影响因素,研究了西双版纳地区寄主植物对3种桑寄生植物叶片功能性状的影响,并分析了桑寄生植物与寄主植物叶片功能性状的相关性。结果表明,不同寄主植物的相同寄生植物叶片功能性状存在显著差异,来自7种寄主植物的五蕊寄生(Dendrophthoe pentandra)的叶片含水量(61.2%～70.1%)、氮含量(9.6～16.0 g/kg)、碳氮比(30.8～48.5)以及缩合单宁含量(3.3%～11.0%)等性状的差异较大;从4种寄主植物上获取的澜沧江寄生(Scurrula chingii var.yunnanensis)的叶片含水量(60.0%～71.7%)、碳含量(431.3～502.3 g/kg)和缩合单宁含量(3.8%～9.9%)等性状也呈现较大种间差异,而在2种寄主植物上的离瓣寄生(Helixanthera parasitica)的叶片功能性状没有显著差异。桑寄生植物与寄主植物的叶片含水量、总碳含量、总氮含量、碳氮比和缩合单宁含量呈显著的正相关。寄主植物作为桑寄生植物营养物质的主要来源,会影响桑寄生植物叶片的相应功能性状。桑寄生植物能从寄主植物获...     

Halving sunlight reveals no carbon limitation of aboveground biomass production in alpine grassland

In temperate alpine environments, the short growing season, low temperature and a slow nutrient cycle may restrict plant growth more than carbon (C) assimilation does. To test whether C is a limiting resource, we applied a shade gradient from ambient light to 44% (maximum shade) of incident photon flux density (PFD) in late successional, Carex curvula‐dominated alpine grassland at 2,580 m elevation in the Swiss central Alps for 3 years (2014–2016). Total aboveground biomass did not significantly decrease under reduced PFD, with a confidence interval ranging from +4% to ?15% biomass in maximum shade. Belowground biomass, of which more than 80% were fine roots, was significantly reduced by a mean of 17.9 ± 4.6% (±SE), corresponding to 228 g/m², in maximum shade in 2015 and 2016. This suggests reduced investments into water and nutrient acquisition according to the functional equilibrium concept. Specific leaf area (SLA) and maximum leaf length of the most abundant species increased with decreasing PFD. Foliar concentration of nonstructural carbohydrates (NSC) was reduced by 12.5 ± 4.3% under maximum shade (mean of eight tested species), while NSC concentration of belowground storage organs were unchanged in the four most abundant forbs. Furthermore, maximum shade lowered foliar δ¹³C by 1.56 ± 0.35‰ and increased foliar nitrogen concentrations per unit dry mass by 18.8 ± 4.1% across six species in 2015. However, based on unit leaf area, N concentrations were lower in shade (effect of higher SLA). Thus, while we found typical morphological and physiological plant responses to lower light, shading did not considerably affect seasonal aboveground biomass production of this alpine plant community within a broad range of PFD. This suggests that C is not a growth‐limiting resource, matching the unresponsiveness to in situ CO₂ enrichment previously reported for this type of grassland.     

Effects of elevated CO2 on growth and carbon/nutrient balance in the deciduous woody shrub Lindera benzoin (L.) Blume (Lauraceae)

We examined the effects of elevated CO₂ on growth and carbon/nutrient balance in a natural population of the deciduous temperate zone shrub Lindera benzoin. Our data concern whole plant, leaf, and stem growth for the first two seasons of a long-term field experiment in which CO₂ levels were manipulated in situ. In addition to growth parameters, we evaluated changes in leaf and stem chemistry, including total nitrogen, nonstructural carbohydrates, and total phenolics. Over the course of this study, L. benzoin appeared to respond to elevated CO₂ primarily by physiological and biochemical changes, with only a slight enhancement in aboveground growth (ramet height). Positive effects on aboveground growth were primarily evident in young (nonreproductive) ramets. Our results suggest that nitrogen limitation may have constrained plants to allocate carbohydrates produced in response to elevated CO₂ primarily to storage and belowground growth, and perhaps to increased secondary chemical production, rather than to increased stem and leaf growth. We discuss our results in terms of changes in carbon/nutrient balance induced by elevated CO₂, and provide predictions for future changes in this system based upon constraints imposed by intrinsic and extrinsic factors and their potential effects on the reallocation of stored reserves.     

Analysis of relationships among leaf functional traits and economics spectrum of plant species in the desert steppe of Nei Mongol

Aims Grassland desertification is being accelerated because of adverse climate change effects and unsustainable land uses, resulting in several major environmental problems. However, there are few studies on the economics spectrum of different plant functional types in desert steppe. The objectives of the current study are to examine the relationships among leaf functional traits of native plant species, to compare the functional traits among different plant functional types, and to determine whether an economic spectrum exists for the majority of species in the desert steppe of Damao Banner, Nei Mongol, China.
Methods Photosynthetic and chlorophyll fluorescence parameters, specific leaf area (SLA), and leaf nitrogen contents across 24 species of different functional types were measured in situ in the desert steppe ecosystem. Non-parametric tests were used to analyze leaf trait differences in plant species of different functional types. Linear regression analysis was used to determine the relationships among leaf traits in different plant species. Finally, a comprehensive analysis on these leaf traits in different plant species was conducted using the principal component analysis. All data analyses were performed using SPSS 16.0 (SPSS, Chicago, USA).
Important findings Significant differences among plant functional types were found in most of the leaf traits. SLA and mass-based nitrogen concentration (N_mass) in grasses were 2.39 and 1.20 folds, respectively, of that in shrubs; area-based photosynthetic capacity (A_area), SLA, and photosynthetic nitrogen use efficiency (PNUE) in annual species were 1.93, 2.13, and 4.24 folds, respectively, of that in perennial species; and A_area, SLA, and PNUE in C₄ species were 2.25, 1.73, and 3.61 folds, respectively, of that in C₃ species. Almost all relationships significantly differed (p < 0.01) among the leaf traits, with exception of the relationships between A_area and area-based nitrogen concentration (N_area) and between quantum yield of PSII electron transport (Φ_PSII) and SLA, implying that an economic spectrum may exist in the desert steppe ecosystem. The relationships of N_area, mass-based photosynthetic capacity (A_mass), and PNUE with SLA were most significantly strong (R² = 0.54, 0.62, 0.60, respectively; p < 0.01). Results in this study suggest that the annuals, grasses, and C₄ species might be located at the end of the leaf economic spectrum with high area-based photosynthetic rate, high nitrogen concentration on mass basis, short leaf lifespan, and high SLA; whereas the perennials, shrubs, and C₃ species could be located at the another end of the economic spectrum with contrasting traits.     

Co-ordination between xylem anatomy,plant architecture and leaf functional traits in response to abiotic and biotic drivers in a nurse cushion plant

Background and AimsPlants in dry Mediterranean mountains experience a double climatic stress: at low elevations, high temperatures coincide with water shortage during summer, while at high elevations temperature decreases and water availability increases. Cushion plants often act as nurses by improving the microclimate underneath their canopies, hosting beneficiary species that may reciprocally modify their benefactors’ microenvironment. We assess how the nurse cushion plant Arenaria tetraquetra subsp. amabilis adjusts its hydraulic system to face these complex abiotic and biotic constraints.MethodsWe evaluated intra-specific variation and co-ordination of stem xylem anatomy, leaf functional traits and plant architecture in response to elevation, aspect and the presence of beneficiary species in four A. tetraquetra subsp. amabilis populations in the Sierra Nevada mountains, southern Spain.Key ResultsXylem anatomical and plant architectural traits were the most responsive to environmental conditions, showing the highest mutual co-ordination. Cushions were more compact and had smaller, more isolated conductive vessels in the southern than in the northern aspect, which allow minimization of the negative impacts of more intense drought. Only vessel size, leaf mass per area and terminal branch length varied with elevation. Nurse cushions co-ordinated plant architecture and xylem traits, having higher canopy compactness, fewer leaves per branch and fewer, more isolated vessels than non-nurse cushions, which reflects the negative effects of beneficiary plants on nurse water status. In non-nurse cushions, plant architecture co-ordinated with leaf traits instead. The interacting effects of aspect and elevation on xylem traits showed that stress due to frost at high elevation constrained xylem anatomy in the north, whereas stress due to drought had a parallel effect in the south.ConclusionsTrait co-ordination was weaker under more demanding environmental conditions, which agrees with the hypothesis that trait independence allows plants to better optimize different functions, probably entailing higher adjustment potential against future environmental changes.     

环境及遗传背景对延河流域植物叶片和细根功能性状变异的影响

研究环境筛选作用和植物系统发育背景对植物群落构建产生的影响,有助于理解植物在生长过程中对资源的分配利用和对环境的适应规律。以延河流域3个植被带(森林带、森林草原带及典型草原带)稳定的自然植物群落为研究对象,调查了31个样地107种植物,隶属于35个科78个属,测量了6种叶片和3种细根性状。分别对3个植被带和不同植物科植物的叶片和细根性状做单因素方差分析,结果表明:叶片氮含量和细根氮含量在3个植被带间无显著差异,叶厚度、比叶面积、叶组织密度、叶片磷含量、叶片氮磷比、比根长、根组织密度在3个植被带间差异极显著。由南向北随着气候干旱的加剧,植物通过调节叶片和细根性状,表现出了不同的适应策略:森林带植物叶片相对生长速率高,根系防御力强;森林草原带植物叶片防御力强,根系相对生长速率快。不同科的植物在相同的环境条件下,对于资源的竞争力和胁迫的忍耐力也有所不同,比如豆科植物具有远远高于其他科的叶片和细根氮含量,但是对养分的利用效率并不高。GLM分析结果说明,所涉及的植物功能性状的空间变异主要来自于年均降雨量的变化及植物科的差异,如16.26%的比叶面积的变异可由年均降雨量变化解释,4.02%可由植物科的差异解释。物种水平上,叶厚度、比叶面积、叶组织密度、比根长、根组织密度、叶片磷含量是对气候干燥度变化响应敏感的植物功能性状,其空间变异主要由环境差异所致。延河流域的植物群落在形成过程中,存在明显的环境筛选效应。这表明,环境异质性在植被恢复实践中必须予以考虑。