Evaluation of the trade-off between water use efficiency and nutrient use efficiency in two semiarid coniferous tree species growing on an organic amended metalliferous mine tailing substrate.

We evaluated the ecophysiological responses of two semiarid coniferous tree species, Pinus halepensis and Tetraclinis articulata, growing on a nutrient-poor metalliferous mine tailings substrate to organic amendments (biochar and/or organic municipal waste). The trees were grown in mesocosms under irrigated conditions for 20 months. Then, a comprehensive characterization of soil and plant parameters (including stable isotopes) was carried out. Treatments containing municipal waste showed better soil fertility indicators (approximately 2-fold higher organic carbon and total nitrogen concentrations) and higher plant biomass (up to 5-fold higher) than unamended and only biochar treatments. Trees in most of the treatments exhibited leaf N/P ratios <14 indicating severe N limitation of plant growth. Metal uptake was below phytotoxic levels across all the treatments. Leaf δ13C values correlated positively with δ18O across treatments for both species indicating increasing water use efficiency with tighter stomatal regulation of water flux, and with T. articulata exhibiting tighter stomatal control (higher δ18O values) than P. halepensis. Trees in treatments containing only biochar did not differ in ecophysiological performance from those in the unamended treatments. In contrast, leaf stable isotopes revealed sharply increased of time-integrated photosynthetic activity (favoured by higher leaf N concentrations) combined with lower time-integrated stomatal conductance in the treatments containing municipal waste, indicating greatly enhanced water use efficiency in better nourished plants. Trade-offs between water use efficiency and nutrient (N and P) use efficiency were evident across treatments, with higher leaf nutrient concentrations associated with higher water use efficiency, at the cost of a lower nutrient use efficiency. These trade-offs were not impaired by the high metal concentrations of the tailings substrate, indicating that ecophysiological adjustments in response to changes in plant nutrient status promoted by the addition of organic amendments are critical for the adaptability of native tree species employed in the phytostabilisation of mine tailings.

Tight coupling between leaf δ13 C and N content along leaf ageing in the N2 -fixing legume tree black locust (Robinia pseudoacacia L.).

N2 -fixing legumes can strongly affect ecosystem functions by supplying nitrogen (N) and improving the carbon-fixing capacity of vegetation. Still, the question of how their leaf-level N status and carbon metabolism are coordinated along leaf ageing remains unexplored. Leaf tissue carbon isotopic composition (δ13 C) provides a useful indicator of time-integrated intrinsic water use efficiency (WUEi). Here, we quantified the seasonal changes of leaf δ13 C, N content on a mass and area basis (Nmass , Narea , respectively), Δ18 O (leaf 18 O enrichment above source water, a proxy of time-integrated stomatal conductance) and morphological traits in an emblematic N2 -fixing legume tree, the black locust (Robinia pseudoacacia L.), at a subtropical site in Southwest China. We also measured xylem, soil and rainwater isotopes (δ18 O, δ2 H) to characterize tree water uptake patterns. Xylem water isotopic data reveal that black locust primarily used shallow soil water in this humid habitat. Black locust exhibited a decreasing δ13 C along leaf ageing, which was largely driven by decreasing leaf Nmass , despite roughly constant Narea . In contrast, the decreasing δ13 C along leaf ageing was largely uncoupled from parallel increases in Δ18 O and leaf thickness. Leaf N content is used as a proxy of leaf photosynthetic capacity; thus, it plays a key role in determining the seasonality in δ13 C, whereas the roles of stomatal conductance and leaf morphology are minor. Black locust leaves can effectively adjust to changing environmental conditions along leaf ageing through LMA increases and moderate stomatal conductance reduction while maintaining constant Narea to optimize photosynthesis and carbon assimilation, despite declining leaf Nmass and δ13 C.

Higher leaf nitrogen content is linked to tighter stomatal regulation of transpiration and more efficient water use across dryland trees.

The least-cost economic theory of photosynthesis shows that water and nitrogen are mutually substitutable resources to achieve a given carbon gain. However, vegetation in the Sahel has to cope with the dual challenge imposed by drought and nutrient-poor soils. We addressed how variation in leaf nitrogen per area (Narea ) modulates leaf oxygen and carbon isotopic composition (δ18 O, δ13 C), as proxies of stomatal conductance and water-use efficiency, across 34 Sahelian woody species. Dryland species exhibited diverging leaf δ18 O and δ13 C values, indicating large interspecific variation in time-integrated stomatal conductance and water-use efficiency. Structural equation modeling revealed that leaf Narea is a pivotal trait linked to multiple water-use traits. Leaf Narea was positively linked to both δ18 O and δ13 C, suggesting higher carboxylation capacity and tighter stomatal regulation of transpiration in N-rich species, which allows them to achieve higher water-use efficiency and more conservative water use. These adaptations represent a key physiological advantage of N-rich species, such as legumes, that could contribute to their dominance across many dryland regions. This is the first report of a robust mechanistic link between leaf Narea and δ18 O in dryland vegetation that is consistent with core principles of plant physiology.

The role of nutritional impairment in carbon-water balance of silver fir drought-induced dieback.

Rear-edge populations at the xeric distribution limit of tree species are particularly vulnerable to forest dieback triggered by drought. This is the case of silver fir (Abies alba) forests located in Southwestern Europe. While silver fir drought-induced dieback patterns have been previously explored, information on the role played by nutritional impairment is lacking despite its potential interactions with tree carbon-water balances. We performed a comparative analysis of radial growth, intrinsic water-use efficiency (iWUE), oxygen isotopes (δ18 O) and nutrient concentrations in leaves of declining (DD) and non-declining (ND) trees in silver fir in four forests in the Spanish Pyrenees. We also evaluated the relationships among dieback predisposition, intraspecific trait variation (wood density and leaf traits) and rhizosphere soil physical-chemical properties. The onset of growth decline in DD trees occurred more than two decades ago, and they subsequently showed low growth resilience against droughts. The DD trees presented consistently lower foliar concentrations of nutrients such as P, K, Cu and Ni than ND trees. The strong effects of foliar nutrient status on growth resilience indices support the key role played by mineral nutrition in tree functioning and growth before, during and after drought. In contrast, variability in wood density and leaf morphological traits, as well as soil properties, showed weak relationships with tree nutritional status and drought performance. At the low elevation, warmer sites, DD trees showed stronger climate-growth relationships and lower δ18 O than ND trees. The uncoupling between iWUE and δ18 O, together with the positive correlations between P and K leaf concentrations and δ18 O, point to deeper soil/bedrock water sources and vertical decoupling between nutrient and water uptake in DD trees. This study provides novel insights into the mechanisms driving silver fir dieback and highlights the need to incorporate tree nutrition into forest dieback studies.

Las poblaciones del límite xérico de distribución de las especies de árboles son particularmente vulnerables al decaimiento forestal inducido por sequía. Este es el caso de los bosques de abeto (Abies alba) situados en el suroeste de Europa. Si bien los patrones de decaimiento provocado por sequía del abeto se han explorado previamente, falta información sobre el papel que desempeña el deterioro nutricional a pesar de sus interacciones potenciales con los balances de agua y carbono de los árboles. En este estudio, hemos realizado un análisis comparativo del crecimiento radial, la eficiencia intrínseca del uso del agua (iWUE), los isótopos de oxígeno (δ18O) y las concentraciones de nutrientes en hojas de árboles decaídos (DD) y no decaídos (ND) en cuatro abetares de los Pirineos españoles. También evaluamos las relaciones entre la predisposición al decaimiento, la variación de rasgos intraespecíficos (densidad de la madera y rasgos de las hojas) y las propiedades físico-químicas de la rizosfera. El inicio de la disminución del crecimiento en los árboles DD ocurrió hace más de dos décadas y posteriormente mostraron una baja resiliencia de crecimiento frente a las sequías. Los árboles DD presentaron concentraciones foliares consistentemente más bajas de nutrientes como P, K, Cu y Ni que los árboles ND. Los fuertes efectos del estado de los nutrientes foliares en los índices de resiliencia del crecimiento respaldan el papel clave que desempeña la nutrición mineral en el funcionamiento y el crecimiento de los árboles antes, durante y después de la sequía. En contraste, la variabilidad en la densidad de la madera y los rasgos morfológicos de las hojas, así como las propiedades del suelo, mostraron una relación débil con el estado nutricional de los árboles y la respuesta del crecimiento a la sequía. En los sitios más cálidos y de baja elevación, los árboles DD mostraron relaciones clima-crecimiento más fuertes y un δ18 O más bajo que los árboles ND. El desacoplamiento entre iWUE y δ18 O, junto con las correlaciones positivas entre las concentraciones foliares de P y K y δ18 O, apuntan a fuentes de agua más profundas del suelo/lecho rocoso y un desacoplamiento vertical entre la absorción de nutrientes y agua en los árboles DD. Este estudio proporciona información novedosa sobre los mecanismos que impulsan el decaimiento del abeto y destaca la necesidad de incorporar la nutrición de los árboles en los estudios de muerte regresiva del bosque.

Contrasting Responses of Arbuscular Mycorrhizal Fungal Families to Simulated Climate Warming and Drying in a Semiarid Shrubland.

We carried out a 4-year manipulative field experiment in a semiarid shrubland in southeastern Spain to assess the impacts of experimental warming (W), rainfall reduction (RR), and their combination (W + RR) on the composition and diversity of arbuscular mycorrhizal fungal (AMF) communities in rhizosphere soil of H. syriacum and G. struthium shrubs using single-molecule real-time (SMRT) DNA sequencing. Across climate treatments, we encountered 109 AMF operational taxonomic units (OTUs) that were assigned to four families: Glomeraceae (93.94%), Gigasporaceae (2.19%), Claroideoglomeraceae (1.95%), and Diversisporaceae (1.92%). AMF community composition and diversity at OTU level were unaffected by the climate manipulation treatments, except for a significant decrease in AMF OTU richness in the W treatment relative to the control. However, we found a significant decrease of AMF family richness in all climate manipulation treatments relative to the control treatment. Members of the Gigasporaceae and Diversisporaceae families appeared to be highly vulnerable to intensification of heat and drought stress, as their abundances decreased by 67% and 77%, respectively, in the W + RR treatment relative to current ambient conditions. In contrast, the relative abundance and dominance of the Glomeraceae family within the AMF community increased significantly under the W + RR treatment, with Glomeraceae being the indicator family for the W + RR treatment. The interaction between warming and rainfall reduction had a significant effect on AMF community structure at family level. These findings provide new insights to help in the conservation of the soil biodiversity facing climate change in dryland ecosystems.

The Economics Spectrum Drives Root Trait Strategies in Mediterranean Vegetation.

Extensive research efforts are devoted to understand fine root trait variation and to confirm the existence of a belowground root economics spectrum (RES) from acquisitive to conservative root strategies that is analogous to the leaf economics spectrum (LES). The economics spectrum implies a trade-off between maximizing resource acquisition and productivity or maximizing resource conservation and longevity; however, this theoretical framework still remains controversial for roots. We compiled a database of 320 Mediterranean woody and herbaceous species to critically assess if the classic economics spectrum theory can be broadly extended to roots. Fine roots displayed a wide diversity of forms and properties in Mediterranean vegetation, resulting in a multidimensional trait space. The main trend of variation in this multidimensional root space is analogous to the main axis of LES, while the second trend of variation is partially determined by an anatomical trade-off between tissue density and diameter. Specific root area (SRA) is the main trait explaining species distribution along the RES, regardless of the selected traits. We advocate for the need to unify and standardize the criteria and approaches used within the economics framework between leaves and roots, for the sake of theoretical consistency.

Lower relative abundance of ectomycorrhizal fungi under a warmer and drier climate is linked to enhanced soil organic matter decomposition.

The aboveground impacts of climate change receive extensive research attention, but climate change could also alter belowground processes such as the delicate balance between free-living fungal decomposers and nutrient-scavenging mycorrhizal fungi that can inhibit decomposition through a mechanism called the Gadgil effect. We investigated how climate change-induced reductions in plant survival, photosynthesis and productivity alter soil fungal community composition in a mixed arbuscular/ectomycorrhizal (AM/EM) semiarid shrubland exposed to experimental warming (W) and/or rainfall reduction (RR). We hypothesised that increased EM host plant mortality under a warmer and drier climate might decrease ectomycorrhizal fungal (EMF) abundance, thereby favouring the proliferation and activity of fungal saprotrophs. The relative abundance of EMF sequences decreased by 57.5% under W+RR, which was accompanied by reductions in the activity of hydrolytic enzymes involved in the acquisition of organic-bound nutrients by EMF and their host plants. W+RR thereby created an enhanced potential for soil organic matter (SOM) breakdown and nitrogen mineralisation by decomposers, as revealed by 127-190% increases in dissolved organic carbon and nitrogen, respectively, and decreasing SOM content in soil. Climate aridification impacts on vegetation can cascade belowground through shifts in fungal guild structure that alter ecosystem biogeochemistry and accelerate SOM decomposition by reducing the Gadgil effect.

Vertical decoupling of soil nutrients and water under climate warming reduces plant cumulative nutrient uptake, water-use efficiency and productivity.

Warming-induced desiccation of the fertile topsoil layer could lead to decreased nutrient diffusion, mobility, mineralization and uptake by roots. Increased vertical decoupling between nutrients in topsoil and water availability in subsoil/bedrock layers under warming could thereby reduce cumulative nutrient uptake over the growing season. We used a Mediterranean semiarid shrubland as model system to assess the impacts of warming-induced topsoil desiccation on plant water- and nutrient-use patterns. A 6 yr manipulative field experiment examined the effects of warming (2.5°C), rainfall reduction (30%) and their combination on soil resource utilization by Helianthemum squamatum shrubs. A drier fertile topsoil ('growth pool') under warming led to greater proportional utilization of water from deeper, wetter, but less fertile subsoil/bedrock layers ('maintenance pool') by plants. This was linked to decreased cumulative nutrient uptake, increased nonstomatal (nutritional) limitation of photosynthesis and reduced water-use efficiency, above-ground biomass growth and drought survival. Whereas a shift to greater utilization of water stored in deep subsoil/bedrock may buffer the negative impact of warming-induced topsoil desiccation on transpiration, this plastic response cannot compensate for the associated reduction in cumulative nutrient uptake and carbon assimilation, which may compromise the capacity of plants to adjust to a warmer and drier climate.

Nutrient limitation determines the suitability of a municipal organic waste for phytomanaging metal(loid) enriched mine tailings with a pine-grass co-culture.

The suitable phytomanaging of mine tailings not only requires an improvement of soil fertility but also the assessment of the biotic interactions between the selected plant species. This study aimed to evaluate the effect of an organic amendment on the response of two plant species of contrasting habit, a tree, Pinus halepensis and a grass, Piptatherum miliaceum growing on a metal(loid)-contaminated substrate collected from mine tailings. Pots containing single plant individuals or their combination, with and without organic amendment (at 10% rate), were established and grown in a greenhouse for 13 months. Plant biomass, foliar ionome, leaf δ15N and metal(loid) concentrations were measured at the end of the experiment. The amendment alleviated P deficiency in the substrate and strongly stimulated biomass production by both plant species (10-fold for pine; 90-fold for the grass), leading to more balanced N/P ratios in leaves (especially for the grass). Co-culture with the grass negatively affected pine growth, decreasing total biomass and leaf δ15N values and inducing severe N deficiency (leaf N/P ratio<10). In contrast, co-culture with pine improved the nutrient status and growth of the grass, but only under non-amended conditions. Needle metal(loid) concentrations in P. halepensis were affected by both amendment addition and co-culture with the grass. High biomass growth with low metal(loid) concentrations in P. miliaceum leaves for the amended treatment makes this grass species suitable for the phytomanagement of metal(loid) polluted tailings, since it achieves high biomass production together with low concentrations of metal(loid)s in edible/senescent parts.

Poor plant performance under simulated climate change is linked to mycorrhizal responses in a semiarid shrubland.

Warmer and drier conditions associated with ongoing climate change will increase abiotic stress for plants and mycorrhizal fungi in drylands worldwide, thereby potentially reducing vegetation cover and productivity and increasing the risk of land degradation and desertification. Rhizosphere microbial interactions and feedbacks are critical processes that could either mitigate or aggravate the vulnerability of dryland vegetation to forecasted climate change.We conducted a four-year manipulative study in a semiarid shrubland in the Iberian Peninsula to assess the effects of warming (~2.5ºC; W), rainfall reduction (~30%; RR) and their combination (W+RR) on the performance of native shrubs (Helianthemum squamatum) and their associated mycorrhizal fungi.Warming (W and W+RR) decreased the net photosynthetic rates of H. squamatum shrubs by ~31% despite concurrent increases in stomatal conductance (~33%), leading to sharp decreases (~50%) in water use efficiency. Warming also advanced growth phenology, decreased leaf nitrogen and phosphorus contents per unit area, reduced shoot biomass production by ~36% and decreased survival during a dry year in both W and W+RR plants. Plants under RR showed more moderate decreases (~10-20%) in photosynthesis, stomatal conductance and shoot growth.Warming, RR and W+RR altered ectomycorrhizal fungal (EMF) community structure and drastically reduced the relative abundance of EMF sequences obtained by high-throughput sequencing, a response associated with decreases in the leaf nitrogen, phosphorus and dry matter contents of their host plants. In contrast to EMF, the community structure and relative sequence abundances of other non-mycorrhizal fungal guilds were not significantly affected by the climate manipulation treatments.Synthesis: Our findings highlight the vulnerability of both native plants and their symbiotic mycorrhizal fungi to climate warming and drying in semiarid shrublands, and point to the importance of a deeper understanding of plant-soil feedbacks to predict dryland vegetation responses to forecasted aridification. The interdependent responses of plants and ectomycorrhizal fungi to warming and rainfall reduction may lead to a detrimental feedback loop on vegetation productivity and nutrient pool size, which could amplify the adverse impacts of forecasted climate change on ecosystem functioning in EMF-dominated drylands.

Importance of intra- and interspecific plant interactions for the phytomanagement of semiarid mine tailings using the tree species Pinus halepensis.

The objective of this work was to evaluate the effects of plant interactions (intra- and interspecific) on the growth and metal(loid) uptake of the tree species Pinus halepensis to determine its suitability for the phytomanagement of semiarid mine tailings. The pioneer tailings colonizer grass Piptatherum miliaceum was selected for assessing interspecific interactions. The experiment was conducted following a pot experimental design employing mine tailings soil. Pots containing single individuals of P. halepensis or P. miliaceum and pots containing combinations with pines (two pines per pot, or one pine and one grass per pot) were used. The analyses included the determination of plant biomass, foliar element status and stable isotope composition, metal(loid) uptake and its translocation to different plant organs. P. halepensis strongly favoured the growth of P. miliaceum by increasing 9-fold the latter's biomass and alleviating its P limitation. In this interspecific treatment P. halepensis showed a strong N limitation (N/P = 7), which negatively affected its growth, (to about half the biomass of that obtained for the other treatments) and exhibited a significant increase in some metals translocation (especially Cd) into aerial parts. Interestingly, P. miliaceum showed a decrease in the root to leaves translocation factor for most of metals when growing together with pines. The effects of the intraespecific combination on growth and metal uptake in P. halepensis were less relevant than those obtained for the interspecific one. Further research should be focused on testing the behaviour of plant co-cultures under the addition of N or P amendments which could alleviate the negative effects of plant competition.

Nurse plants transfer more nitrogen to distantly related species.

Plant facilitative interactions enhance co-occurrence between distant relatives, partly due to limited overlap in resource requirements. We propose a different mechanism for the coexistence of distant relatives based on positive interactions of nutrient sharing. Nutrients move between plants following source-sink gradients driven by plant traits that allow these gradients to establish. Specifically, nitrogen (N) concentration gradients can arise from variation in leaf N content across plants species. As many ecologically relevant traits, we hypothesize that leaf N content is phylogenetically conserved and can result in N gradients promoting N transfer among distant relatives. In a Mexican desert community governed by facilitation, we labelled nurse plants (Mimosa luisana) with 15 N and measured its transfer to 14 other species in the community, spanning the range of phylogenetic distances to the nurse plant. Nurses established steeper N source-sink gradients with distant relatives, increasing 15 N transfer toward these species. Nutrient sharing may provide long-term benefits to facilitated plants and may be an overlooked mechanism maintaining coexistence and increasing the phylogenetic diversity of plant communities.

Metal(loid) allocation and nutrient retranslocation in Pinus halepensis trees growing on semiarid mine tailings.

The goal of this study was to evaluate internal metal(loid) cycling and the risk of metal(loid) accumulation in litter from Pinus halepensis trees growing at a mine tailing disposal site in semiarid Southeast Spain. Internal nutrient retranslocation was also evaluated in order to gain insight into the ability of pine trees to cope with the low-fertility soil conditions at the tailings. We measured metal(loid) concentrations in the foliage (young and old needles), woody stems and fresh leaf litter of pine trees growing on tailings. The nutrient status and stable isotope composition of pine foliage (δ(13)C, δ(15)N, δ(18)O as indicators of photosynthesis and water use efficiency) were also analyzed. Tailing soil properties in vegetation patches and in adjacent bare soil patches were characterized as well. Significant amounts of metal(loid)s such us Cd, Cu, Pb and Sb were immobilized in the woody stems of Pinus halepensis trees growing on tailings. Leaf litterfall showed high concentrations of As, Cd, Sb, Pb and Zn, which thereby return to the soil. However, water extractable metal(loid) concentrations in tailing soils were similar between vegetation patches (mineral soil under the litter layer) and bare soil patches. The pines growing on mine tailings showed very low foliar P concentrations in all leaf age classes, which suggests severe P deficiency. Young (current year) needles showed lower accumulation of metal(loid)s, higher nutrient concentrations (P and K), and higher water use efficiency (as indicated by and δ(13)C and δ(18)O data) than older needles. Substantial nutrient resorption occurred before leaf litterfall, with 46% retranslocation efficiency for P and 89% for K. In conclusion, phytostabilization of semiarid mine tailings with Pinus halepensis is feasible but would require careful monitoring of the trace elements released from litterfall, in order to assess the long term risk of metal(loid) transfer to the food chain.

Usefulness of pioneer vegetation for the phytomanagement of metal(loid)s enriched tailings: grasses vs. shrubs vs. trees.

The goal of this work was to assess the selection of the most suitable combination of plant species for the phytomanagement of mine tailings, by comparing among different plant life-forms (grasses, shrubs and trees). A comparison on induced rhizosphere changes generated by four plant species (the grass Piptatherum miliaceum, the shrub Helichrysum decumbens, and the trees, Pinus halepensis and Tetraclinis articulata) and high density vegetation patches (fertility islands) at a mine tailing located at Southeast Spain and the description of their physiological status employing stable isotopes analyses were carried out. The edaphic niches for plant growth were determined by salinity, organic matter and total soil nitrogen while metal(loid)s concentrations played a minor role. Induced changes in plant rhizospheres had a significant impact in soil microbiology. While grasses and shrubs may play an important role in primary ecological succession, trees seem to be the key to the development of fertility islands. The low δ(15)N values (-8.00‰) in P. halepensis needles may reflect higher ectomycorrhizal dependence. Large differences in leaf δ(18)O among the plant species indicated contrasting and complementary water acquisition strategies. Leaf δ(13)C values (-27.6‰) suggested that T. articulata had higher water use efficiency than the rest of species (-29.9‰). The implement of a diverse set of plant species with contrasting life forms for revegetating tailings may result in a more efficient employment of water resources and a higher biodiversity not only in relation to flora but soil microbiology too.

Isotopes reveal contrasting water use strategies among coexisting plant species in a Mediterranean ecosystem.

Variation in the stable carbon and oxygen isotope composition (δ13C, Δ18O) of co-occurring plant species may reflect the functional diversity of water use strategies present in natural plant communities. We investigated the patterns of water use among 10 coexisting plant species representing diverse taxonomic groups and life forms in semiarid southeast Spain by measuring their leaf δ13C and Δ18O, the oxygen isotope ratio of stem water and leaf gas exchange rates. Across species, Δ18O was tightly negatively correlated with stomatal conductance (gs), whereas δ13C was positively correlated with intrinsic water use efficiency (WUEi). Broad interspecific variation in Δ18O, δ13C and WUEi was largely determined by differences in gs, as indicated by a strong positive correlation between leaf δ13C and Δ18O across species The 10 co-occurring species segregated along a continuous ecophysiological gradient defined by their leaf δ13C and Δ18O, thus revealing a wide spectrum of stomatal regulation intensity and contrasting water use strategies ranging from 'profligate/opportunistic' (high gs, low WUEi) to 'conservative' (low gs, high WUEi). Coexisting species maintained their relative isotopic rankings in 2 yr with contrasting rainfall, suggesting the existence of species-specific 'isotopic niches' that reflect ecophysiological niche segregation in dryland plant communities.

Stand structure modulates the long-term vulnerability of Pinus halepensis to climatic drought in a semiarid Mediterranean ecosystem.

We investigated whether stand structure modulates the long-term physiological performance and growth of Pinus halepensis Mill. in a semiarid Mediterranean ecosystem. Tree radial growth and carbon and oxygen stable isotope composition of latewood (δ(13)C(LW) and δ(18)O(LW), respectively) from 1967 to 2007 were measured in P. halepensis trees from two sharply contrasting stand types: open woodlands with widely scattered trees versus dense afforested stands. In both stand types, tree radial growth, δ(13)C(LW) and δ(18)O(LW) were strongly correlated with annual rainfall, thus indicating that tree performance in this semiarid environment is largely determined by inter-annual changes in water availability. However, trees in dense afforested stands showed consistently higher δ(18)O(LW) and similar δ(13)C(LW) values compared with those in neighbouring open woodlands, indicating lower stomatal conductance and photosynthesis rates in the former, but little difference in water use efficiency between stand types. Trees in dense afforested stands were more water stressed and showed lower radial growth, overall suggesting greater vulnerability to drought and climate aridification compared with trees in open woodlands. In this semiarid ecosystem, the negative impacts of intense inter-tree competition for water on P. halepensis performance clearly outweigh potential benefits derived from enhanced infiltration and reduced run-off losses in dense afforested stands.

No tillage affects the phosphorus status, isotopic composition and crop yield of Phaseolus vulgaris in a rain-fed farming system.

BACKGROUND: Conservation tillage promotes the accretion of soil organic matter and often leads to improved soil fertility and moisture availability. However, few studies have looked at the physiological response of crop plants to different tillage practices. It was therefore hypothesised that measuring the nutrient concentrations and stable isotope composition (δ(13)C, δ(18)O, δ(15)N) of shoots could help evaluate the physiological response of common bean (Phaseolus vulgaris L.) to different tillage treatments (no tillage (NT) and mouldboard ploughing (MP)) in a rain-fed farming system in northern Mexico. RESULTS: NT significantly enhanced shoot phosphorus concentration in bean plants. Tillage exerted a negative effect on the extent of root colonisation (%) by arbuscular mycorrhizal fungi (AMF). Lower shoot δ(18)O but unchanged δ(13)C values in plants from the NT system suggest enhanced stomatal conductance but also enhanced photosynthetic rate, which overall resulted in unchanged water use efficiency. Bean plants in the NT system showed lower shoot δ(15)N values, which suggests that a larger proportion of total plant nitrogen was obtained through atmospheric nitrogen fixation in this treatment. CONCLUSION: Greater diversity of AMF soil communities and heavier colonisation of roots by AMF in the NT compared with the MP system appeared to contribute to improved crop nutrition, water relations and yield in this rain-fed agroecosystem.

Efflux of hydraulically lifted water from mycorrhizal fungal hyphae during imposed drought.

Apart from improving plant and soil water status during drought, it has been suggested that hydraulic lift (HL) could enhance plant nutrient capture through the flow of mineral nutrients directly from the soil to plant roots, or by maintaining the functioning of mycorrhizal fungi. We evaluated the extent to which the diel cycle of water availability created by HL covaries with the efflux of HL water from the tips of extramatrical (external) mycorrhizal hyphae, and the possible effects on biogeochemical processes. Phenotypic mycorrhizal fungal variables, such as total and live hyphal lengths, were positively correlated with HL efflux from hyphae, soil water potential (dawn), and plant response variables (foliar (15)N). The efflux of HL water from hyphae was also correlated with bacterial abundance and soil enzyme activity (P), and the moistening of soil organic matter. Such findings indicate that the efflux of HL water from the external mycorrhizal mycelia may be a complementary explanation for plant nutrient acquisition and survival during drought.

Common mycorrhizal networks provide a potential pathway for the transfer of hydraulically lifted water between plants.

Plant roots may be linked by shared or common mycorrhizal networks (CMNs) that constitute pathways for the transfer of resources among plants. The potential for water transfer by such networks was examined by manipulating CMNs independently of plant roots in order to isolate the role(s) of ectomycorrhizal (EM) and arbuscular mycorrhizal fungal (AMF) networks in the plant water balance during drought (soil water potential -5.9 MPa). Fluorescent tracer dyes and deuterium-enriched water were used to follow the pathways of water transfer from coastal live oak seedlings (Quercus agrifolia Nee; colonized by EM and AMF) conducting hydraulic lift (HL) into the roots of water-stressed seedlings connected only by EM (Q. agrifolia) or AMF networks (Q. agrifolia, Eriogonum fasciculatum Benth., Salvia mellifera Greene, Keckiella antirrhinoides Benth). When connected to donor plants by hyphal linkages, deuterium was detected in the transpiration flux of receiver oak plants, and dye-labelled extraradical hyphae, rhizomorphs, mantles, and Hartig nets were observed in receiver EM oak roots, and in AMF hyphae of Salvia. Hyphal labelling was scarce in Eriogonum and Keckiella since these species are less dependent on AMF. The observed patterns of dye distribution also indicated that only a small percentage of mycorrhizal roots and extraradical hyphae were involved with water transfer among plants. Our results suggest that the movement of water by CMNs is potentially important to plant survival during drought, and that the functional ecophysiological traits of individual mycorrhizal fungi may be a component of this mechanism.