Which root traits determine nitrogen uptake by alpine plant species on the Tibetan Plateau?

Background and aims

Nitrogen (N) is one of the most important limiting factors influencing plant growth and reproduction in alpine and tundra ecosystems. However, in situ observations of the effects of root traits on N absorption by alpine plant species are still lacking.

Methods

We investigated the rates of N uptake and the effect of root characteristics in ten common herbaceous alpine plant species using a ¹⁵N isotope tracer technique and the root systems of plants growing in a semi-arid steppe environment on the Tibetan Plateau. Our objective was to determine the root traits (root biomass, volume, surface area, average diameter, length, specific root length and specific root area) that make the largest contribution to the total uptake of N (¹⁵N–NO₃ ^?, ¹⁵N–NH₄ ⁺ or ¹⁵N–glycine) by alpine plant species.

Results

Monocotyledonous species had higher absorption rates for ¹⁵N–NH₄ ⁺, ¹⁵N–NO₃ ^?, ¹⁵N–glycine and total ¹⁵N than dicotyledonous species (P < 0.05). The root biomass, volume, surface area and average diameter were negatively correlated with the absorption capacity for ¹⁵N–NH₄ ⁺, ¹⁵N–NO₃ ^? and total ¹⁵N across the ten alpine plant species. However, the specific root length and the specific root area had significantly positive effects on the uptake of N.

Conclusions

In contrast with traditional views on the uptake of N, the N uptake rate was not improved by a larger root volume or root surface area for these alpine plant species in a high-altitude ecosystem. Root morphological traits had greater impacts on N absorption than traits related to the root system size in alpine herbaceous plants.

     

Root morphology and its contribution to a large root system for phosphorus uptake by <Emphasis Type="Italic">Rytidosperma</Emphasis> species (wallaby grass)

Aims

Rytidosperma species are native Australian grasses which have different growth rates and phosphorus (P) requirements. This study examined the role of root morphology traits in response to P supply.

Methods

Nine Rytidosperma species ranging from slow- to fast-growth were examined along with Lolium perenne and Bromus hordeaceus. Plants were grown in a glasshouse for 47 days in soil supplied with six levels of P between 0 and 60 mg P per pot. Root mass, length and diameter, root hair length and density, and extent of mycorrhizal colonisation were measured.

Results

Across all species there was a positive correlation (P < 0.001) between P uptake and root mass, length and root hair cylinder volume (RHCV; estimated using root diameter, root hair length and root length) at all levels of P supply. An exception was the RHCV of B. hordeaceus, where expected P uptake was not achieved due to a markedly reduced root length at low-P supply. For the Rytidosperma species, morphological plasticity for specific root length, root mass fraction and root hair length ranged from 1.5-fold to 2.7-fold between high- and low-P supply. However, across all species and P levels no single root morphological trait was identified for universally increasing the size of the root system and P uptake.

Conclusions

Fast-growing species took up more P as a result of an overall larger root mass, greater root length and larger RHCV.

     

Threshold effect of the groundwater depth on the photosynthetic efficiency of <Emphasis Type="Italic">Tamarix chinensis</Emphasis> in the Yellow River Delta

Background and aims

Functional traits may underlie differences in niches, which promote plant species co-existence, but also differences in competitive ability, which drive competitive exclusion. Empirical evidence concerning the contribution of different traits to niche differentiation and the ability to supress and tolerate competitors is very limited, particularly when considering belowground interactions.

Methods

We grew 26 temperate grassland species along a density gradient of interspecific competitors to determine which belowground traits a) explain species’ ability to suppress and tolerate neighbours and b) contribute to niche differentiation, such that species with dissimilar trait values experience reduced competition.

Results

We found that having larger root systems with extensive horizontal spread and lower root tissue density enabled efficient suppression of neighbours but did not significantly contribute to the ability to tolerate competition. Species with deeper root systems, lower specific root length and less branched roots were better at tolerating competition, but these traits did not significantly affect the ability to suppress neighbours. None of the measured traits contributed significantly to niche differentiation, either individually or in combination.

Conclusions

This study provides little support for belowground traits contributing to species co-existence through niche differentiation. Instead, different sets of weakly correlated traits enable plants to either suppress or tolerate their competitors.

     

archiDART: an R package for the automated computation of plant root architectural traits

Background and aims

In order to analyse root system architectures (RSAs) from captured images, a variety of manual (e.g. Data Analysis of Root Tracings, DART), semi-automated and fully automated software packages have been developed. These tools offer complementary approaches to study RSAs and the use of the Root System Markup Language (RSML) to store RSA data makes the comparison of measurements obtained with different (semi-) automated root imaging platforms easier. The throughput of the data analysis process using exported RSA data, however, should benefit greatly from batch analysis in a generic data analysis environment (R software).

Methods

We developed an R package (archiDART) with five functions. It computes global RSA traits, root growth rates, root growth directions and trajectories, and lateral root distribution from DART-generated and/or RSML files. It also has specific plotting functions designed to visualise the dynamics of root system growth.

Results

The results demonstrated the ability of the package’s functions to compute relevant traits for three contrasted RSAs (Brachypodium distachyon [L.] P. Beauv., Hevea brasiliensis Müll. Arg. and Solanum lycopersicum L.).

Conclusions

This work extends the DART software package and other image analysis tools supporting the RSML format, enabling users to easily calculate a number of RSA traits in a generic data analysis environment.

     

Cultivar differences in spatial root distribution during early growth in soil,and its relation to nutrient uptake - a study of wheat,onion and lettuce

Background and aims

A study was made to quantify early root development, soil exploitation and nutrient uptake in spring wheat, onion and lettuce, and their variation among cultivars. The goal was to study genetic variation in root traits making cultivars better adapted to organic production systems or other low-input systems.

Methods

Six cultivars of each species were grown in transparent tubes to allow direct observation of early root growth. The tubes were 0.3 m deep, and 0.24 m in diameter. By placing the plants close to the edge rather than at the centre of the tubes, we could quantify the spatial distribution of the root systems as well as the general root growth and nutrient uptake.

Results

Root growth of wheat and lettuce was faster than root growth of onion, and onion showed little capacity for horizontal root system development. Significant variation in early root growth and horizontal spread of the root system was found among cultivars of all three species. In general, cultivars with strong growth and high volume of soil exploitation showed higher average nutrient concentrations.

Conclusion

Early shoot growth, root growth and nutrient uptake are intrinsically linked, making it difficult to determine whether improved root growth was the primary cause of improved performance. However, we did find cultivars where the strong root growth and superior root distribution seemed to be the driver for improved overall growth.

     

<Emphasis Type="Italic">Trichoderma</Emphasis>-enriched organic fertilizer can mitigate microbiome degeneration of monocropped soil to maintain better plant growth

Background and aims

Given the worldwide effort to improve the nitrogen (N) economy of crops, it is critical to understand the mechanisms of improved N uptake which have resulted from selection pressure for grain yield in Australian wheat (Triticum aestivum L.). Changes in root system traits and N uptake were examined in nine Australian wheat varieties released between 1958 and 2007.

Methods

Wheat varieties were grown in rhizo-boxes in a glasshouse. We measured nitrogen uptake and mapped root growth and proliferation to quantify root length density (RLD), root length per plant, root biomass, specific root length, and plant nitrogen uptake per unit root length.

Results

Selection for yield reduced total RLD and total root length, and increased N uptake per unit root length that overrode the reduction in root system size, effectively explaining the increase in N uptake. Importantly, N uptake in our experiment under controlled conditions matched field measurements, reinforcing the agronomic significance of the present study.

Conclusions

Wheat varieties released in Australia between 1958 and 2007 increased their N uptake, not because of increasing their root length and RLD, but for progressively increasing the efficiency of their root system in capturing N. Our collection of varieties is therefore an interesting model to probe for variation in the affinity of the root system for nitrate.

     

Root morphology acclimation to phosphorus supply by six cultivars of <Emphasis Type="Italic">Trifolium subterraneum</Emphasis> L

Aims

Trifolium subterraneum L. is the predominant annual pasture legume in southern Australia. Cultivars with improved phosphorus (P) foraging ability would improve the P-use efficiency of agricultural systems. We therefore investigated variation in root traits related to P-uptake among six cultivars.

Methods

Micro-swards were grown at six levels of P in field soil with indigenous arbuscular mycorrhizal (AM) fungi for six weeks. Dry matter yield, tissue P concentration, rhizosphere carboxylates, AM fungal colonisation and root morphological traits were measured.

Results

The cultivars showed similar shoot and root yield responses to P supply. Average root diameter did not change, specific root length (SRL) increased and root tissue density (RTD) decreased with increased P supply. Amounts of total rhizosphere carboxylates were low (<1.2 nmol cm^?1 root). The percentage of root length colonised by AM fungi was greatest (29–43 %) at an intermediate level (8 mg kg^?1 dry soil) of P supply.

Conclusions

Most differences among cultivars were reasonably consistent across P supply levels, indicating greater numbers of lines could be screened reliably at a single P level. Low colonisation by AM fungi at low P supply deserves consideration when selecting soils for cultivar comparisons. Increased SRL and decreased RTD at high P supply likely result from self-shading within the micro-swards and warrant further investigation.

     

Trapping of lead (Pb) by corn and pea root border cells

Aims

Most plants produce a root tip extracellular matrix that includes viable border cell populations programmed to disperse into soil. Like neutrophils, border cells export structures that trap pathogens and prevent root tip infection. Border cells also trap metals. The goal of this study was to determine if border cells trap Pb.

Methods

Border cell responses to Pb were observed microscopically. Border cell impact on Pb-induced injury to roots was assessed using root growth assays. Pb removal from solution was measured using inductively coupled plasma mass spectrometry (ICP-MS). Speciation of Pb associated with border cells was evaluated by synchrotron X-ray absorption spectroscopy (XAS).

Results

Increased border cell trap size and number occurred within minutes in response to Pb but not silicon (Si). Transient immersion of root tips into Pb after border cells were removed resulted in growth inhibition. Immersion of root tips and border cells into Pb solution resulted in significant removal of Pb. Si levels in the presence of root tips remained unchanged. The Pb speciation, measured with Pb L_III XAS, altered when reacted with border cells, indicating that direct binding by extracellular traps occurred.

Conclusions

Border cells can trap Pb and prevent damage to the root tip.

     

Comparative effect of inorganic N on plant growth and N<Subscript>2</Subscript> fixation of ten legume crops: towards a better understanding of the differential response among species

Aims

A better understanding of how plant growth, N nutrition and symbiotic nitrogen fixation (SNF) are influenced by soil inorganic N availability, for a wide range of legume species, is crucial to optimise legume productivity, N₂ fixation, while limiting environmental risks such as N leaching.

Methods

A comparative analysis was performed for ten legume crops, grown in a field experiment and supplied with four N fertiliser rates. Dry matter, N concentration and SNF were measured. In parallel, root elongation rates were studied in a greenhouse experiment.

Results

For most species, N fertilisation had little effect on plant growth and N accumulation. SNF was reduced by soil inorganic N available at sowing but with large differences in the magnitude of the response among species. The response varied according to plant N requirements for growth and plant ability to retrieve inorganic N. Accordingly, root lateral expansion rate measured in RhizoTubes was highly correlated with plant ability to retrieve inorganic N measured in the field experiment.

Conclusion

Combining SNF response to soil inorganic N, shoot N and plant ability to retrieve inorganic N, allowed a robust evaluation of differential response to soil inorganic N among a wide range of legume species.

     

<Emphasis Type="Italic">Sedum</Emphasis> root foraging in layered green roof substrates

Background and aims

Layered profiles of designed soils may provide long-term benefits for green roofs, provided the vegetation can exploit resources in the different layers. We aimed to quantify Sedum root foraging for water and nutrients in designed soils of different texture and layering.

Methods

In a controlled pot experiment we quantified the root foraging ability of the species Sedum album (L.) and S. rupestre (L.) in response to substrate structure (fine, coarse, layered or mixed), vertical fertiliser placement (top or bottom half of pot) and watering (5, 10 or 20 mm week^?1).

Results

Water availability was the main driver of plant growth, followed by substrate structure, while fertiliser placement only had marginal effects on plant growth. Root foraging ability was low to moderate, as also reflected in the low proportion of biomass allocated to roots (5–13%). Increased watering reduced the proportion of root length and root biomass in deeper layers.

Conclusions

Both S. album and S. rupestre had a low ability to exploit water and nutrients by precise root foraging in substrates of different texture and layering. Allocation of biomass to roots was low and showed limited flexibility even under water-deficient conditions.

     

Modelling facilitation or competition within a root system: importance of the overlap of root depletion and accumulation zones

Background and aims

Relevant soil properties and nutrient distributions influencing crop root growth might be different under no-till (NT) and mouldboard plough (MP) management. The possible different root systems within different managements might have key impact on crop nutrient uptake and consequently crop production. Our objective was to assess the long-term combined effects of tillage and phosphorus (P) fertilization on corn (Zea mays L.) root distribution and morphology.

Methods

Corn root and soil samples were collected during the silking stage at five depths (0–5, 5–10, 10–20, 20–30 and 30–40 cm) and three horizontal distances perpendicular to the corn row (5, 15 and 25 cm) under MP and NT with three P fertilizations (0, 17.5, and 35 kg P ha^?1) for a long-term (22 years) experiment in eastern Canada. Root morphology and soil properties were determined.

Results

NT practice decreased corn root biomass by ?26 % compared to MP, mainly by decreasing the primary and secondary roots. Additionally, corn roots in NT tend to be more expansive on the surface layer with higher root length and surface densities for the depth of 0–5 cm at two sampling distances of 15 and 25 cm. The 35 kg P ha^?1 rate increased the root biomass by 26 and 41 % compared to the 0 and 17.5 kg P ha^?1 rates.

Conclusions

No-tillage practice and low rates of P fertilization reduce corn roots. This is probably caused by the weed competition in NT and the continued downward P status with low P rates over 22 years.

     

Effects of <Emphasis Type="Italic">Penicillium bilaii</Emphasis> on maize growth are mediated by available phosphorus

Background and aims

Functional traits are promising indicators of global changes and ecosystem processes. Trait responses to environmental conditions have been examined widely in vascular plants. In contrast, few studies have focused on soil lichens and mosses composing biocrusts. We aimed to evaluate the potential of biocrust tissue traits as indicators of changes in climate and soil properties.

Methods

Isotope ratios and nutrient content in biocrust tissue were analyzed in 13 Mediterranean shrublands along an aridity gradient. Differences in tissue traits between biocrust groups (lichens and mosses), and relationships between tissue traits and climatic and soil variables were examined.

Results

Lichens and mosses differed in δ¹³C, δ¹⁵N and N content, indicating distinct physical and physiological attributes. Tissue traits correlated strongly with numerous climatic variables, likely due to a modulator effect on biocrust water relations and metabolism. We found contrasting responses of lichen and moss traits to climate, although they responded similarly to soil properties. Overall, the most responsive trait was δ¹⁵N, suggesting this trait is the best to reflect integrated processes occurring in the atmosphere and soil.

Conclusions

Biocrust tissue traits arise as cost-effective, integrative ecological indicators of global change drivers in Mediterranean ecosystems, with potential applications in response-effect trait frameworks.

     

Colonization cues of leaf- and root-inhabiting bacterial microbiota of <Emphasis Type="Italic">Atractylodes lancea</Emphasis> derived in vitro and in vivo

Background and aims

Earthworms effect on plant growth is mediated by their dejections or “casts”, a complex mixture of organic matter, minerals and microbes. In casts, different processes such as organic matter mineralization and signal molecule production follow a complex temporal dynamics. An adaptation of root morphology to cast dynamics could allow an efficient nitrogen capture by the plant.

Methods

The plant Brachypodium distachyon was grown in a laboratory experiment with different proportions of casts of increasing ages. Casts were labelled with ¹⁵N to quantify the plant N uptake from the casts. Plant biomass and morphology, especially root system structure, were analysed.

Results

The age of casts had an effect on fine root length, highlighting the importance of the dynamics of cast maturation in root adaptation. Plant biomass production was affected by the interaction between the age and proportion of casts. A positive correlation between the ¹⁵N proportion in plant tissues and plant biomasses indicated that plants were more efficient in foraging N in casts than in the bulk soil.

Conclusions

Our results suggested that both a timely adaptation of the root system structure and a significant proportion of casts are necessary to observe a positive effect of casts on plant growth.

     

Co-managing soil and plant pathogens: effects of organic amendments on soil fertility and fungal pathogen survival

Backgrounds and aims

Interactions between plants can be both positive and negative, denoting facilitation and competition. Although facilitative effects of having legume neighbours (focus on yield productivity) are well studied, a better mechanistic understanding of how legumes interact with non-legumes in terms of root distribution is needed. We tested the effects of neighbour identity, its spatial location, as well as the effects of plant order of arrival on above and belowground traits and root distribution.

Methods

We performed a rhizotron experiment (4 weeks duration) in which we grew maize alone, with only a legume, only another grass, or with both species and tracked roots of the plant species using green and red fluorescent markers.

Results

Maize grew differently when it had a neighbour, with reduced development when growing with wheat compared to alone. Growing with a legume generally equated to the same outcome as not having a neighbour. Roots grew towards the legume species and away from the wheat. Order of arrival affected aboveground traits to a certain extent, but its effects on maize roots were dependent on spatial location.

Conclusions

Our study provides evidence of facilitation, showing the importance of the identity of the neighbours, together with their spatial location, and how order of arrival can modulate the outcome of these initial interactions.

     

Tree development in structural soil – an empirical below-ground in-situ study of urban trees in Copenhagen,Denmark

Aims

The purpose of this study is to evaluate root and stem growth, nutrient status and soil properties of a tree planting in structural soil.

Methods

Root and stem growth were measured at an 10-year-old urban planting of Tilia x europaea L. ‘Pallida’ established in structural soil on a paved square in Copenhagen, Denmark.Root abundance was quantified and soil profiles were established at three different distances from 10 trees (1.0, 1.9, and 2.9 m). Soil and leaf samples were analysed for nutrient concentrations.

Results

Abundant root presence was measured across the entire profile of structural soil (160?×?60 cm) at all distances, provided that structural soil layers were constructed correctly. Erroneously constructed structural soil layers however, resulted in impaired tree and root growth. Coarse root morphology was affected by the confined growing space in the voids of the stone matrix. The soil was relatively alkaline with an average pH (CaCl₂) of 7.3. Foliar analysis revealed deficiencies of K and Mn. These deficiencies might be due to a combined effect of high pH and low soil concentrations of these nutrients.

Conclusions

The study documents that correctly constructed structural soils allow and encourage root growth in load bearing layers.

     

Impact of plant growth-promoting bacteria on grain yield,protein content,and urea-<Superscript>15</Superscript> N recovery by maize in a Cerrado Oxisol

Background and aims

The inoculation of cereal crops with plant growth-promoting bacteria (PGPB) is a potential strategy to improve fertilizer-N acquisition by crops in soils with low capacity to supply N. A study was conducted to assess the impact of three inoculants on grain yield, protein content, and urea-¹⁵ N recovery in maize (Zea mays L.) under Cerrado soil and climate conditions.

Methods

The main treatments included inoculants containing (i) Azospirillum brasilense strain Sp245, (ii) A. brasilense strains AbV5 + AbV6, (iii) Herbaspirillum seropedicae strain ZAE94, and (iv) a non-inoculated control. The subtreatments were (i) urea-N fertilization (100 kg N ha^?1) at 30 days after sowing and (ii) no N addition at the stage. To determine fertilizer-N recovery, ¹⁵N–labelled urea was applied in microplots.

Results

Inoculants carrying A. brasilense improved urea-¹⁵ N acquisition efficiency in maize and also improved grain yield compared to the non-inoculated control, while urea-N fertilization enhanced grain quality by providing higher protein content.

Conclusion

Our results suggest that the inoculation of maize grains with PGPB represents a strategy to improve fertilizer-N recovery and maize yield in Cerrado soil with a low capacity to supply N.

     

Regulation of dauciform root formation and root phosphatase activities of sedges (<Emphasis Type="Italic">Carex</Emphasis>) by nitrogen and phosphorus

Aims

Dauciform roots (DR) are formed by some Cyperaceae under phosphorus (P) deficiency. To advance our understanding of their physiological function, I ask: Is DR formation regulated by shoot P status or external P supply? How does it respond to nitrogen (N)? Do DR enhance root monoesterase, diesterase or phytase activities and ability to utilize organic P?

Methods

Greenhouse experiments were carried out with two Carex species grown in sand with (1) different combinations of N and P supply, (2) local supply of N or P to root halves, and (3) different organic P forms.

Results

Carex flava produced DR in all treatments. The density of DR and phosphatase activities increased with N supply; they were regulated by shoot P status and external N (but not P) supply. All phosphatase activities increased with DR density. Carex muricata produced no DR and had lower diesterase activity than C. flava but both species grew equally well with diester-P.

Conclusions

DR and phosphatase activities are regulated by both N and P supply. Similar growth responses to nutrients in both species suggest small costs and benefits of DR under experimental conditions but confirmation is needed for plants grown on natural soils.

     

<Emphasis Type="Italic">Parthenium hysterophorus</Emphasis> L. (Asteraceae) invasion had limited impact on major soil nutrients and enzyme activity: Is the null effect real or reflects data insensitivity?

Background and aims

Competition from the annual grass Bromus tectorum threatens aridland perennial bunchgrass communities. Unlike annuals, perennials must allocate part of their first year nitrogen (N) budget to storage rather than growth, potentially placing them at a competitive disadvantage.

Methods

We evaluated N acquisition and conservation for two perennial bunchgrasses, Agropyron desertorum and Pseudoroegneria spicata, at the seedling stage to investigate potential trade-offs between storage and growth when grown with and without B. tectorum under two levels of soil N.

Results

Agropyron desertorum had higher growth rates, N uptake, and N productivity than P. spicata when grown without B. tectorum, but trait values were similarly low for both species under competition. Without competition, N resorption was poor under high soil N, but it was equally proficient among species under competition.

Conclusions

A. desertorum had higher growth rates and N productivity than P. spicata without competition, suggesting these traits may in part promote its greater success in restoration programs. However, B. tectorum neighbors reduced its trait advantage. As plant traits become more integral to restoration ecology, understanding how N capture and conservation traits vary across candidate species and under competition may improve our ability to select species with the highest likelihood of establishing in arid, nutrient-limited systems.

     

Kinetics of metal toxicity in plant roots and its effects on root morphology

Aims

We evaluated the efficacy of biochar application for suppressing bacterial wilt of tomato and identified the potential underlying mechanisms involved in the disease control.

Methods

We measured the impact of two different sized biochar (53–120 μm and 380–830 μm) on bacterial wilt incidence in a greenhouse experiment. The efficiency of different sized biochar for the adsorption of tomato root exudates and the pathogen was further examined in vitro. We also quantified the effects of biochar and tomato root exudates on two pathogen virulence factors, chemotaxis, swarming motility and examined the effect of biochar on pathogen root colonization.

Results

Fine biochar application (3%; w:w) significantly decreased the bacterial wilt incidence by 19.9%. Biochar with different particle size had similar adsorption capacity for root exudates, while fine biochar was efficient (91%) in pathogen adsorption. Root exudates and fine biochar increased the chemotaxis ability of pathogen, while fine biochar reduced pathogen swarming motility and rhizosphere colonization.

Conclusions

Application of fine biochar can significantly decreased bacterial wilt incidence. This was mechanistically explained by biochar ability to 1) adsorb pathogen directly and indirectly via adsorption of root exudates (based on pathogen chemotaxis) and to 2) directly suppress pathogen swarming motility and subsequent root colonization.

     

Plant-soil feedbacks and root responses of two Mediterranean oaks along a precipitation gradient

Aims

Plant-soil feedbacks (PSFs) have been shown to be relevant drivers of forest community dynamics. However, few studies have explored variation of PSFs along environmental gradients. In a framework of climate change, there is a great need to understand how interactions between plants and soil microbes respond along climatic gradients. Therefore, we compared PSFs along a precipitation gradient in Mediterranean oak forests and included trait responses. Following the Stress Gradient Hypothesis (SGH), we expected less negative or even positive PSFs in the physically harsh dry end of our gradient and more negative PSFs in the wettest end.

Methods

We grew Quercus ilex and Quercus suber acorns on soil inoculated with microbes sampled under adults of both species in six sites ranging in annual precipitation. After 4 months, we measured shoot biomass and allocation and morphological traits above and belowground.

Results

We found negative PSFs for Q. ilex independent of precipitation, whereas for Q. suber PSFs ranged from positive in dry sites to negative in wet sites, in agreement with the SGH. The leaf allocation showed patterns similar to shoot biomass, but belowground allocation and morphological traits revealed responses which could not be detected aboveground.

Conclusions

We provide first evidence for context-dependent PSFs along a precipitation gradient. Moreover, we show that measuring root traits can help improve our understanding of climate-dependent PSFs. Such understanding helps to predict plant soil microbe interactions, and their role as drivers of plant community dynamics under ongoing climate change.