Linking fine root lifespan to root chemical and morphological traits-A global analysis.

Fine root lifespan is a critical trait associated with contrasting root strategies of resource acquisition and protection. Yet, its position within the multidimensional "root economics space" synthesizing global root economics strategies is largely uncertain, and it is rarely represented in frameworks integrating plant trait variations. Here, we compiled the most comprehensive dataset of absorptive median root lifespan (MRL) data including 98 observations from 79 woody species using (mini-)rhizotrons across 40 sites and linked MRL to other plant traits to address questions of the regulators of MRL at large spatial scales. We demonstrate that MRL not only decreases with plant investment in root nitrogen (associated with more metabolically active tissues) but also increases with construction of larger diameter roots which is often associated with greater plant reliance on mycorrhizal symbionts. Although theories linking organ structure and function suggest that root traits should play a role in modulating MRL, we found no correlation between root traits associated with structural defense (root tissue density and specific root length) and MRL. Moreover, fine root and leaf lifespan were globally unrelated, except among evergreen species, suggesting contrasting evolutionary selection between leaves and roots facing contrasting environmental influences above vs. belowground. At large geographic scales, MRL was typically longer at sites with lower mean annual temperature and higher mean annual precipitation. Overall, this synthesis uncovered several key ecophysiological covariates and environmental drivers of MRL, highlighting broad avenues for accurate parametrization of global biogeochemical models and the understanding of ecosystem response to global climate change.

The mycorrhiza-specific ammonium transporter ZmAMT3;1 mediates mycorrhiza-dependent nitrogen uptake in maize roots.

Most plant species can form symbioses with arbuscular mycorrhizal fungi (AMFs), which may enhance the host plant's acquisition of soil nutrients. In contrast to phosphorus nutrition, the molecular mechanism of mycorrhizal nitrogen (N) uptake remains largely unknown, and its physiological relevance is unclear. Here, we identified a gene encoding an AMF-inducible ammonium transporter, ZmAMT3;1, in maize (Zea mays) roots. ZmAMT3;1 was specifically expressed in arbuscule-containing cortical cells and the encoded protein was localized at the peri-arbuscular membrane. Functional analysis in yeast and Xenopus oocytes indicated that ZmAMT3;1 mediated high-affinity ammonium transport, with the substrate NH4+ being accessed, but likely translocating uncharged NH3. Phosphorylation of ZmAMT3;1 at the C-terminus suppressed transport activity. Using ZmAMT3;1-RNAi transgenic maize lines grown in compartmented pot experiments, we demonstrated that substantial quantities of N were transferred from AMF to plants, and 68%-74% of this capacity was conferred by ZmAMT3;1. Under field conditions, the ZmAMT3;1-dependent mycorrhizal N pathway contributed >30% of postsilking N uptake. Furthermore, AMFs downregulated ZmAMT1;1a and ZmAMT1;3 protein abundance and transport activities expressed in the root epidermis, suggesting a trade-off between mycorrhizal and direct root N-uptake pathways. Taken together, our results provide a comprehensive understanding of mycorrhiza-dependent N uptake in maize and present a promising approach to improve N-acquisition efficiency via plant-microbe interactions.

Role of mycorrhizas and root exudates in plant uptake of soil nutrients (calcium, iron, magnesium, and potassium): has the puzzle been completely solved?

Anthropogenic global change is driving an increase in the frequency and intensity of drought and flood events, along with associated imbalances and limitation of several soil nutrients. In the context of an increasing human population, these impacts represent a global-scale challenge for biodiversity conservation and sustainable crop production to ensure food security. Plants have evolved strategies to enhance uptake of soil nutrients under environmental stress conditions; for example, symbioses with fungi (mycorrhization) in the rhizosphere and the release of exudates from roots. Although crop cultivation is managed for the effects of limited availability of nitrogen (N) and phosphorus (P), there is increasing evidence for limitation of plant growth and fitness because of the low availability of other soil nutrients such as the metals potassium (K), calcium (Ca), magnesium (Mg), and iron (Fe), which may become increasingly limiting for plant productivity under global change. The roles of mycorrhizas and plant exudates on N and P uptake have been studied intensively; however, our understanding of the effects on metal nutrients is less clear and still inconsistent. Here, we review the literature on the role of mycorrhizas and root exudates in plant uptake of key nutrients (N, P, K, Ca, Mg, and Fe) in the context of potential nutrient deficiencies in crop and non-crop terrestrial ecosystems, and identify knowledge gaps for future research to improve nutrient-uptake capacity in food crop plants.

Leaf phosphorus fractions vary with leaf economic traits among 35 Australian woody species.

Adaptations of plants to phosphorus (P) deficiency include reduced investment of leaf P in storage (orthophosphates in vacuoles), nucleic acids and membrane lipids. Yet, it is unclear how these adaptations are associated with plant ecological strategies. Five leaf P fractions (orthophosphate P, Pi ; metabolite P, PM ; nucleic acid P, PN ; lipid P, PL ; and residual P, PR ) were analysed alongside leaf economic traits among 35 Australian woody species from three habitats: one a high-P basalt-derived soil and two low-P sandstone-derived soils, one undisturbed and one disturbed by human activities with artificial P inputs. Species at the undisturbed low-P site generally exhibited lower concentrations of total leaf P ([Ptotal ]), primarily associated with lower concentrations of Pi , and PN . The relative allocation of P to each fraction varied little among sites, except that higher PL per [Ptotal ] (rPL ) was recorded at the undisturbed low-P site than at the high-P site. This higher rPL , reflecting relative allocation to membranes, was primarily associated with lower concentrations of leaf nitrogen at the undisturbed low-P site than at the high-P site. Associations between leaf P fractions and leaf nitrogen may provide a basis for understanding the variation in plant ecological strategies dependent on soil P availability.

Facilitative and competitive interactions between mycorrhizal and nonmycorrhizal plants in an extremely phosphorus-impoverished environment: role of ectomycorrhizal fungi and native oomycete pathogens in shaping species coexistence.

Nonmycorrhizal cluster root-forming species enhance the phosphorus (P) acquisition of mycorrhizal neighbours in P-impoverished megadiverse systems. However, whether mycorrhizal plants facilitate the defence of nonmycorrhizal plants against soil-borne pathogens, in return and via their symbiosis, remains unknown. We characterised growth and defence-related compounds in Banksia menziesii (nonmycorrhizal) and Eucalyptus todtiana (ectomycorrhizal, ECM) seedlings grown either in monoculture or mixture in a multifactorial glasshouse experiment involving ECM fungi and native oomycete pathogens. Roots of B. menziesii had higher levels of phytohormones (salicylic and jasmonic acids, jasmonoyl-isoleucine and 12-oxo-phytodienoic acid) than E. todtiana which further activated a salicylic acid-mediated defence response in roots of B. menziesii, but only in the presence of ECM fungi. We also found that B. menziesii induced a shift in the defence strategy of E. todtiana, from defence-related secondary metabolites (phenolic and flavonoid) towards induced phytohormone response pathways. We conclude that ECM fungi play a vital role in the interactions between mycorrhizal and nonmycorrhizal plants in a severely P-impoverished environment, by introducing a competitive component within the facilitation interaction between the two plant species with contrasting nutrient-acquisition strategies. This study sheds light on the interplay between beneficial and detrimental soil microbes that shape plant-plant interaction in severely nutrient-impoverished ecosystems.

Global change progressively increases foliar nitrogen-phosphorus ratios in China's subtropical forests.

Globally increased nitrogen (N) to phosphorus (P) ratios (N/P) affect the structure and functioning of terrestrial ecosystems, but few studies have addressed the variation of foliar N/P over time in subtropical forests. Foliar N/P indicates N versus P limitation in terrestrial ecosystems. Quantifying long-term dynamics of foliar N/P and their potential drivers is crucial for predicting nutrient status and functioning in forest ecosystems under global change. We detected temporal trends of foliar N/P, quantitatively estimated their potential drivers and their interaction between plant types (evergreen vs. deciduous and trees vs. shrubs), using 1811 herbarium specimens of 12 widely distributed species collected during 1920-2010 across China's subtropical forests. We found significant decreases in foliar P concentrations (23.1%) and increases in foliar N/P (21.2%). Foliar N/P increased more in evergreen species (22.9%) than in deciduous species (16.9%). Changes in atmospheric CO2 concentrations ( P CO 2 $$ {\mathrm{P}}_{{\mathrm{CO}}_2} $$ ), atmospheric N deposition and mean annual temperature (MAT) dominantly contributed to the increased foliar N/P of evergreen species, while P CO 2 $$ {\mathrm{P}}_{{\mathrm{CO}}_2} $$ , MAT, and vapor pressure deficit, to that of deciduous species. Under future Shared Socioeconomic Pathway (SSP) scenarios, increasing MAT and P CO 2 $$ {\mathrm{P}}_{{\mathrm{CO}}_2} $$ would continuously increase more foliar N/P in deciduous species than in evergreen species, with more 12.9%, 17.7%, and 19.4% versus 6.1%, 7.9%, and 8.9% of magnitudes under the scenarios of SSP1-2.6, SSP3-7.0, and SSP5-8.5, respectively. The results suggest that global change has intensified and will progressively aggravate N-P imbalance, further altering community composition and ecosystem functioning of subtropical forests.

Enhanced rock weathering increased soil phosphorus availability and altered root phosphorus-acquisition strategies.

Enhanced rock weathering (ERW) has been proposed as a measure to enhance the carbon (C)-sequestration potential and fertility of soils. The effects of this practice on the soil phosphorus (P) pools and the general mechanisms affecting microbial P cycling, as well as plant P uptake are not well understood. Here, the impact of ERW on soil P availability and microbial P cycling functional groups and root P-acquisition traits were explored through a 2-year wollastonite field addition experiment in a tropical rubber plantation. The results show that ERW significantly increased soil microbial carbon-use efficiency and total P concentrations and indirectly increased soil P availability by enhancing organic P mobilization and mineralization of rhizosheath carboxylates and phosphatase, respectively. Also, ERW stimulated the activities of P-solubilizing (gcd, ppa and ppx) and mineralizing enzymes (phoADN and phnAPHLFXIM), thus contributing to the inorganic P solubilization and organic P mineralization. Accompanying the increase in soil P availability, the P-acquisition strategy of the rubber fine roots changed from do-it-yourself acquisition by roots to dependence on mycorrhizal collaboration and the release of root exudates. In addition, the direct effects of ERW on root P-acquisition traits (such as root diameter, specific root length, and mycorrhizal colonization rate) may also be related to changes in the pattern of belowground carbon investments in plants. Our study provides a new insight that ERW increases carbon-sequestration potential and P availability in tropical forests and profoundly affects belowground plant resource-use strategies.

Adenanthos species (Proteaceae) in phosphorus-impoverished environments use a variety of phosphorus-acquisition strategies and achieve high-phosphorus-use efficiency.

BACKGROUND AND AIMS: Soils in south-western Australia are severely phosphorus (P) impoverished, and plants in this region have evolved a variety of P-acquisition strategies. Phosphorus acquisition by Adenanthos cygnorum (Proteaceae) is facilitated by P-mobilizing neighbours which allows it to extend its range of habitats. However, we do not know if other Adenanthos species also exhibit a strategy based on facilitation for P acquisition in P-impoverished environments. METHODS: We collected leaf and soil samples of Adenanthosbarbiger, A. cuneatus, A.meisneri,A. obovatus, A. sericeus and Adenanthos sp. Whicher Range (G.J. Keighery 9736) growing in their natural habitats at different locations within the severely P-limited megadiverse environment of south-western Australia. Hydroponic experiments were conducted to collect the carboxylates exuded by cluster roots. Pot experiments in soil were carried out to measure rhizosheath phosphatase activity. KEY RESULTS: We found no evidence for facilitation of P uptake in any of the studied Adenanthos species. Like most Proteaceae, A. cuneatus, A. meisneri, A. obovatus, A. sericeus and Adenanthos sp. Whicher Range (G.J. Keighery 9736) expressed P-mining strategies, including the formation of cluster roots. Cluster roots of A. obovatus were less effective than those of the other four Adenanthos species. In contrast to what is known for most Proteaceae, we found no cluster roots for A. barbiger. This species probably expressed a post-fire P-acquisition strategy. All Adenanthos species used P highly efficiently for photosynthesis, like other Proteaceae in similar natural habitats. CONCLUSIONS: Adenanthos is the first genus of Proteaceae found to express multiple P-acquisition strategies. The diversity of P-acquisition strategies in these Proteaceae, coupled with similarly diverse strategies in Fabaceae and Myrtaceae, demonstrates that caution is needed in making family- or genus-wide extrapolations about the strategies exhibited in severely P-impoverished megadiverse ecosystems.

Engineering microalgae for water phosphorus recovery to close the phosphorus cycle.

As a finite and non-renewable resource, phosphorus (P) is essential to all life and crucial for crop growth and food production. The boosted agricultural use and associated loss of P to the aquatic environment are increasing environmental pollution, harming ecosystems, and threatening future global food security. Thus, recovering and reusing P from water bodies is urgently needed to close the P cycle. As a natural, eco-friendly, and sustainable reclamation strategy, microalgae-based biological P recovery is considered a promising solution. However, the low P-accumulation capacity and P-removal efficiency of algal bioreactors restrict its application. Herein, it is demonstrated that manipulating genes involved in cellular P accumulation and signalling could triple the Chlamydomonas P-storage capacity to ~7% of dry biomass, which is the highest P concentration in plants to date. Furthermore, the engineered algae could recover P from wastewater almost three times faster than the unengineered one, which could be directly used as a P fertilizer. Thus, engineering genes involved in cellular P accumulation and signalling in microalgae could be a promising strategy to enhance P uptake and accumulation, which have the potential to accelerate the application of algae for P recovery from the water body and closing the P cycle.

Extraradical hyphae alleviate nitrogen deposition-induced phosphorus deficiency in ectomycorrhiza-dominated forests.

The continuous imbalance between nitrogen (N) and phosphorus (P) deposition is expected to shift many ecosystems from N- to P limitation. Extraradical hyphae of ectomycorrhizal (ECM) fungi play important roles in plant nutrient acquisition under nutrient deficiency. However, whether and how ECM hyphae enhance soil P availability to alleviate N-induced P deficiency remains unclear. We investigated the impacts of ECM hyphae on transformations among different soil P fractions and underlying mechanisms under N deposition in two ECM-dominated forests. Ectomycorrhizal hyphae enhanced soil P availability under N addition by stimulating mineralization of organic P (Po) and desorption and solubilization of secondary mineral P, as indicated by N-induced increase in positive hyphal effect on plant-available P pool and negative hyphal effects on Po and secondary mineral P pools. Moreover, ECM hyphae increased soil phosphatase activity and abundance of microbial genes associated with Po mineralization and inorganic P solubilization, while decreasing concentrations of Fe/Al oxides. Our results suggest that ECM hyphae can alleviate N-induced P deficiency in ECM-dominated forests by regulating interactions between microbial and abiotic factors involved in soil P transformations. This advances our understanding of plant acclimation strategies via mediating plant-mycorrhiza interactions to sustain forest production and functional stability under changing environments.

Soil phosphorus availability alters the correlations between root phosphorus-uptake rates and net photosynthesis of dominant C3 and C4 species in a typical temperate grassland of Northern China.

Phosphorus (P) fertilization can alleviate a soil P deficiency in grassland ecosystems. Understanding plant functional traits that enhance P uptake can improve grassland management. We measured impacts of P addition on soil chemical and microbial properties, net photosynthetic rate (Pn ) and nonstructural carbohydrate concentrations ([NSC]), and root P-uptake rate (PUR), morphology, anatomy, and exudation of two dominant grass species: Leymus chinensis (C3 ) and Cleistogenes squarrosa (C4 ). For L. chinensis, PUR and Pn showed a nonlinear correlation. Growing more adventitious roots compensated for the decrease in P transport per unit root length, so that it maintained a high PUR. For C. squarrosa, PUR and Pn presented a linear correlation. Increased Pn was associated with modifications in root morphology, which further enhanced its PUR and a greater surplus of photosynthate and significantly stimulated root exudation (proxied by leaf [Mn]), which had a greater impact on rhizosheath micro-environment and microbial PLFAs. Our results present correlations between the PUR and the Pn of L. chinensis and C. squarrosa and reveal that NSC appeared to drive the modifications of root morphology and exudation; they provide more objective basis for more efficient P-input in grasslands to address the urgent problem of P deficiency.

Phosphorus fractions in leaves.

Leaf phosphorus (P) comprises four major fractions: inorganic phosphate (Pi ), nucleic acids, phospholipids, P-containing metabolites and a residual fraction. In this review paper, we investigated whether allocation of P fractions varies among groups of terrestrial vascular plants, and is indicative of a species' strategy to use P efficiently. We found that as leaf total P concentration increases, the Pi fraction increases the most, without a plateau, while other fractions plateau. Variability of the concentrations of leaf P fractions is greatest among families > species(family) > regions > plant life forms. The percentage of total P allocated to nucleic acid-P (20-35%) and lipid-P (14-34%) varies less among families/species. High photosynthetic P-use efficiency is associated with low concentrations of all P fractions, and preferential allocation of P to metabolite-P and mesophyll cells. Sequential resorption of P from senescing leaves starts with Pi , followed by metabolite-P, and then other organic P fractions. Allocation of P to leaf P fractions varies with season. Leaf phytate concentrations vary considerably among species, associated with variation in photosynthesis and defence. Plasticity of P allocation to its fractions is important for acclimation to low soil P availability, and species-specific P allocation is needed for co-occurrence with other species.

A novel proxy to examine interspecific phosphorus facilitation between plant species.

Resource complementarity can contribute to enhanced ecosystem functioning in diverse plant communities, but the role of facilitation in the enhanced complementarity is poorly understood. Here, we use leaf manganese concentration ([Mn]) as a proxy for rhizosheath carboxylate concentration to explore novel mechanisms of complementarity mediated by phosphorus (P) facilitation. In pot experiments, we showed that mixtures involving Carex korshinskyi, an efficient P-mobilizing species, exhibited greater biomass and relative complementarity effect than combinations without C. korshinskyi on P-deficient soils. Compared with monocultures, leaf [Mn] and [P] of species that are inefficient at P mobilization increased by 27% and 21% when grown with C. korshinskyi (i.e. interspecific P facilitation via carboxylates) rather than next to another inefficient P-mobilizing species. This experimental result was supported by a meta-analysis including a range of efficient P-mobilizing species. Phosphorus facilitation enhanced the relative complementarity effect in low-P environments, related to a greater change in several facilitated species of their root morphological traits relative to those in monoculture. Using leaf [Mn] as a proxy, we highlight a vital mechanism of interspecific P facilitation via belowground processes and provide evidence for the pivotal role of P facilitation mediated by the plasticity of root traits in biodiversity research.

Differences in foliar phosphorus fractions, rather than in cell-specific phosphorus allocation, underlie contrasting photosynthetic phosphorus use efficiency among chickpea genotypes.

Although significant intraspecific variation in photosynthetic phosphorus (P) use efficiency (PPUE) has been shown in numerous species, we still know little about the biochemical basis for differences in PPUE among genotypes within a species. Here, we grew two high PPUE and two low PPUE chickpea (Cicer arietinum) genotypes with low P supply in a glasshouse to compare their photosynthesis-related traits, total foliar P concentration ([P]) and chemical P fractions (i.e. inorganic P (Pi), metabolite P, lipid P, nucleic acid P, and residual P). Foliar cell-specific nutrient concentrations including P were characterized using elemental X-ray microanalysis. Genotypes with high PPUE showed lower total foliar [P] without slower photosynthetic rates. No consistent differences in cellular [P] between the epidermis and mesophyll cells occurred across the four genotypes. In contrast, high PPUE was associated with lower allocation to Pi and metabolite P, with PPUE being negatively correlated with the percentage of these two fractions. Furthermore, a lower allocation to Pi and metabolite P was correlated with a greater allocation to nucleic acid P, but not to lipid P. Collectively, our results suggest that a different allocation to foliar P fractions, rather than preferential P allocation to specific leaf tissues, underlies the contrasting PPUE among chickpea genotypes.

Growth-defence trade-off in rice: fast-growing and acquisitive genotypes have lower expression of genes involved in immunity.

Plant ecologists and molecular biologists have long considered the hypothesis of a trade-off between plant growth and defence separately. In particular, how genes thought to control the growth-defence trade-off at the molecular level relate to trait-based frameworks in functional ecology, such as the slow-fast plant economics spectrum, is unknown. We grew 49 phenotypically diverse rice genotypes in pots under optimal conditions and measured growth-related functional traits and the constitutive expression of 11 genes involved in plant defence. We also quantified the concentration of silicon (Si) in leaves to estimate silica-based defences. Rice genotypes were aligned along a slow-fast continuum, with slow-growing, late-flowering genotypes versus fast-growing, early-flowering genotypes. Leaf dry matter content and leaf Si concentrations were not aligned with this axis and negatively correlated with each other. Live-fast genotypes exhibited greater expression of OsNPR1, a regulator of the salicylic acid pathway that promotes plant defence while suppressing plant growth. These genotypes also exhibited greater expression of SPL7 and GH3.2, which are also involved in both stress resistance and growth. Our results do not support the hypothesis of a growth-defence trade-off when leaf Si and leaf dry matter content are considered, but they do when hormonal pathway genes are considered. We demonstrate the benefits of combining ecological and molecular approaches to elucidate the growth-defence trade-off, opening new avenues for plant breeding and crop science.

Extraradical hyphae exhibit more plastic nutrient-acquisition strategies than roots under nitrogen enrichment in ectomycorrhiza-dominated forests.

Ectomycorrhizal (ECM) functional traits related to nutrient acquisition are impacted by nitrogen (N) deposition. However, less is known about whether these nutrient-acquisition traits associated with roots and hyphae differentially respond to increased N deposition in ECM-dominated forests with different initial N status. We conducted a chronic N addition experiment (25 kg N ha-1 year-1 ) in two ECM-dominated forests with contrasting initial N status, that is, a Pinus armandii forest (with relatively low N availability) and a Picea asperata forest (with relatively high N availability), to assess nutrient-mining and nutrient-foraging strategies associated with roots and hyphae under N addition. We show that nutrient-acquisition strategies of roots and hyphae differently respond to increased N addition. Root nutrient-acquisition strategies showed a consistent response to N addition, regardless of initial forest nutrient status, shifting from organic N mining toward inorganic N foraging. In contrast, the hyphal nutrient-acquisition strategy showed diverse responses to N addition depending on initial forest N status. In the Pinus armandii forest, trees increased belowground carbon (C) allocation to ECM fungi thus enhancing hyphal N-mining capacity under increased N availability. By comparison, in the Picea asperata forest, ECM fungi enhanced both capacities of P foraging and P mining in response to N-induced P limitation. In conclusion, our results demonstrate that ECM fungal hyphae exhibit greater plasticity in nutrient-mining and nutrient-foraging strategies than roots do in response to changes of nutrient status induced by N deposition. This study highlights the importance of ECM associations in tree acclimation and forest function stability under changing environments.

Faster accumulation and greater contribution of glomalin to the soil organic carbon pool than amino sugars do under tropical coastal forest restoration.

Microbial metabolic products play a vital role in maintaining ecosystem multifunctionality, such as soil physical structure and soil organic carbon (SOC) preservation. Afforestation is an effective strategy to restore degraded land. Glomalin-related soil proteins (GRSP) and amino sugars are regarded as stable microbial-derived C, and their distribution within soil aggregates affects soil structure stability and SOC sequestration. However, the information about how afforestation affects the microbial contribution to SOC pools within aggregates is poorly understood. We assessed the accumulation and contribution of GRSP and amino sugars within soil aggregates along a restoration chronosequence (Bare land, Eucalyptus exserta plantation, native species mixed forest, and native forest) in tropical coastal terraces. Amino sugars and GRSP concentrations increased, whereas their contributions to the SOC pool decreased along the restoration chronosequence. Although microaggregates harbored greater microbial abundances, amino sugars and GRSP concentrations were not significantly affected by aggregate sizes. Interestingly, the contributions of amino sugars and GRSP to SOC pools decreased with decreasing aggregate size which might be associated with increased accumulation of plant-derived C. However, the relative change rate of GRSP was consistently greater in all restoration chronosequences than that of amino sugars. The accumulation of GRSP and amino sugars in SOC pools was closely associated with the dynamics of soil fertility and the microbial community. Our findings suggest that GRSP accumulates faster and contributes more to SOC pools during restoration than amino sugars did which was greatly affected by aggregate sizes. Afforestation substantially enhanced soil quality with native forest comprising species sequestering more SOC than the monoculture plantation did. Such information is invaluable for improving our mechanistic understanding of microbial control over SOC preservation during degraded ecosystem restoration. Our findings also show that plantations using arbuscular mycorrhizal plants can be an effective practice to sequester more soil carbon during restoration.

Coordination of photosynthetic traits across soil and climate gradients.

"Least-cost theory" posits that C3 plants should balance rates of photosynthetic water loss and carboxylation in relation to the relative acquisition and maintenance costs of resources required for these activities. Here we investigated the dependency of photosynthetic traits on climate and soil properties using a new Australia-wide trait dataset spanning 528 species from 67 sites. We tested the hypotheses that plants on relatively cold or dry sites, or on relatively more fertile sites, would typically operate at greater CO2 drawdown (lower ratio of leaf internal to ambient CO2 , Ci :Ca ) during light-saturated photosynthesis, and at higher leaf N per area (Narea ) and higher carboxylation capacity (Vcmax 25 ) for a given rate of stomatal conductance to water vapour, gsw . These results would be indicative of plants having relatively higher water costs than nutrient costs. In general, our hypotheses were supported. Soil total phosphorus (P) concentration and (more weakly) soil pH exerted positive effects on the Narea -gsw and Vcmax 25 -gsw slopes, and negative effects on Ci :Ca . The P effect strengthened when the effect of climate was removed via partial regression. We observed similar trends with increasing soil cation exchange capacity and clay content, which affect soil nutrient availability, and found that soil properties explained similar amounts of variation in the focal traits as climate did. Although climate typically explained more trait variation than soil did, together they explained up to 52% of variation in the slope relationships and soil properties explained up to 30% of the variation in individual traits. Soils influenced photosynthetic traits as well as their coordination. In particular, the influence of soil P likely reflects the Australia's geologically ancient low-relief landscapes with highly leached soils. Least-cost theory provides a valuable framework for understanding trade-offs between resource costs and use in plants, including limiting soil nutrients.

Morphological and physiological traits of dominant plant species in response to mowing in a temperate steppe.

Mowing is common grassland management to feed livestock during winter by harvesting hay in many high-latitude regions in autumn. The trait-based approach has been used to explain the responses of the plant community to disturbance resulting from environmental changes and human activities. However, few studies have focused on the mechanisms underlying the responses of grassland ecosystems to mowing from the perspective of plant traits. Here, we investigated the effects of mowing on the plant community of a temperate steppe in Inner Mongolia of northern China by field experiments to dissect the trade-off between morphological and physiological traits in response to short-term (4 years) and long-term (16 years) mowing. Specifically, we evaluated the two strategies associated with the nutrient acquisition of two dominant species in response to mowing by measuring leaf and root morphological traits and physiological traits of root carboxylate exudation, arbuscular mycorrhizal fungi (AMF) colonization and soil microbial community. We found that long-term mowing, but not short-term mowing, led to an increase in species richness. In addition, mowing decreased the overall plant biomass of the grassland community, but enhanced and suppressed the growth of forbs and grasses, respectively. However, the ratio of forbs to grasses in the community was dependent on mowing duration, such that forbs became more dominant than the grasses under long-term mowing. Our results revealed that short-term mowing reduced soil microbial biodiversity and root colonization of AMF in the grass Stipa krylovii, while the root AMF colonization and carboxylate exudation in the forb Artemisia frigida were enhanced by short-term mowing. In long-term mowing, the functional traits associated with leaf resource conservation (i.e., leaf tissue density) and root resource acquisition were reduced in the grass, while the functional traits related to leaf resource acquisition and root resource conservation were increased in the forb, highlighting the species specificity and divergence in leaf and root traits in the grass and forb of temperate steppe in response to mowing. These novel findings demonstrate that physiological and morphological strategies are the main drivers for dominant species in response to mowing in temperate grasslands.

Enhanced nodulation and phosphorus acquisition from sparingly-soluble iron phosphate upon treatment with arbuscular mycorrhizal fungi in chickpea.

The coordination/trade-off among below-ground strategies for phosphorus (P) acquisition, including root morphology, carboxylate exudation and colonisation by arbuscular mycorrhizal fungi (AMF), is not well understood. This is the first study investigating the relationships between root nodulation, morphology, carboxylates and colonisation by an indigenous community of AMF under varying P levels and source. Two chickpea genotypes with contrasting amounts of rhizosheath carboxylates were grown in pots at six P levels (from 0 to 160 µg g-1 ) as KH2 PO4 (KP, highly soluble) or FePO4 (FeP, sparingly soluble), with or without AMF (±AMF) treatment. Under both FeP and KP, the presence of AMF inhibited shoot growth and shoot branching, decreased total root length and specific root length, increased mean root diameter and root tissue density and reduced carboxylates. However, the role of AMF in acquiring P differed between the two P sources, with the enhanced P acquisition under FeP while not under KP. Co-inoculation of AMF and rhizobia enhanced nodulation under FeP, but not under KP. Our results suggest that the effects of AMF on shoot branching were mediated by cytokinins as the reduced shoot branching in FeP40 and KP40 under +AMF relative to -AMF coincided with a decreased concentration of cytokinins in xylem sap for both genotypes.