Variation in root system traits among African semi‐arid savanna grasses: Implications for drought tolerance

In arid to semi‐arid grasslands and savannas, plant growth, population dynamics, and productivity are consistently and strongly limited by soil water and nutrient availability. Adaptive traits of the root systems of grasses in these ecosystems are crucial to their ability to cope with strong water and/or nutrient limitation and the increasing drought stress associated with ecosystem degradation or projected climate change. We studied 18 grass species in semi‐arid savanna of the Kalahari region of Botswana to quantify interspecific variation in three important root system traits including root system architecture, rhizosheath thickness and mycorrhizal colonization. Drought‐tolerant species and shorter‐lived species showed greater rhizosheath thickness and fine root development but lower mycorrhizal colonization compared to later successional climax grasses and those characteristic of wetter sites. In addition, there was a significant positive correlation between root fibrousness index and rhizosheath thickness among species and a weak negative correlation between root fibrousness index and mycorrhizal colonization. These patterns suggest that an extensive fine root system and rhizosheath development may be important complementary traits of grasses coping with drought conditions, the former aiding in the acquisition of water by the grass plant and the latter aiding in water uptake and retention, and reducing water loss in the rhizosphere. Within species, both rhizosheath development and mycorrhizal colonization were significantly greater in a wet year than in a year with below‐average precipitation. The observed patterns suggest that the primary benefit of rhizosheath development in African savanna grasses is improved drought tolerance and that it is a plastic trait that can be adjusted annually to changing environmental conditions. The functioning of mycorrhizal symbiosis is likely to be relatively more important in infertile savannas where nutrient limitation is higher relative to water limitation.     

Contrasting effects of plant inter‐ and intraspecific variation on community trait responses to restoration of a sandy grassland ecosystem

Changes in plant community traits along an environmental gradient are caused by interspecific and intraspecific trait variation. However, little is known about the role of interspecific and intraspecific trait variation in plant community responses to the restoration of a sandy grassland ecosystem. We measured five functional traits of 34 species along a restoration gradient of sandy grassland (mobile dune, semi‐fixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China. We examined how community‐level traits varied with habitat changes and soil gradients using both abundance‐weighted and non‐weighted averages of trait values. We quantified the relative contribution of inter‐ and intraspecific trait variation in specific leaf area (SLA), leaf dry matter content (LDMC), leaf carbon content (LCC), leaf nitrogen content (LNC), and plant height to the community response to habitat changes in the restoration of sandy grassland. We found that five weighted community‐average traits varied significantly with habitat changes. Along the soil gradient in the restoration of sandy grassland, plant height, SLA, LDMC, and LCC increased, while LNC decreased. For all traits, there was a greater contribution of interspecific variation to community response in regard to habitat changes relative to that of intraspecific variation. The relative contribution of the interspecific variation effect of an abundance‐weighted trait was greater than that of a non‐weighted trait with regard to all traits except LDMC. A community‐level trait response to habitat changes was due largely to species turnover. Though the intraspecific shift plays a small role in community trait response to habitat changes, it has an effect on plant coexistence and the maintenance of herbaceous plants in sandy grassland habitats. The context dependency of positive and negative covariation between inter‐ and intraspecific variation further suggests that both effects of inter‐ and intraspecific variation on a community trait should be considered when understanding a plant community response to environmental changes in sandy grassland ecosystems.     

Linking root respiration to chemistry and morphology across species

Root respiration is a critical physiological trait involved in root resource acquisition strategies, yet it is less represented in root trait syndrome. Here we compiled a large dataset of root respiration associated with root chemical and morphological traits from 245 plant species. Our results demonstrated that root respiration correlated positively with root nitrogen concentration (RNC) and negatively with root tissue density (RTD) across and within woody and non‐woody species. However, the relationships between root respiration and specific root length (SRL) and root diameter (RD) were weak or even insignificant. Such root respiration–traits relationships were not completely in line with predictions by the root economics spectrum (RES). Furthermore, the principal component analysis showed that root trait syndrome was multidimensional. Root respiration was associated more strongly with the RNC‐RTD axis (the classical RES) than with the orthogonal SRL‐RD axis for woody species, but not for non‐woody species. Collectively, the linkages of root physiological, chemical, and morphological traits provide a better understanding of root trait covariation and root resource acquisition strategies.     

Root exudation rate as functional trait involved in plant nutrient‐use strategy classification

Plants adopt a variety of life history strategies to succeed in the Earth's diverse environments. Using functional traits which are defined as “morphological, biochemical, physiological, or phonological” characteristics measurable at the individual level, plants are classified according to their species’ adaptative strategies, more than their taxonomy, from fast growing plant species to slower‐growing conservative species. These different strategies probably influence the input and output of carbon (C)‐resources, from the assimilation of carbon by photosynthesis to its release in the rhizosphere soil via root exudation. However, while root exudation was known to mediate plant‐microbe interactions in the rhizosphere, it was not used as functional trait until recently. Here, we assess whether root exudate levels are useful plant functional traits in the classification of plant nutrient‐use strategies and classical trait syndromes? For this purpose, we conducted an experiment with six grass species representing along a gradient of plant resource‐use strategies, from conservative species, characterized by low biomass nitrogen (N) concentrations and a long lifespans, to exploitative species, characterized by high rates of photosynthesis and rapid rates of N acquisition. Leaf and root traits were measured for each grass and root exudate rate for each planted soil sample. Classical trait syndromes in plant ecology were found for leaf and root traits, with negative relationships observed between specific leaf area and leaf dry matter content or between specific root length and root dry matter content. However, a new root trait syndrome was also found with root exudation levels correlating with plant resource‐use strategy patterns, specifically, between root exudation rate and root dry matter content. We therefore propose root exudation rate can be used as a key functional trait in plant ecology studies and plant strategy classification.     

Long‐term farming systems and last crop sown shape the species and functional composition of the arable weed seed bank

Questions: The assembly of arable weed communities is the result of local filtering by agricultural management and crop competition. Therefore, soil seed banks can reflect the effects of long‐term cumulative field management and crop sequences on weed communities. Moreover, soil seed banks provide strong estimates of future weed problems but also of potential arable plant diversity and associated ecological functions. For this, we evaluated the effects of different long‐term farming systems under the same crop rotation sequence on the abundance, diversity and community assembly of weed seed bank, as well as on the functional diversity and composition. Location: DOK (biodynamic [D], bioorganic [O], conventional [K]) long‐term trial, Therwil, Switzerland. Methods: The effects of long‐term contrasted farming systems (i.e., biodynamic, organic, conventional, mineral and unfertilised systems) and last crop sown (i.e., wheat and maize) were evaluated on different indicators of species and functional diversity and composition of the weed soil seed bank. Results: The results showed significant influences of 40 years of contrasted farming systems on the diversity and composition of the seed bank, with higher diversities being found in unfertilised and organic farming systems, but also higher abundances than those found under conventional systems. Organic farming also allowed higher functional richness, dispersion and redundancy. Different farming systems triggered shifts in species and functional assemblies. Conclusions: The results highlight the importance of organic management for the maintenance of a diverse arable plant community and its functions. However, such results emphasise the need for appropriate yearly management to reduce the abundance of settled weediness and prevent affecting crop production. The farm management filtered community composition based on functional traits. Although the soil seed bank buffers the long‐term farming and crop sequence, the last crop sown and, thus, the yearly management were important determinants of seed bank composition.     

New insights into leaf and fine‐root trait relationships: implications of resource acquisition among 23 xerophytic woody species

Functional traits of leaves and fine root vary broadly among different species, but little is known about how these interspecific variations are coordinated between the two organs. This study aims to determine the interspecific relationships between corresponding leaf and fine‐root traits to better understand plant strategies of resource acquisition. SLA (Specific leaf area), SRL (specific root length), mass‐based N (nitrogen) and P (phosphorus) concentrations of leaves and fine roots, root system, and plant sizes were measured in 23 woody species grown together in a common garden setting. SLA and SRL exhibited a strong negative relationship. There were no significant relationships between corresponding leaf and fine‐root nutrient concentrations. The interspecific variations in plant height and biomass were tightly correlated with root system size characteristics, including root depth and total root length. These results demonstrate a coordinated plant size‐dependent variation between shoots and roots, but for efficiency, plant resource acquisition appears to be uncoupled between the leaves and fine roots. The different patterns of leaf and fine‐root traits suggest different strategies for resource acquisition between the two organs. This provides insights into the linkage between above‐ and belowground subsystems in carbon and nutrient economy.     

Phenotypic plasticity of the maize root system in response to heterogeneous nitrogen availability

Mineral nutrients are distributed in a non-uniform manner in the soil. Plasticity in root responses to the availability of mineral nutrients is believed to be important for optimizing nutrient acquisition. The response of root architecture to heterogeneous nutrient availability has been documented in various plant species, and the molecular mechanisms coordinating these responses have been investigated particularly in Arabidopsis, a model dicotyledonous plant. Recently, progress has been made in describing the phenotypic plasticity of root architecture in maize, a monocotyledonous crop. This article reviews aspects of phenotypic plasticity of maize root system architecture, with special emphasis on describing (1) the development of its complex root system; (2) phenotypic responses in root system architecture to heterogeneous N availability; (3) the importance of phenotypic plasticity for N acquisition; (4) different regulation of root growth and nutrients uptake by shoot; and (5) root traits in maize breeding. This knowledge will inform breeding strategies for root traits enabling more efficient acquisition of soil resources and synchronizing crop growth demand, root resource acquisition and fertilizer application during crop growing season, thereby maximizing crop yields and nutrient-use efficiency and minimizing environmental pollution.     

Fine root tradeoffs between nitrogen concentration and xylem vessel traits preclude unified whole‐plant resource strategies in Helianthus

Recent work suggests variation in plant growth strategies is governed by a tradeoff in resource acquisition and use, ranging from a rapid resource acquisition strategy to a resource‐conservative strategy. While evidence for this tradeoff has been found in leaves, knowledge of root trait strategies, and whether they reflect adaptive differentiation across environments, is limited. In the greenhouse, we investigated variation in fine root morphology (specific root length and tissue density), chemistry (nitrogen concentration and carbon:nitrogen), and anatomy (root cross‐sectional traits) in populations of 26 Helianthus species and sister Phoebanthus tenuifolius. We also compared root trait variation in this study with leaf trait variation previously reported in a parallel study of these populations. Root traits varied widely and exhibited little phylogenetic signal, suggesting high evolutionary lability. Specific root length and root tissue density were weakly negatively correlated, but neither was associated with root nitrogen, providing little support for a single axis of root trait covariation. Correlations between traits measured in the greenhouse and native site characteristics were generally weak, suggesting a variety of equally viable root trait combinations exist within and across environments. However, high root nitrogen was associated with lower xylem vessel number and cross‐sectional area, suggesting a tradeoff between nutrient investment and water transport capacity. This led to correlations between root and leaf traits that were not always consistent with an acquisition–conservation tradeoff at the whole‐plant level. Given that roots must balance acquisition of water and nutrients with functions like anchorage, exudation, and microbial symbioses, the varied evidence for root trait covariation likely reflects the complexity of interacting selection pressures belowground. Similarly, the lack of evidence for a single acquisition–conservation tradeoff at the whole‐plant level likely reflects the vastly different selection pressures shaping roots and leaves, and the resources they are optimized to obtain.     

Intraspecific trait variation can weaken interspecific trait correlations when assessing the whole‐plant economic spectrum

The worldwide plant economic spectrum hypothesis predicts that leaf, stem, and root traits are correlated across vascular plant species because carbon gain depends on leaves being adequately supplied with water and nutrients, and because construction of each organ involves a trade‐off between performance and persistence. Despite its logical and intuitive appeal, this hypothesis has received mixed empirical support. If traits within species diverge in their responses to an environmental gradient, then interspecific trait correlations could be weakened when measured in natural ecosystems. To test this prediction, we measured relative growth rates (RGR) and seven functional traits that have been shown to be related to fluxes of water, nutrients, and carbon across 56 functionally diverse tree species on (1) juveniles in a controlled environment, (2) juveniles in forest understories, and (3) mature trees in forests. Leaf, stem, and fine root traits of juveniles grown in a controlled environment were closely correlated with each other, and with RGR. Remarkably, the seven leaf, stem, and fine root tissue traits spanned a single dimension of variation when measured in the controlled environment. Forest‐grown juveniles expressed lower leaf mass per area, but higher wood and fine root tissue density, than greenhouse‐grown juveniles. Traits and growth rates were decoupled in forest‐grown juveniles and mature trees. Our results indicate that constraints exist on the covariation, not just the variation, among vegetative plant organs; however, divergent responses of traits within species to environmental gradients can mask interspecific trait correlations in natural environments. Correlations among organs and relationships between traits and RGR were strong when plants were compared in a standardized environment. Our results may reconcile the discrepancies seen among studies, by showing that if traits and growth rates of species are compared across varied environments, then the interorgan trait correlations observed in controlled conditions can weaken or disappear.     

Plasticity in above‐ and belowground resource acquisition traits in response to single and multiple environmental factors in three tree species

Functional trait plasticity is a major component of plant adjustment to environmental stresses. Here, we explore how multiple local environmental gradients in resources required by plants (light, water, and nutrients) and soil disturbance together influence the direction and amplitude of intraspecific changes in leaf and fine root traits that facilitate capture of these resources. We measured population‐level analogous above‐ and belowground traits related to resource acquisition, i.e. “specific leaf area”–“specific root length” (SLA–SRL), and leaf and root N, P, and dry matter content (DMC), on three dominant understory tree species with contrasting carbon and nutrient economics across 15 plots in a temperate forest influenced by burrowing seabirds. We observed similar responses of the three species to the same single environmental influences, but partially species‐specific responses to combinations of influences. The strength of intraspecific above‐ and belowground trait responses appeared unrelated to species resource acquisition strategy. Finally, most analogous leaf and root traits (SLA vs. SRL, and leaf versus root P and DMC) were controlled by contrasting environmental influences. The decoupled responses of above‐ and belowground traits to these multiple environmental factors together with partially species‐specific adjustments suggest complex responses of plant communities to environmental changes, and potentially contrasting feedbacks of plant traits with ecosystem properties. We demonstrate that despite the growing evidence for broadly consistent resource‐acquisition strategies at the whole plant level among species, plants also show partially decoupled, finely tuned strategies between above‐ and belowground parts at the intraspecific level in response to their environment. This decoupling within species suggests a need for many species‐centred ecological theories on how plants respond to their environments (e.g. competitive/stress‐tolerant/ruderal and response‐effect trait frameworks) to be adapted to account for distinct plant‐environment interactions among distinct individuals of the same species and parts of the same individual.     

Plant functional traits and soil microbial biomass in different vegetation zones on the Loess Plateau

Plant functional traits built the relationships between plant diversity, species composition, and physiology along with the environmental changes, thus influencing soil microbial community. As the sensitivity indicators, soil microbial biomass and plant functional traits responses soil micro-organism and plant characteristics in direct way. Ten plant functional traits of 149 species and soil microbial biomass (carbon, nitrogen, and phosphorus) were analyzed across the different vegetation types (forest, forest-steppe, and steppe) that are divided by environmental gradient (temperature and precipitation), aimed to find the correlations among them. Our results confirmed the greatest values of plant functional traits (except the leaf density and the fine root density) that were distributed in the steppe zone, mainly due to the different mean annual temperature and mean annual precipitation conditions. For different plant growth forms, the plant functional traits were significant differences among the vegetation zones. The advantages of higher rate nutrient cycling, plentiful biomass supplements, and favorite habit conditions lead to the forest-steppe zone with the highest C_mic and N_mic concentrations. The canonical correlation analysis indicated that leaf nitrogen, root nitrogen, and fine root densities were correlated with root exudate and tissue which affected the concentrations of soil organic carbon (SOC) and total nitrogen (N), consequently impacting soil microbial biomass carbon (C_mic) and soil microbial biomass nitrogen (N_mic). Soil is the medium that connects micro-organism and plant root system that influenced leaf nitrogen, root nitrogen, and fine root density of plant functional traits, the concentrations of SOC and total N that plant feedback are consequently influencing C_mic and N_mic.     

Impact of plant architecture versus leaf quality on attack by leaf-tying caterpillars on five oak species

Because shelter-building herbivorous insect species often consider structural features of their host plants in selecting construction sites, their probability of attack is likely to be a function of some combination of plant architectural traits and leaf quality factors. We tested the hypothesis that plant architecture, in the form of the number of touching leaves, influences interspecific variation in attack by leaf-tying caterpillars in five species of sympatric Missouri oaks (Quercus). We compared colonization on control branches, in which both architecture and leaf quality were potentially important, with colonization on experimental branches for which we controlled for the effects of architecture by creating equal numbers of artificial ties. Colonization of artificial ties was highly correlated with natural colonization on neighboring control branches, suggesting that leaf quality factors and not architecture influenced interspecific variation in attack by leaf-tying caterpillars. Of the leaf quality factors measured (water, protein-binding capacity, nitrogen, specific leaf area, pubescence, and toughness), nitrogen was the most explanatory. With the exception of white oak, natural leaf tie colonization was positively correlated with nitrogen availability (ratio of nitrogen to protein-binding capacity), and negatively correlated with protein-binding capacity of leaf extracts. Both host plant species and subgenus oak influenced the community composition of leaf-tying caterpillars and the non-tying symbionts colonizing the ties. Host plant differences in leaf nitrogen content were positively correlated with pupal weight of one of two caterpillar species reared on all five host plant species. Thus, interspecific differences in nitrogen, nitrogen availability, and protein-binding capacity of leaf extracts are the best predictors at this time of interspecific differences in attack by leaf-tying caterpillars, in turn affecting their success on individual host plants in the laboratory.     

Prediction of in situ root decomposition rates in an interspecific context from chemical and morphological traits

Background and Aims

We quantitatively relate in situ root decomposition rates of a wide range of trees and herbs used in agroforestry to root chemical and morphological traits in order to better describe carbon fluxes from roots to the soil carbon pool across a diverse group of plant species.

Methods

In situ root decomposition rates were measured over an entire year by an intact core method on ten tree and seven herb species typical of agroforestry systems and were quantified using decay constants (k values) from Olson''s single exponential model. Decay constants were related to root chemical (total carbon, nitrogen, soluble carbon, cellulose, hemicellulose, lignin) and morphological (specific root length, specific root length) traits. Traits were measured for both absorbing and non-absorbing roots.

Key Results

From 61 to 77 % of the variation in the different root traits and 63 % of that in root decomposition rates was interspecific. N was positively correlated, but total carbon and lignin were negatively correlated with k values. Initial root traits accounted for 75 % of the variation in interspecific decomposition rates using partial least squares regressions; partial slopes attributed to each trait were consistent with functional ecology expectations.

Conclusions

Easily measured initial root traits can be used to predict rates of root decomposition in soils in an interspecific context.     

Patterns in root traits of woody species hosting arbuscular and ectomycorrhizas: implications for the evolution of belowground strategies

Root traits vary enormously among plant species but we have little understanding of how this variation affects their functioning. Of central interest is how root traits are related to plant resource acquisition strategies from soil. We examined root traits of 33 woody species from northeastern US forests that form two of the most common types of mutualisms with fungi, arbuscular mycorrhizas (AM) and ectomycorrhizas (EM). We examined root trait distribution with respect to plant phylogeny, quantifying the phylogenetic signal (K statistic) in fine root morphology and architecture, and used phylogenetically independent contrasts (PICs) to test whether taxa forming different mycorrhizal associations had different root traits. We found a pattern of species forming roots with thinner diameters as species diversified across time. Given moderate phylogenetic signals (K = 0.44–0.68), we used PICs to examine traits variation among taxa forming AM or EM, revealing that hosts of AM were associated with lower branching intensity (r_PIC = −0.77) and thicker root diameter (r_PIC = −0.41). Because EM evolved relatively more recently and intermittently across plant phylogenies, significant differences in root traits and colonization between plants forming AM and EM imply linkages between the evolution of these biotic interactions and root traits and suggest a history of selection pressures, with trade-offs for supporting different types of associations. Finally, across plant hosts of both EM and AM, species with thinner root diameters and longer specific root length (SRL) had less colonization (r_PIC = 0.85, −0.87), suggesting constraints on colonization linked to the evolution of root morphology.     

Disentangling the relative importance of species occurrence,abundance and intraspecific variability in community assembly: a trait‐based approach at the whole‐plant level in Mediterranean forests

Understanding which factors and rules govern the process of assembly in communities constitutes one of the main challenges of plant community ecology. The presence of certain functional strategies along broad environmental gradients can help to understand the patterns observed in community assembly and the filtering mechanisms that take place. We used a trait‐based approach, quantifying variations in aboveground (leaf and stem) and belowground (root) functional traits along environmental gradients in Mediterranean forest communities (south Spain). We proposed a new practical method to quantify the relative importance of species turnover (distinguishing between species occurrence and abundance) versus intraspecific variation, which allowed us to better understand the assemblage rules of these plant communities along environmental gradients. Our results showed that the functional structure of the studied plant communities was highly determined by soil environment. Results from our modelling approach based on maximum likelihood estimators showed a predominant influence of soil water storage on most of the community functional traits. We found that changes in community functional structure along environmental gradients were mainly promoted by species turnover rather than by intraspecific variability. Specifically, our new method of variance decomposition demonstrated that between‐site trait variation was the result of changes in species occurrence rather than in the abundance of certain dominant species. In conclusion, this study showed that water availability promoted the predominance of specific trait values (both in above and belowground fractions) associated to a resource acquisition or conservation strategy. In addition, we provided evidence that changes on community functional structure along the environmental gradient were mainly promoted by a process of species replacement, which represent a crucial step towards a more general understanding of the relative importance of intraspecific versus interspecific trait variation in these woody Mediterranean communities.     

常绿阔叶林外生和丛枝菌根树种细根形态和构型性状对氮添加的可塑性响应

以中亚热带常绿阔叶林外生菌根树种罗浮栲和丛枝菌根树种木荷为研究对象,采用根袋法进行野外原位氮添加试验,研究了细根形态性状(比根长、比表面积、组织密度、平均根直径)和构型性状(分枝数、分枝比、根长增长速率、根尖密度、分枝密度),分析不同菌根树种细根形态和构型性状对氮沉降的响应。结果表明: 随序级增加,外生和丛枝菌根树种细根比根长、比表面积和分枝数对氮添加的塑性响应逐渐降低,组织密度则相反;这反映了不同分枝等级细根的养分获取与资源维持在序级间存在权衡关系。不同菌根树种对土壤氮有效性的变化采取不同的适应对策: 氮添加后,罗浮栲细根采取机会主义策略,依靠细根本身来提高养分吸收效率、增强空间扩展和就地养分吸收能力,以快速的养分吸收策略为主;而木荷通过养分吸收效率和根系构建成本之间的权衡,并未改变细根形态性状,更多地依赖于菌根菌和细根构型之间的互补性进行养分获取。外生和丛枝菌根树种维持和构建细根碳(C)成本的差异,导致细根采取最适合自身的养分捕获方式,以达到生存的最优策略。     

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.     

Intraspecific trait variation and allocation strategies of calcareous grassland species: Results from a restoration experiment

Intra- and interspecific trait variation express the response of plants dealing with different environmental conditions. We measured root and leaf traits on 14 species of calcareous grasslands in a restoration experiment. We aimed at identifying intraspecific differences in biomass allocation and functional plant traits under contrasting soil conditions by comparing plants growing in ancient grassland and two restored grasslands on ex-arable land, one of them with topsoil removal. Relative importance of trait variation within and among species, and among site was assessed by variance partitioning. Interspecific variation was more important than intraspecific variation, but the contribution of the latter to total variation was considerable, especially for specific leaf area. Changes in soil properties due to topsoil removal resulted in lower values of plant height, specific leaf area and specific root length compared to the control (ancient grassland). Soil fertility found in the treatment without top soil removal did not affect plant plasticity compared to the control. The study species showed two allocation strategies in relation to resource stress, while the responses of individual traits to the soil treatments were consistent across species. We conclude that caution must be taken when using mean trait values for plastic species or when working with environmental gradients.     

The ecological significance of plasticity in root weight ratio in response to nitrogen: Opinion

We analyzed data on root weight ratio from a range of experimental studies documenting plant allocation changes in response to altered nitrogen availability. Our goal was to determine the degree to which plasticity in allocation between roots and shoots exists and to search for patterns in such plasticity among species. Our survey included 77 studies representing 206 cases and 129 species. As expected, we found that root weight ratio decreased with increased nitrogen availability in the majority of cases examined, and this response was most consistent when plants were grown individually or in intraspecific competition (versus interspecific competition). Surprisingly, however, we found no evidence to support existing hypotheses that fast-growing species adapted to high soil fertilities exhibit the highest levels of morphological plasticity, or that plasticity is positively associated with competitive ability. Rather, we found that average amounts of plasticity in root weight ratio in response to nitrogen availability were similar among species grouped by maximum relative growth rate and habitat fertility. Similar results were obtained for species categorized by life form, life history or root weight ratio itself, and plasticity in root weight ratio also had no consistent relationship with competitive ability. Numerous difficulties are associated with the attempt to search for pattern using independent studies, however our results lead to the conclusion that strong patterns in plasticity of root weight ratio in response to nitrogen availability among species do not exist. We discuss two reasons for this: (1) the costs of plasticity relative to its benefits are lower than previously predicted and (2) plasticity in traits other than root weight ratio is more important to plant foraging ability.