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Chilling restrains the distribution of mangroves. We tested whether foliar phosphorus (P) fractions and gene expression are associated with cold tolerance in mangrove species. We exposed seedlings of six mangrove populations from different latitudes to favorable, chilling and recovery treatments, and measured their foliar P concentrations and fractions, photochemistry, nighttime respiration, and gene expression. A Kandelia obovata (KO; 26.45°N) population completely and a Bruguiera gymnorhiza (Guangxi) (BGG; 21.50°N) population partially (30%) survived chilling. Avicennia marina (24.29°N), and other B. gymnorhiza (26.66°N, 24.40°N, and 19.62°N) populations died after chilling. Photosystems of KO and photosystem I of BGG were least injured. During chilling, leaf P fractions, except nucleic acid P in three populations, declined and photoinhibition and nighttime respiration increased in all populations, with the greatest impact in B. gymnorhiza. Leaf nucleic acid P was positively correlated with photochemical efficiency during recovery and nighttime respiration across populations for each treatment. Relatively high concentrations of nucleic acid P and metabolite P were associated with stronger chilling tolerance in KO. Bruguiera gymnorhiza exhibited relatively low concentrations of organic P in favorable and chilling conditions, but its partially survived population showed stronger compensation in nucleic acid P and Pi concentrations and gene expression during recovery.

Background and Aims Initiation of cluster roots in white lupin (Lupinus albus) under phosphorus (P) deficiency requires auxin signalling, whereas flavonoids inhibit auxin transport. However, little information is available about the interactions between P deficiency and flavonoids in terms of cluster-root formation in white lupin. Methods Hydroponic and aeroponic systems were used to investigate the role of flavonoids in cluster-root formation, with or without 75 μm P supply. Key Results Phosphorus-deficiency-induced flavonoid accumulation in cluster roots depended on developmental stage, based on in situ determination of fluorescence of flavonoids and flavonoid concentration. LaCHS8, which codes for a chalcone synthase isoform, was highly expressed in cluster roots, and silencing LaCHS8 reduced flavonoid production and rootlet density. Exogenous flavonoids suppressed cluster-root formation. Tissue-specific distribution of flavonoids in roots was altered by P deficiency, suggesting that P deficiency induced flavonoid accumulation, thus fine-tuning the effect of flavonoids on cluster-root formation. Furthermore, naringenin inhibited expression of an auxin-responsive DR5:GUS marker, suggesting an interaction of flavonoids and auxin in regulating cluster-root formation. Conclusions Phosphorus deficiency triggered cluster-root formation through the regulation of flavonoid distribution, which fine-tuned an auxin response in the early stages of cluster-root development. These findings provide valuable insights into the mechanisms of cluster-root formation under P deficiency.

Low availability of phosphorus (P) is a key limiting factor for the growth of many crops. Selenium (Se) is a nutrient for humans that is acquired predominantly from plants. Localised P and Se supply may affect P- and Se-uptake efficiency. Our aim was to examine the mechanisms of alfalfa (Medicago sativa L.) to acquire P and Se when the elements are heterogeneously or homogeneously distributed in soil, and how P and Se supply affect plant growth and uptake of P and Se. We conducted a split-root experiment growing alfalfa in a loess soil with two distribution patterns (i.e. heterogeneous and homogeneous) of P and Se. The application rates of P (KH2PO4) and Se (Na2SeO3) were 0 and 20 mg P kg−1, and 0 and 1 mg Se kg−1, respectively. Our results showed that plants absorbed more Se when both P and Se were supplied homogeneously than when supplied heterogeneously. Supplying Se had a positive effect on plant P content. Localised P supply resulted in the exudation of more carboxylates by roots than homogeneous P supply did. Soil microbial biomass P was significantly greater when P was supplied homogeneously. Shoot-to-root translocation of Se had a positive effect on P-uptake efficiency. These results indicated that, compared with homogeneous P supply, localised P supply promoted P and Se uptake by increasing the amount of rhizosheath carboxylates and weakening the competition between roots and microbes. Translocation of Se within plant organs was promoted by the application of P, thus enhancing the P-uptake efficiency of alfalfa.

Carboxylate release by roots has been considered a strategy for mobilisation and acquisition of phosphorus (P). However, recently, it was argued that carboxylate release may be a way to discharge surplus carbon produced under conditions that limit plant growth. Plant P status may not be the main factor driving carboxylate release by roots. Instead, plant nitrogen (N) status and/or N:P ratio of the soil or plant may play a more important role in enhancing carboxylate release. A greenhouse pot experiment was performed to grow alfalfa in a P-deficient soil, supplied with two rates of P (0 and 20 mg kg−1) in combination with four forms of nitrogen (N) at five rates (0, 25, 50, 75, and 100 mg kg−1), to explore the effects of P rate, N form, N rate, and their interactions on plant growth, P and N status, and carboxylate release, and to determine the factors driving carboxylate release. Nitrogen addition weakened the positive effect of P addition on plant growth, and increased plant N and P concentrations; P addition increased plant P concentration, but weakened the effect of N addition on plant N concentration. The amount of tartrate increased dramatically with increasing N rate, which decreased shoot growth, depending on N form. At high P supply, tartrate exudation correlated negatively with shoot biomass. Nitrogen addition to P-deficient alfalfa decreased shoot growth and enhanced the release of tartrate, likely to discharge surplus carbon; and the effects varied with N form.

Invasive plants rapidly spread in habitats with low soil phosphorus (P) availability and have triggered a sharp decline in the diversity of native species. However, no studies have explored how widespread invasive species acclimate to low soil P availability via changing foliar P fractions, especially under elevated atmospheric CO2 concentrations ([CO2 ]) and nitrogen (N) deposition. Here, an open-top chamber experiment was conducted to explore the effect of nutrient addition and elevated [CO2 ] on leaf traits and foliar functional P fractions (i.e. Pi, metabolite P, lipid P, nucleic acid P, and residual P) of two aggressive invasive species (Mikania micranatha and Chromolaena odorata). We found that foliar N/P ratios were more than 20, and P addition significantly increased plant biomass. Both results indicated P-limited plant growth at our studied site. Elevated [CO2 ], N and N+P addition greatly increased plant biomass, photosynthetic rates, and photosynthetic P-use efficiency (PPUE) in invasive species, but PPUE decreased with increasing P addition. Nitrogen addition slightly decreased the concentration of leaf total P, decreased foliar residual P, but increased metabolite P concentrations in invasive species. Similar changes in foliar P fractions were found under N+P addition. Phosphorus addition increased foliar P concentrations, which was strongly correlated with an increase in metabolite P concentrations in invasive species. Elevated [CO2 ] alleviated these effects and increased PPUE. The present results suggest that future elevated [CO2 ] and N deposition allow the invasive species to acclimate to low soil P availability and support their successful invasion by greatly reducing P allocation to non-metabolite foliar P fractions (i.e. nucleic acid P and residual P) to meet their demand of metabolite P for photosynthesis and exhibit a high PPUE. This article is protected by copyright. All rights reserved.

Estimation of leaf nutrient composition of dominant plant species from contrasting habitats (i.e., karst and nonkarst forests) provides an opportunity to understand how plants are adapted to karst habitats from the perspective of leaf traits. Here, we measured leaf traits—specific leaf area (SLA), concentrations of total carbon ([TC]), nitrogen ([TN]), phosphorus ([TP]), calcium ([Ca]), magnesium ([Mg]), manganese ([Mn]), minerals ([Min]), soluble sugars, soluble phenolics, lipids, and organic acids ([OA])—and calculated water‐use efficiency (WUE), construction costs (CC), and N/P ratios, and searched for correlations between these traits of 18 abundant plant species in karst and nonkarst forests in southwestern China. Variation in leaf traits within and across the abundant species was both divergent and convergent. Leaf [TC], [Ca], [Min], [OA], and CC were habitat‐dependent, while the others were not habitat‐ but species‐specific. The correlations among [TN], [TP], SLA, [TC], CC, [Min], WUE, [OA], and CC were habitat‐independent, and inherently associated with plant growth and carbon allocation; those between [CC] and [Lip], between [Ca] and [Mg], and between [Mg] and [WUE] were habitat‐dependent. Habitat significantly affected leaf [Ca] and thus indirectly affected leaf [OA], [Min], and CC. Our results indicate that plants may regulate leaf [Ca] to moderate levels via adjusting leaf [OA] under both high and low soil Ca availability, and offer new insights into the abundance of common plant species in contrasting habitats., Variation in leaf traits within and across the common tree species was both divergent and convergent, and the causal effects of leaf traits were habitat‐dependent or habitat‐independent. Habitat had significant effects on leaf [Ca] and thus indirectly affected leaf [OA], [Min], and CC. Our study revealed leaf [OA] plays an important role in adjusting leaf concentrations and enhances adaptability of dominant plant species in karst habitats, offering new insights into the abundance of common tree species in contrasting habitats, which would be useful in karst forest restoration.

Plants deploying a phosphorus (P)-mobilising strategy via carboxylate release have relatively high leaf manganese concentrations ([Mn]). Thus, leaf [Mn] is a proxy for the amount of rhizosheath carboxylates. Whether the concentrations of other leaf micronutrient, such as iron ([Fe]), zinc ([Zn]) and copper ([Cu]), show a similar signal for rhizosheath carboxylates is unclear. We grew a large number of chickpea genotypes in two glasshouse studies with different growth media, P sources and P levels. Seven weeks after sowing, we determined concentrations of micronutrients in mature leaves, and the quantity and composition of rhizosheath carboxylates. For 100 genotypes grown in river sand with low P supply, leaf [Fe] (R2 = 0.36) and [Zn] (R2 = 0.22), like leaf [Mn] (R2 = 0.38), were positively correlated with the total amount of rhizosheath carboxylates. For 20 genotypes grown in a soil mixture, leaf [Fe], [Zn], [Cu] and [Mn] showed positive correlations with total rhizosheath carboxylates that were stronger under moderately low P (R2 = 0.59, 0.59, 0.54, 0.72) than severely low P (R2 = 0.39, 0.28, 0.20, 0.36) or sufficient P (R2 = 0.36, 0.00, 0.01, 0.50) supply. Malonate was the predominant carboxylate in the rhizosheath and was significantly correlated with leaf micronutrient concentrations in both experiments. In addition to leaf [Mn], leaf [Fe] and [Zn] can be used as alternative and easily measurable proxies for belowground carboxylate-releasing processes in chickpea under low-P supply, particularly on moderately low-P soils.

Seeds of crops contain large amounts of phosphorus (P), mostly as phytate. To lower seed phytate concentration is the ultimate breeding aim and highly desirable as the benefit of humans and many animals, given seed size is not sacrificed, i.e. leading to reduced seed P content. This study tested whether low seed P content affects early chickpea growth, and whether P supply during early growth compensates for early growth from seeds with low seed P content. Twenty-three chickpea genotypes with contrasting seed P content were selected for three treatments that were compared with a control: 1) seedlings supplied with a small amount of starter P (P10), 2) one cotyledon removed to reduce P content (Remove 1), and 3) both cotyledons removed to further reduce P content (Remove 2). Genotypes with higher seed P content had more vigorous early growth than genotypes with lower seed P, as did intact seedlings compared with those cotyledons removed. Application of P fertiliser significantly enhanced early growth. Low seed P genotypes had a more economic strategy, with a greater specific leaf area and specific root length, than genotypes with medium or high seed P content. Our results provided useful information for breeders that breeding chickpea genotypes with reduced seed P, does not affect germination and seedling vigour, provided starter P is supplied at the early development stage.

Residues of antibiotics such as oxytetracycline (OTC) in soil can affect microbial compositions and activities, thus affecting soil P availability, and consequently plant P uptake and growth. A pot experiment was performed to grow alfalfa in a loess soil with different doses of P (0, 25, 50, and 100 mg kg−1) and OTC (0, 25, 50, and 100 mg kg−1). Plant dry mass, shoot and root P concentrations, bulk soil and rhizosheath pH, rhizosheath carboxylates, and bulk soil alkaline phosphatase activity were determined. Shoot dry mass and root dry mass increased with increasing P dose, while shoot dry mass decreased with increasing OTC dose, especially at lower P doses (0 and 25 mg kg−1). Addition of OTC slightly reduced P concentrations in shoots and roots, but did not reduce plant P content consistently. Increasing OTC dose significantly reduced bulk soil alkaline phosphatase activity at 0P and strongly reduced rhizosheath tartrate amount at all P doses. The effects of OTC on plant growth and P uptake depended on both OTC and P doses in soil. High OTC dose had negative effects on shoot P uptake and growth, especially at lower P doses, while it had a positive effect on root growth at higher P doses.

Zinc (Zn) and phosphorus (P) often interact negatively with each other in soil-plant systems. We investigated the effects of P-Zn interaction on Zn and P accumulation and partitioning in alfalfa. Plants were grown in a calcareous soil supplied with different rates of Zn (0, 200, and 800 mg kg−1) and P (0, 20, and 80 mg kg−1). Plant dry mass, Zn and P concentrations in shoots and roots, bulk soil and rhizosheath pH, rhizosheath carboxylates, and DTPA-extractable Zn concentration in the bulk soil were determined. Phosphorus-Zn interaction significantly affected DTPA-extractable Zn concentration, plant dry mass, accumulation of Zn and P, and partitioning of Zn in alfalfa, but did not affect rhizosheath pH or the amounts of rhizosheath carboxylates. Increasing P rate promoted plant growth at all soil Zn rates and might enhance the plants’ capacity to cope with excessive Zn; it resulted in a lower rhizosheath pH, which likely contributed to greater Zn and P uptake. Zinc deficiency enhanced exudation of citrate, malonate and malate, while the release of tartrate was related with P deficiency. There are strong P-Zn interactions in calcareous soil-plant system, such interactions significantly affect Zn bioavailability, plant growth, accumulation of Zn and P, and partitioning of Zn in alfalfa. Rational P fertilization should be considered for efficient Zn biofortification on Zn-deficient soils and phytoremediation of Zn-contaminated soils.

Nutrient-poor ecosystems globally exhibit high plant diversity. One mechanism enabling the co-existence of species in such ecosystems is facilitation among plants with contrasting nutrient-acquisition strategies. The ecophysiological processes underlying these interactions remain poorly understood. We hypothesized that root positioning plays a role between sympatric species in nutrient-poor vegetation. We investigated how the growth traits of the focal mycorrhizal non-cluster-rooted Hibbertia racemosa change when grown in proximity of non-mycorrhizal Banksia attenuata, which produces cluster roots that increase nutrient availability, compared with growth with conspecifics. Focal plants were placed in the centre of rhizoboxes, and biomass allocation, root system architecture, specific root length (SRL), and leaf nutrient concentration were assessed. When grown with B. attenuata, focal plants decreased root investment, increased root growth towards B. attenuata, and positioned their roots near B. attenuata cluster roots. SRL was greater, and the degree of localized root investment correlated positively with B. attenuata cluster-root biomass. Total nutrient contents in the focal individuals were greater when grown with B. attenuata. Focal plants directed their root growth towards the putatively facilitating neighbour's cluster roots, modifying root traits and investment. Preferential root positioning and root morphological traits play important roles in positive plant-plant interactions.

We investigated the effects of water stress under low phosphorus (P) supply on P-acquisition by chickpea, and identified a genotype with faster relative growth and P-acquisition rates. We grew four genotypes in pots filled with a sand and soil mixture with a low P availability in a glasshouse. Plants were either well-watered or water-stressed, imposed at the reproductive stage. Plants were harvested when water-stressed plants fully closed their stomata. For all four genotypes, water stress reduced shoot and root growth, root mass ratio, and shoot P content, while it increased specific root length (except in ICC 456), water-use efficiency and the amount of rhizosheath carboxylates per gram root dry weight. A faster relative shoot P-acquisition rate in ICC 2884 was associated with a greater specific root length, a smaller mean root diameter and a greater increase in the amount of rhizosheath carboxylates in response to water stress under low P supply. Interestingly, under water stress ICC 2884 also maintained a similar physiological P-use efficiency to that of the well-watered plants. ICC 2884 is recommended as a parental genotype in chickpea breeding programs to develop cultivars for low-P and terminal drought environments.

Root-released carboxylates enhance the availability of manganese (Mn), which enters roots through transporters with low substrate specificity. Leaf Mn concentration ([Mn]) has been proposed as a signature for phosphorus (P)-mobilising carboxylates in the rhizosphere. Here we test whether leaf [Mn] provides a signature for root functional types related to P acquisition. Across 727 species at 66 sites in Australia and New Zealand, we measured leaf [Mn] as related to root functional type, while also considering soil and climate variables. To further assess the specific situations under which leaf [Mn] is a suitable proxy for rhizosphere carboxylate concentration, we studied leaf [Mn] along a strong gradient in water availability on one representative site. In addition, we focused on two systems where a species produced unexpected results. Controlling for background site-specific variation in leaf [Mn] with soil pH and mean annual precipitation, we established that mycorrhizal species have significantly lower leaf [Mn] than non-mycorrhizal species with carboxylate-releasing root structures, e.g., cluster roots. In exception to the general tendency, leaf [Mn] did not provide information about root functional types under seasonally waterlogged conditions, which increase iron availability and thereby interfere with Mn-uptake capacity. Two further exceptions were scrutinised, leading to the conclusion that they were ‘anomalous’ in not functioning like typical species in their families, as expected according to the literature. Leaf [Mn] variation provides considerable insights on differences in belowground functioning among co-occurring species. Using this approach, we concluded that, within typical mycorrhizal families, some species actually depend on a carboxylate-releasing P-mobilising strategy. Likewise, within families that are known to produce carboxylate-releasing cluster roots, some do not produce functional cluster roots when mature. An analysis of leaf [Mn] can alert us to such ‘anomalous’ species.

Reducing nitrogen (N) input can improve crop productivity in cereal-legume intercrops, but the impact on phosphorus (P) acquisition is unclear. A 10-year (2009–2018) field experiment was conducted to quantify how P acquisition by sugarcane (Saccharum officinarum) was affected by intercropping with soybean (Glycine max (L.) Merrill at 1:1 and 1:2) with two N inputs (300 kg ha−1 [reduced], 525 kg ha−1 [conventional]). Nitrogen was supplied only to the sugarcane crop, and soybean received no N. There was a significantly higher land-equivalent ratio of sugarcane-soybean intercropping than of the sole cropping, and the intercropping advantage was more pronounced under reduced N input which can be associated with high degree of complementary N use. Furthermore, soybean intercropping with reduced N input stimulated acid phosphomonoesterase activity and depleted organic P in the rhizosphere of sugarcane, resulting in increased sugarcane stem P concentration and system P-use efficiency. The interspecific facilitation of P acquisition could be associated with the increased symbiotic N2 fixation in soybean, soil microbial biomass and activity under reduced N input. In conclusion, soybean intercropping with reduced N input to sugarcane enhanced rhizosphere enzymatic organic P transformation and sugarcane P acquisition, which may contribute to maintaining a sustainable sugarcane production under low N supply. The findings advance our understanding of interactions between N and P cycling and provide new evidence for the value of cereal-legume intercrops in reducing fertilizer input.

Plant growth is often limited by low soil phosphorus (P) availability, soil nitrogen (N) availability may affect plants’ responses to P supply. We studied the growth and physiological responses of alfalfa to soil P supply at different N levels. Alfalfa (Medicago sativa L.) plants were grown in an alkaline soil supplied with different levels of P (0, 5, and 20 mg kg−1) as monopotassium phosphate, and N (50 and 100 mg kg−1) as ammonium nitrate. Plant biomass and P concentrations always showed positive responses to P addition but not to N addition, nodulation was inhibited by lower P supply and higher N supply. Roots released more phosphatase and carboxylates, mainly tartrate, into the rhizosheath at lower soil P supply and higher N supply. Roots always acidified the rhizosheath, but rhizosheath pH did not vary considerably among treatments. This study demonstrates the release of tartrate as a major carboxylate as affected by soil P supply and N supply, and highlights the importance of investigating plant adaptive strategies for P acquisition from soil with different N availability for proper application of P and N fertilizers.

Soil alkalinity imposes important limitations to lupin productivity; however, little attention has been paid to investigate the effects of soil alkalinity on plant growth and development. Many lupins are sensitive to alkaline soils, butLupinus albusmaterial from Egypt was found to have tolerance to limed soils. The aim of this study was to compare the growth response of two cultivars ofL.albusL. – an Egyptian cultivar, P27734, and an Australian cultivar, Kiev Mutant, to different soil pH levels and to understand the physiological mechanisms underlying agronomic alkalinity tolerance of P27734. Plants were grown under three pH levels (5.1, 6.7, and 7.8) in a temperature-controlled glasshouse. For both cultivars, the greatest dry mass production and carboxylate exudation from roots were observed at alkaline pH. The better performance of the Egyptian cultivar at high pH was entirely accounted for by its greater seed weight. From a physiological perspective, the Australian cultivar was as alkaline-tolerant as the Egyptian cultivar. These findings highlight the agronomic importance of seed weight for sowing, and both cultivars can be used in alkaline soils.

Phosphorus (P) is essential for productivity of alpine grassland ecosystems, which are sensitive to global warming. We tested the hypotheses that (1) mobilized 'calcium-bound inorganic P' (Ca-Pi ) is a major source of plant-available P in alpine meadows with alkaline soils after long-term warming, (2) mobilization of Ca-Pi is linked to effective plant carboxylate-releasing P-acquisition strategies under warming, and (3) the mobilization is also related to plant nitrogen (N)-acquisition. We conducted an 8-year warming experiment in an alpine meadow (4635 m above sea level) on the Qinghai-Tibetan Plateau. A significant increase in P concentration in both aboveground and belowground biomass indicates an increased mobilization and assimilation of P by plants under warming. We observed a significant decrease in Ca-Pi , no change in moderately-labile organic P, and an increase in highly resistant organic P after warming. There was no increase in phosphatase activities. Our results indicate that Ca-Pi , rather than organic P was the major source of plant-available P for alpine meadows under warming. Higher leaf manganese concentrations of sedges and forbs after warming indicate that carboxylates released by these plants are a key mechanism of Ca-Pi mobilization. The insignificant increase in Rhizobiales after warming and the very small cover of legumes show a minor role of N-acquisition strategies in solubilizing phosphate. The insignificant change in relative abundance of mycorrhizal fungi and bacteria related to P cycling after warming shows a small contribution of microorganisms to Ca-Pi mobilization. The significant increase in leaf N and P concentrations and N:P ratio of grasses and no change in sedge leaf N:P ratio reflect distinct responses of plant nutrient status to warming due to differences in P-acquisition strategies. We highlight the important effects of belowground P-acquisition strategies, especially plant carboxylate-releasing P-acquisition strategies on responses of plants to global changes in alpine meadows.

Aims Phosphorus (P) fertilizers are often considered an important source of cadmium (Cd) in crop plants. However, increased plant Cd concentrations are not strictly related to the Cd content of P fertilizers. Considering this, we hypothesized that, alternatively, reduction of arbuscular mycorrhizal colonization by P fertilization enhances Cd accumulation in plants. Methods Wheat and canola as mycorrhizal and non-mycorrhizal species, respectively, were grown under greenhouse conditions with and without soil sterilization. Phosphorus fertilizers with 0.09, 5, and 28 mg Cd kg(-1) were applied at different rates with varied zinc (Zn) fertilization. Results In wheat, all three P fertilizers markedly increased shoot and grain Cd concentrations as P supply was increased, irrespective of the Cd concentration in the fertilizers. These increases were pronounced with soil sterilization or at low zinc supply. Adding mycorrhizal fungi to sterilized soil substantially decreased shoot Cd concentrations. We found a strong negative relationship in wheat between mycorrhizal colonization and shoot Cd concentration, for both high- and low-Cd fertilizers. In contrast to wheat, shoot Cd concentrations in canola showed virtually no response to P supply or soil sterilization. High Zn application also reduced plant Cd concentrations, especially at high P rates. Conclusion Our findings demonstrate the critical importance of mycorrhizal colonization in reducing Cd accumulation in wheat, and suggest that factors suppressing root mycorrhizal activity including P fertilization, will increase Cd uptake in mycorrhizal plants.

Aims Carboxylate release by roots has been considered a strategy for mobilization and acquisition of phosphorus (P). However, recently, it was argued that carboxylate release may be a way to discharge surplus carbon produced under conditions that limit plant growth. Plant P status may not be the main factor driving carboxylate release by roots. Instead, plant nitrogen (N) status and/or N:P ratio of the soil or plant may play a more important role in enhancing carboxylate release. Methods A greenhouse pot experiment was performed to grow alfalfa in a P-deficient soil, supplied with two rates of P (0 and 20 mg kg− 1) in combination with four forms of nitrogen (N) at five rates (0, 25, 50, 75, and 100 mg kg− 1), to explore the effects of P rate, N form, N rate, and their interactions on plant growth, P and N status, and carboxylate release, and to determine the factors driving carboxylate release. Results Nitrogen addition weakened the positive effect of P addition on plant growth, and increased plant N ([N]) and P concentrations ([P]); P addition increased plant [P], but weakened the effect of N addition on plant [N]. The amount of tartrate increased dramatically with increasing N rate, which decreased shoot growth, depending on N form. At high P supply, tartrate exudation correlated negatively with shoot biomass. Conclusions Nitrogen addition to P-deficient alfalfa decreased shoot growth and enhanced the release of tartrate, likely to discharge surplus carbon; and the effects varied with N form.

Background and Aims Soil phosphorus (P) deficiency and salinity are constraints to crop productivity in arid and semiarid regions. Salinity may weaken the effect of P fertilization on plant growth. We investigated the interactive effects of soil P availability and salinity on plant growth, P nutrition and salt tolerance of two alfalfa (Medicago sativa) cultivars. Methods A pot experiment was carried out to grow two cultivars of alfalfa in a loess soil under a combination of different rates of added P (0, 40, 80 and 160 mg P kg−1 soil as monopotassium phosphate) and sodium chloride (0, 0.4, 0.8 and 1.6 g NaCl kg−1 soil). Plant biomass, concentrations of P ([P]), sodium ([Na]) and potassium ([K]) were determined, and rhizosheath carboxylates were analysed. Key Results There were significant interactions between soil P availability and salinity on some, but not all, of the parameters investigated, and interactions depended on cultivar. Plant growth and P uptake were enhanced by P fertilization, but inhibited by increased levels of salinity. Increasing the salinity resulted in decreased plant P-uptake efficiency and [K]/[Na]. Only soil P availability had a significant effect on the amount of tartrate in the rhizosheath of both cultivars. Conclusions Increased salinity aggravated P deficiency. Appropriate application of P fertilizers improved the salt tolerance of alfalfa and increased its productivity in saline soils.

Summary • The influence of soil P level and plant development on carboxylate concentrations in the rhizosphere of lateral roots of chickpea (Cicer arietinum) cultivars Heera and Tyson were studied to investigate the relationship between carboxylate exudation and P acquisition by chickpea. • Two chickpea cultivars were grown in soil supplied with 0–100 µg P g−1. Plants were harvested four times during plant development. In two other experiments, carboxylate concentrations were measured along root sections, using plants grown in soil or in hydroponics. • Carboxylate concentrations in the rhizosphere of lateral roots steadily increased with plant development for cv. Heera, but not for Tyson. Carboxylate concentrations increased with increasing distance from the root apex in soil. This increase might be due to accumulation, because an experiment with plants in hydroponics showed that older segments released at least as much carboxylates as younger segments. • Carboxylate concentrations in the rhizosphere of chickpea appear to be developmentally controlled. Unlike some other plant species, there is no simple correlation with plant or soil P status.

SUMMARY Two inbred lines of Plantago major L., differing in relative growth rate (RGR), were studied in order to elucidate physiological factors responsible for this difference. The slow growing inbred line of P. major ssp. major L. (line W9) had a lower specific leaf area (SLA) compared with that of the fast growing P. major ssp. pleiosperma Pilger (line A4). Photosynthetic activity per unit leaf area was higher than that of A4, but expressed per unit leaf dry weight W9 exhibited a lower activity than A4. Photosynthetic capacity (per unit leaf dry weight or nitrogen) was identical for both lines. The amount of chlorophyll a per unit dry weight or leaf area was higher for line W9. Differences in the chemical composition of the leaf dry weight of the two lines did not explain the lower SLA or lower photosynthetic activity per unit leaf dry weight of the slow growing line. The lower SLA of W9 was mainly caused by the lower water content per unit dry weight. A higher amount of cell wall material per unit leaf dry weight was found. We conclude that the lower SLA of W9 resulted in a greater shading within the leaf, a lower light absorption and lower photosynthetic rate per unit leaf dry weight. Consequently, this resulted in a lower RGR. The significance of these characters for growth under trampling and soil compaction is discussed.

Mine tailings are challenging substrates for ecological restoration, as the establishment of diverse native plant communities can be constrained by a range of edaphic factors. Thus, the ability to restore native vegetation communities will depend upon developing a clear evidence-base as to what types of species and communities are likely sustainably reinstated on such altered substrates. As global tailings production and the cumulative footprint of tailings storage facilities continue to grow, understanding the effect of edaphic filters on community establishment is foundational for developing effective restoration solutions for tailings. We standardised growth rate estimates derived from nine root and shoot parameters for plants grown in magnetite tailings and natural topsoil, using crops (eight species) to characterise previously identified plant responses and native plants (40 species) to understand the impact of edaphic conditions on the species pool available for restoration. The edaphic conditions of unweathered magnetite tailings select against the majority of native plant species and nutrient-acquisition guilds (approximately 75% of reference floristic biodiversity), with plant development on tailings compared with natural topsoil compromised in a number of variables in all but six species. Plant growth on tailings was limited by a lack of available nitrogen (N) and high alkalinity (pH >9), and seedling growth and development was positively associated with seed N concentration. Calcicole species and species from N2-fixing and cluster root-producing strategies performed better on tailings than calcifuge species and species without specialised nutrient-acquisition strategy or those reliant upon mycorrhizal associations. The return of plant communities native to highly weathered, acidic soils on magnetite tailings is likely unsuccessful, unless strategies to ameliorate substrate hostility through acidification of the soil profile and improving N availability are prioritised.

Silicon (Si) is widely recognized as an important regulator of the global carbon (C) cycle via its effect on diatom productivity in oceans and the weathering of silicate minerals on continents. Si is also a beneficial plant nutrient, improving resistance to herbivory and pathogens and mitigating the negative effects of several abiotic stresses, including nutrient limitation. However, changes in Si sources and cycling during long-term development of terrestrial ecosystems remain poorly understood. We studied Si in soils and plants along two 2-Ma coastal dune chronosequences in southwestern Australia (Jurien Bay and Guilderton). Soil development along these chronosequences includes carbonate leaching in Holocene soils, formation of secondary Si-bearing minerals in Mid-Pleistocene soils, followed by their loss via dissolution, to yield quartz-rich soils of Early-Pleistocene age. The chronosequences also exhibit an extreme gradient of soil fertility in terms of rock-derived nutrients, and shifts from nitrogen (N) to phosphorus (P) limitation of plant productivity as soils age. Along each chronosequence, we quantified the pools of reactive Si-bearing phases and plant-available Si in the soils, and physically extracted soil phytoliths (amorphous silica formed in plant tissues). We also quantified Si, macronutrients and total phenols in the most abundant plants growing along the best-studied of the two chronosequences (Jurien Bay). We found that plant-available Si was lowest in young and carbonate-rich soils, because carbonates weathering reduces the weathering of silicate minerals by consuming protons, and Si is strongly sorbed by secondary minerals in alkaline soils. Plant-available Si increased in intermediate-age soils during the formation of secondary minerals (kaolinite), and finally decreased in old, quartz-rich soils, due to continuous desilication. As pedogenic Si pools became depleted with increasing soil age, Si availability was increasingly determined by soil phytoliths. At Jurien Bay, foliar Si increased continuously as soils aged, in contrast with foliar macronutrients that declined markedly in strongly weathered soils. Finally, foliar phenol concentrations declined with increasing soil age and were negatively correlated with foliar Si at the community and individual species level, suggesting a tradeoff between these two leaf defense strategies. Our results highlight a nonlinear response of plant-available Si to long-term pedogenesis, with an increase during carbonate loss and a decrease in the silicates weathering domain. They also demonstrate that the retention of Si by plants during ecosystem retrogression sustains its terrestrial cycling by leveraging the high reactivity of soil phytoliths compared with soil-derived aluminosilicates. Moreover, the continuous increase of plant Si concentrations as rock-derived nutrients are depleted suggests important plant benefits associated with Si in P-impoverished environments. This is in line with the resource availability hypothesis, which predicts that plants adapted to infertile soils have high levels of anti-herbivore leaf defenses. In particular, old and P-depleted soils increased the relative expression of Si-based defenses, while young soils where plant productivity is limited by N promoted leaf phenol accumulation. Overall, our results demonstrate that long-term ecosystem and soil development strongly influence soil-plant Si dynamics, with cascading effects on plant ecology and global Si and C biogeochemistry.

Made available in DSpace on 2021-06-25T15:03:38Z (GMT). No. of bitstreams: 0 Previous issue date: 2021-03-01 Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) UWA Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG) Questions Environmental filters limit the set of potentially coexisting species in plant communities. Paradoxically, some of the world's most biodiverse communities are subjected to strong abiotic filters. We explored how environmental heterogeneity provides conditions for niche segregation in a harsh megadiverse ecosystem, focusing on fine-scale factors that drive seedling recruitment. Due to the environmental stress (mainly nutrient and water scarcity), we expected the prevalence of positive plant interactions over negative ones to contribute to species coexistence. Location Ironstone outcrop (canga) at Serra da Calcada, southeast Brazil. Methods We characterised microhabitats with one or neither of two dominant shrubs Mimosa calodendron (Fabaceae) and Lychnophora pinaster (Asteraceae), and assessed which micro-environmental attributes affect seedling survival, growth, and abundance, as well as adult richness in the ecosystem. Results Spatial heterogeneity in the community was mostly driven by differences in substrate properties, and plots with Mimosa calodendron and Lychnophora pinaster exhibited different biotic and abiotic conditions from those lacking these species. Microhabitats under both shrubs had greater adult richness, and those occupied by Mimosa calodendron had greater seedling abundance, suggesting positive effects of environmental conditions associated with these plants. Nevertheless, we identified two potential negative plant-plant interactions: larger crown area of the two dominant shrubs was associated with lower species richness, and greater understorey plant cover with lower seedling abundance. Over one year, 63% of the recorded seedlings survived, but measured attributes failed to predict survival. Seedling relative growth rates were species-specific and lower in microsites where the rocky substrate is fragmented. The most abundant seedling taxa occurred in association with specific microhabitats. Conclusions Our study suggested that fine-scale spatial heterogeneity determines the outcome of plant-plant interactions in a stressful ecosystem. Micro-environmental heterogeneity allows for the spatial segregation of species recruitment, thereby broadening the range of viable strategies within a harsh megadiverse ecosystem. Univ Fed Minas Gerais, Dept Bot, Av Antonio Carlos 6627, BR-31270110 Belo Horizonte, MG, Brazil Univ Western Australia, Sch Biol Sci, Perth, WA, Australia Univ Estadual Paulista, Dept Zool, Rio Claro, Brazil Univ Fed Minas Gerais, Dept Genet Ecol & Evolucao, Belo Horizonte, MG, Brazil Univ Estadual Paulista, Dept Zool, Rio Claro, Brazil CAPES: 88881.068071/2014-01 FAPESP: 2017/07954-0

Tailings are among the most challenging mined substrates for plant re-establishment, in particular because of a lack of soil-like structure and nitrogen. Potential pioneer plants are sometimes found in such disturbed and infertile sites. We present a group of pioneer species from the genus Maireana (Chenopodiaceae) that are promising candidates for the restoration of magnetite tailings. We found that these Maireana species did not rely on biologically fixed N from the atmosphere, but exhibited an exceptionally high leaf N-resorption efficiency (about 95%) during leaf senescence, at the same time effectively scavenging trace amount of N from the substrate, in part through rapid transpiration.

Phosphorus (P) deficiency induces cluster-root formation and carboxylate exudation in most Proteaceae. However, how external P supply regulates these root traits in Macadamia integrifolia remains unclear. Macadamia plants were grown hydroponically with seven P levels to characterize biomass allocation, cluster-root development, and exudation of carboxylates and acid phosphatases. Plant biomass increased with increasing P supply, peaking at 5 μM P, was the same at 5–25 μM P, and declined at 50–100 μM P. Leaf P concentration increased with increasing P supply, but shoot biomass was positively correlated with leaf P concentration up to 0.7–0.8 mg P g–1 dry weight (DW), and declined with further increasing leaf P concentration. The number of cluster roots declined with increasing P supply, with a critical value of leaf P concentration at 0.7–0.8 mg P g–1 DW. We found a similar trend for carboxylate release, with a critical value of leaf P concentration at 0.5 mg g–1 DW, but the activity of acid phosphatases showed a gradually-decreasing trend with increasing P supply. Our results suggest that leaf P concentration regulates the development and functioning of cluster roots, with a critical P concentration of 0.5–0.8 mg g–1, above which macadamia growth is inhibited.

Phytate-phosphorus (P) in food and feed is not efficiently utilized by humans and non-ruminant livestock, potentially contributing to high losses of P to the environment. Crops with high P-acquisition efficiency can access soil P effectively. It remains elusive whether crop genotypes with high P-acquisition efficiency can also have low seed phytate concentrations. A core collection of 256 soybean [Glycine max (L.) Merr.] genotypes from China with diverse genetic background were grown in the same environment and seeds were sampled to screen for seed phytate-P concentration. Some of these genotypes were also grown in a low-P soil in the glasshouse to measure root morphological and physiological traits related to P acquisition. Large genotypic variation was found in seed phytate-P concentration (0.69–5.49 mg P g–1 dry weight), total root length, root surface area, rhizosheath carboxylates, and acid phosphatase activity in rhizosheath soil. Geographically, seed phytate-P concentration was the highest for the genotypes from Hainan Province, whereas it was the lowest for the genotypes from Inner Mongolia. Seed phytate-P concentration showed no correlation with any desirable root traits associated with enhanced P acquisition. Two genotypes (Siliyuan and Diliuhuangdou-2) with both low phytate concentrations and highly desirable P-acquisition traits were identified. This is the first study to show that some soybean genotypes have extremely low seed phytate concentrations, combined with important root traits for efficient P acquisition, offering material for breeding genotypes with low seed phytate-P concentrations.

Plant growth is usually constrained by the availability of nutrients, water, or temperature, rather than photosynthetic carbon (C) fixation. Under these conditions leaf growth is curtailed more than C fixation, and the surplus photosynthates are exported from the leaf. In plants limited by nitrogen (N) or phosphorus (P), photosynthates are converted into sugars and secondary metabolites. Some surplus C is translocated to roots and released as root exudates or transferred to root-associated microorganisms. Surplus C is also produced under low moisture availability, low temperature, and high atmospheric CO2 concentrations, with similar below-ground effects. Many interactions among above- and below-ground ecosystem components can be parsimoniously explained by the production, distribution, and release of surplus C under conditions that limit plant growth. Non

Arachis hypogaea (peanut) is a globally important oilseed crop with high nutritional value. However, upon exposure to overnight chilling stress, it shows poor growth and seedling necrosis in many cultivation areas worldwide. Calcium (Ca2+) enhances chilling resistance in various plant species. We undertook a pot experiment to investigate the effects of exogenous Ca2+ and a calmodulin (CaM) inhibitor on growth and photosynthetic characteristics of peanut exposed to low night temperature (LNT) stress following warm sunny days. The LNT stress reduced growth, leaf extension, biomass accumulation, gas exchange rates, and photosynthetic electron transport rates. Following LNT stress, we observed larger starch grains and a concomitant increase in nonstructural carbohydrates and hydrogen peroxide (H2O2) concentrations. The LNT stress further induced photoinhibition and caused structural damage to the chloroplast grana. Exogenous Ca2+ enhanced plant growth following LNT stress, possibly by allowing continued export of carbohydrates from leaves. Foliar Ca2+ likely alleviated the nocturnal chilling-dependent feedback limitation on photosynthesis in the daytime by increasing sink demand. The foliar Ca2+ pretreatment protected the photosystems from photoinhibition by facilitating cyclic electron flow (CEF) and decreasing the proton gradient (ΔpH) across thylakoid membranes during LNT stress. Foliar application of a CaM inhibitor increased the negative impact of LNT stress on photosynthetic processes, confirming that Ca2+–CaM played an important role in alleviating photosynthetic inhibition due to the overnight chilling-dependent feedback.

Soil property impacts the efficacy of rhizobial inoculation and rhizosphere processes. However, very few comparative studies have been done to analyze the effects of rhizobial inoculants on plant nitrogen (N) and phosphorus (P) in different soils. The aim of the present study was, therefore, to evaluate N fixation and P acquisition of inoculated soybean (Glycine max) plants in acid and calcareous soils in the field as well as in a pot culture. Our results demonstrate that soil pH determined the effectiveness of rhizobial inoculation. Rhizobial inoculation resulted in greater effects on nodule traits and plant N content in calcareous soil than in acid soil compared with those in non-inoculated plants. A low P availability significantly decreased nodule number and dry weight in both soils relative to P-application treatments. Rhizobial inoculation did not affect root morphological traits and arbuscular mycorrhizal colonization, but strongly increased the amount of carboxylates in the rhizosheath and plant P content. It also increased plant dry weight and yield in calcareous soil. In conclusion, the beneficial effects of rhizobial inoculation on N and P content and soybean growth were particularly pronounced in calcareous soil. The mining strategy of P acquisition was responsible for increases in P content in inoculated soybean plants compared with non-inoculated controls.

Phosphorus (P) deficiency induces cluster-root formation and carboxylate exudation in most Proteaceae. However, how external P supply regulates these root traits in Macadamia integrifolia remains unclear. Macadamia plants were grown hydroponically with seven P levels to characterize biomass allocation, cluster-root development, and exudation of carboxylates and acid phosphatases. Plant biomass increased with increasing P supply, peaking at 5 μM P, was the same at 5–25 μM P, and declined at 50–100 μM P. Leaf P concentration increased with increasing P supply, but shoot biomass was positively correlated with leaf P concentration up to 0.7–0.8 mg P g–1 dry weight (DW), and declined with further increasing leaf P concentration. The number of cluster roots declined with increasing P supply, with a critical value of leaf P concentration at 0.7–0.8 mg P g–1 DW. We found a similar trend for carboxylate release, with a critical value of leaf P concentration at 0.5 mg g–1 DW, but the activity of acid phosphatases showed a gradually-decreasing trend with increasing P supply. Our results suggest that leaf P concentration regulates the development and functioning of cluster roots, with a critical P concentration of 0.5–0.8 mg g–1, above which macadamia growth is inhibited.

Autotoxicity and allelopathy affect the respiration and yield of GL66 and GL72, two populations of perennial ryegrass (Lolium perenne L. cv. S23) that were originally selected for contrasting rates of mature-leaf dark respiration under conditions where allelopathic effects could not occur and autotoxic effects were minimal. The aim of this study was to further investigate growth and biomass allocation of these two populations in relation to their autotoxic and allelopathic properties. To this end, plants were subjected to two conditions (monoculture and mixed culture) and two treatments (growth in 'renewed' and 'replenished' nutrient solution, allowing for short- and long-term accumulation of allelochemicals, respectively). The fast-respiring population, GL66, showed a reduced total yield due to allelopathic effects only when long-term accumulation of allelochemicals was allowed (mixed culture, replenished). However, short-term accumulation (mixed culture, renewed) of allelochemicals was sufficient to affect allocation of biomass to leaf sheaths. The slow-respiring population, GL72, suffered from autotoxicity only when long-term accumulation was allowed (monoculture, replenished), and from allelopathy under both short- and long-term accumulation (mixed culture, either renewed or replenished). The predominant allelopathic and autotoxic effect was on dry matter percentage and dry weight of leaf sheaths. We conclude that the roots of both populations release one or more chemical compounds that primarily affect biomass allocation to leaf sheaths, both of the same and of the other population. Sensitivity to the putative inhibitor(s) released by the other population was greater than sensitivity to the inhibitor(s) released by a population's own roots.

The 2-fold difference in final length of leaf number three on the main stem between the fast-growing Aegilops tauschii L. and the slow-growing Aegilops caudata L. is correlated with a difference in leaf elongation rate (LER), and not in duration of leaf elongation. In this paper the cellular basis of inherent differences in LER between these species was investigated.The dynamics of abaxial epidermal cells along the growth zone of leaf number three on the main stem of both species was analysed by means of a kinematic analysis. The faster LER of Ae. tauschii compared with that of Ae.�caudata was associated with (i) a larger leaf basal meristem and cell elongation-only zone, and (ii) a faster cell production rate owing to a larger number of dividing cells. Cell division rate, mature cell size and cell elongation rate did not differ between the two species. The lack of variation in cell expansion rate between the species was supported by a similar capacity of both species to extend their isolated cell walls upon acidification.These data suggest that differences in the number of dividing cells can bring about differences in the number of simultaneously elongating cells, and hence in LER.

We compared the use of phosphorus (P) and nitrogen (N) in slow-growing Banksia attenuata (Proteaceae), which resprouts after fire and naturally occurs on deep sand, with faster-growing opportunistic B. sessilis, which is killed by fire and occurs on shallow sand over laterite or limestone. We carried out pot experiments with plants on substrates with different P availability. We measured foliar nutrient concentrations, and P allocated to major biochemical fractions. The two species had similar foliar total P concentrations, but distinct patterns of P allocation to P-containing fractions. The foliar total N concentration of B. sessilis was greater than that of B. attenuata on all substrates. The foliar total P and N concentrations in both species decreased with decreasing P availability. The relative growth rate of both species was positively correlated with both foliar nucleic acid P and total N concentrations, but there was no correlation with other P and N fractions. Faster-growing B. sessilis allocated more P to nucleic acids than B. attenuata did, but other fractions were similar. We conclude that the nutrient-allocation patterns in faster-growing opportunistic B. sessilis and slower-growing B. attenuata revealed different strategies in response to soil P availability, which matched their contrasting growth strategy.

Peanut (Arachis hypogaea L.) is a globally important oil crop, which often experiences poor growth and seedling necrosis under low nocturnal temperatures (LNT). This study assessed the effects of supplementary calcium (Ca2+) and a calmodulin inhibitor on peanut growth and photosynthetic characteristics of plants exposed to LNT, followed by recovery at a higher temperature. We monitored key growth and photosynthetic parameters in a climate-controlled chamber in pots containing soil. LNT reduced peanut growth and dry matter accumulation, enhanced leaf nonstructural carbohydrates concentrations and non-photochemical quenching, decreased the electron transport rate, increased the transmembrane proton gradient, and decreased gas exchange rates. In peanuts subjected to LNT, foliar application of Ca2+ restored growth, dry matter production and leaf photosynthetic capacity. In particular, the foliar Ca2+ application restored temperature-dependent photosynthesis feedback inhibition due to improved growth/ sink demand. Foliar sprays of a calmodulin inhibitor further deteriorated the effects of LNT which validated the protective role of Ca2+ in facilitating LNT tolerance of peanuts.

Previous studies have shown that pH, rather than calcium (Ca), is the main reason why some Lupinus species are sensitive to nutrient solutions mimicking calcareous soils; however, a hydroponic system is quite different from soil systems, and plants may respond differently to these two growing conditions. Thus, studies with Lupinus species grown in calcareous soils are needed. Two calcicole and two calcifuge species were grown in river sand with different Ca forms and amounts, pH levels, and [bicarbonate (HCO3−)] (HCO3− concentration, which is produced by calcium carbonate (CaCO3)). Leaf symptoms, leaf area, gas exchange, biomass, and root morphology were recorded; whole leaf and root nutrient concentrations were analysed. We observed leaf chlorosis of the youngest leaves under high pH (adjusted by KOH) and high pH + high Ca (representing high [HCO3−], high pH and high Ca) treatments for all Lupinus species. However, after 2 weeks, leaf chlorosis of all Lupinus species under high pH started to disappear, with calcicole species fully, and calcifuge species only partly recovering. Leaf chlorosis symptoms of calcicole species under high pH + high Ca partly disappeared as well, while those of calcifuge species did not disappear at all. High pH (resulting from either KOH or HCO3−) inhibited root growth, and subsequently uptake of some nutrients and shoot growth of Lupinus species. However, the strong buffering capacity of HCO3− is the key factor determining if Lupinus species can survive in calcareous soils. Among all studied Lupinus species, L. pilosus was the most tolerant to high [HCO3−] and/or high pH, followed by L. cosentinii and L. angustifolius, while L. hispanicus was the most sensitive.

Hakea prostrata R.Br. (Proteaceae) shows a ‘delayed greening’ strategy of leaf development characterised by reddish young leaves that become green as they mature. This trait may contribute to efficient use of phosphorus (P) during leaf development by first investing P in the development of leaf structure followed by maturation of the photosynthetic machinery. In this study, we investigated the properties of delayed greening in a highly P-efficient species to enhance our understanding of the ecological significance of this trait as a nutrient-saving and photoprotective strategy. In glasshouse-grown plants, we assessed foliar pigments, fatty acids and nutrient composition across five leaf developmental stages. Young leaves had higher concentrations of anthocyanin, P, nitrogen (N), copper (Cu), xanthophyll-cycle pigments and saturated fatty acids than mature leaves. As leaves developed, the concentration of anthocyanins decreased, whereas that of chlorophyll and the double bond index of fatty acids increased. In mature leaves, ~60% of the fatty acids was α-linolenic acid (C18:3 n-3). Mature leaves also had higher concentrations of aluminium (Al), calcium (Ca) and manganese (Mn) than young leaves. We conclude that delayed greening in H. prostrata is a strategy that saves P as well as N and Cu through sequential allocation of these resources, first to cell production and structural development, and then to supplement chloroplast development. This strategy also protects young leaves against photodamage and oxidative stress during leaf expansion under high-light conditions.

Soils used for rice (Oryza sativa L.) cultivation in some areas contain high concentrations of arsenic (As) due to irrigation with groundwater containing As and intensive use of agrochemicals or industrial residues containing As. To restrict rice uptake of As in these soils, approaches to reduce As input and bioavailability must be considered. One approach to reduce As input into rice soils or uptake by rice is cultivating rice under aerobic, intermittent flooding, or alternate wetting and drying (AWD) conditions, rather than in submerged soils, or use of irrigation water low in As. For reducing As bioavailability in soil, aerobic or AWD rice culture and application of biochar, sulfur (S), and/or rice polish to soil are promising. Moreover, use of As-hyperaccumulating plant species (e.g., Pteris vittata L.) in rotation or combinations with favourable plant species (e.g., Azolla, Chlorella, or Nannochloropsis species) can also be promoted, in addition to using rice cultivars that are tolerant to As. Though applications of high doses of phosphorus (P), iron (Fe), and silicon (Si) fertilizers have shown promise in many instances, these methods have to be practiced carefully, because negative effects have also been reported, although such incidents are rare. Major factors affecting As speciation and bioavailability in soil are chemical properties such as redox status, pH, and Fe, P, Si, and S concentrations, physical properties such as texture and organic matter, and biological properties such as methylation activity by soil microorganisms. However, as many of these factors interact, long-term examination under field conditions is needed before measures are recommended for and implemented in farmers' fields.

Root foraging and root physiology such as exudation of carboxylates into the rhizosphere are important strategies for plant phosphorus (P) acquisition. We used 100 chickpea (Cicer arietinum) genotypes with diverse genetic backgrounds to study the relative roles of root morphology and physiology in P acquisition. Plants were grown in pots in a low-P sterilized river sand supplied with 10 mu g P g(-1) soil as FePO4, a poorly soluble form of P. There was a large genotypic variation in root morphology (total root length, root surface area, mean root diameter, specific root length and root hair length), and root physiology (rhizosheath pH, carboxylates and acid phosphatase activity). Shoot P content was correlated with total root length, root surface area and total carboxylates per plant, particularly malonate. A positive correlation was found between mature leaf manganese (Mn) concentration and carboxylate amount in rhizosheath relative to root DW. This is the first study to demonstrate that the mature leaf Mn concentration can be used as an easily measurable proxy for the assessment of belowground carboxylate-releasing processes in a range of chickpea genotypes grown under low-P, and therefore offers an important breeding trait, with potential application in other crops.

Orthophosphate (H2 PO4- , Pi) is an essential macronutrient integral to energy metabolism as well as a component of membrane lipids, nucleic acids, including ribosomal RNA, and therefore essential for protein synthesis. The Pi concentration in the solution of most soils worldwide is usually far too low for maximum growth of crops, including rice. This has prompted the massive use of inefficient, polluting, and nonrenewable phosphorus (P) fertilizers in agriculture. We urgently need alternative and more sustainable approaches to decrease agriculture's dependence on Pi fertilizers. These include manipulating crops by (a) enhancing the ability of their roots to acquire limiting Pi from the soil (i.e. increased P-acquisition efficiency) and/or (b) increasing the total biomass/yield produced per molecule of Pi acquired from the soil (i.e. increased P-use efficiency). Improved P-use efficiency may be achieved by producing high-yielding plants with lower P concentrations or by improving the remobilization of acquired P within the plant so as to maximize growth and biomass allocation to developing organs. Membrane lipid remodelling coupled with hydrolysis of RNA and smaller P-esters in senescing organs fuels P remobilization in rice, the world's most important cereal crop.

Phosphorus deficiency often limits crop productivity, while phosphate rock, which is used to produce phosphorus fertilisers, is a non-renewable resource. Formation of cluster roots is an adaptation to nutrient-poor soils in Lupinus species, including L. albus. Lupinus species also produce nodules, which require a large investment of phosphorus. Our aim is to test whether nodulation promotes cluster-root formation in L. albus. Seedlings of L. albus, either inoculated with rhizobia or non-inoculated, were grown in nutrient solution with a low phosphorus supply. Non-inoculated plants were provided with the same amount of nitrogen in the form of nitrate as the nodulated ones acquired, from both air and nutrient solution, based on preliminary experiments. We measured biomass, phosphorus and nitrogen concentrations as well as photosynthesis just prior to each harvest. Nodulated plants and non-nodulated control plants produced the same amount of biomass. Nodulated plants had, on average, 86% more cluster roots than non-nodulated ones at the four harvests. As hypothesised, nodulation significantly promoted cluster-root formation; it also enhanced rates of photosynthesis. Nodulation promoted cluster-root formation and photosynthesis, presumably because nodules are significant sinks for phosphorus and photosynthates. Our results do not provide evidence for a trade-off between investment of resources in nodules and cluster roots.

Plants exhibit respiratory bypasses (e.g., the alternative oxidase [AOX]) and increase the synthesis of carboxylates in their organs (leaves and roots) in response to phosphorus (P) deficiency, which increases P uptake capacity. They also show differential expression of high-affinity inorganic phosphorus (Pi) transporters, thus avoiding P toxicity at a high P availability. The association between AOX and carboxylate synthesis was tested in Solanum lycopersicum plants grown at different soil P availability, by using plants grown under P-sufficient and P-limiting conditions and by applying a short-term (24 hr) P-sufficient pulse to plants grown under P limitation. Tests were also performed with plants colonized with arbuscular mycorrhizal fungi, which increased plant P concentration under reduced P availability. The in vivo activities of AOX and cytochrome oxidase were measured together with the concentration of carboxylates and the P concentration in plant organs. Gene transcription of Pi transporters (LePT1 and LePT2) was also studied. A coordinated response between plant P concentration with these traits was observed, indicating that a sufficient P availability in soil led to a suppression of both AOX activity and synthesis of citrate and a downregulation of the transcription of genes encoding high-affinity Pi transporters, presumably to avoid P toxicity.

A global data analysis shows that rice grain arsenic (As) concentrations increase with increasing soil As concentrations until about 60 mg As kg−1soil and then decreases. Of the total grain As, 54% is composed of inorganic As. Therefore, when considering the WHO-permissible grain inorganic As concentration, i.e. 0.2 mg As kg−1, the permissible grain total As concentrations is 0.37 mg total As kg−1grain. Soil total As concentration when grain total As concentration reaches permissible level is 5.5 mg As kg−1soil. Therefore, the suitable soil As concentrations for screening rice cultivars in rice agroecosystems for As resistance is 5–60 mg As kg−1soil. Rice has traits to reduce uptake and translocation of As to grains. Cultivars with higher root porosity, radial oxygen loss, or formation of iron plaques bind more As to iron plaques, reducing As uptake (i.e. As avoidance). Once taken up, glutathione/glutaredoxin-mediated As reduction, and phytochelatin-dependent complexation and sequestration in vacuoles result in less translocation of As to the grain. Moreover, generation of reactive oxygen species and the production of antioxidant enzymes further reduce As toxicity (i.e. As resistance). These resistance mechanisms in rice agroecosystems are further enhanced when adequate concentrations of silicon and sulfur are present in soils and tissues, and when plants are associated with arbuscular mycorrhizal fungi, particularly under aerobic or intermittent-aerobic soil condition. Therefore, As concentrations in rice ecosystems decrease in the order of: roots > leaves > grains, and in grains: hull > bran polish > brown rice > raw rice> polished rice > cooked rice. Within the grain, As speciation is affected by the location in the grain, forms of As species, the grain-filling stage, geographic origin, ecosystem management and cultivars used. Indica type accumulates more As in their grains than japonica type. Rice grain production, within safe limits of As, requires the consideration of soil As dynamics including soil management, cultivar responses including uptake and translocation, and post-harvest processing techniques.

Low-molecular-weight organic anions (carboxylates) influence rhizosphere processes and may enhance plant phosphorus acquisition. We examined the root exudate profile of a range of pasture and grain legumes and focused on the little-investigated carboxylate, citramalate. Twelve species of pasture legumes and herbs, including four Lotus spp. and two crop legumes, Cicer arietinum and Lupinus albus, were grown in a glasshouse for six weeks. The composition and amounts of carboxylates were measured in rhizosheath soil as well as in root exudates from roots washed free of their rhizosheath. Citrate and malate were found in the rhizosheath of all species. However, citramalate was present in the rhizosheath of only Lotus species (10–82% of total exuded carboxylates). Cicer arietinum had the largest amount of carboxylates in its rhizosheath and fastest rate of carboxylate exudation into the trap solution. Citrate and malate were found in the rhizosheath of all species in this study, but citramalate was only found in the root exudates and rhizosheath of Lotus spp. Further investigation into the role of citramalate in Lotus spp. is merited.

It has been demonstrated that intercropped crops may benefit from inoculation with rhizobia or arbuscular mycorrhizal (AM) fungi in soybean/maize intercropping systems under sterilized conditions. Little is known about the relative contribution of indigenous and introduced AM fungi and rhizobia to nutrient acquisition of intercropped soybean and maize in unsterilized soils. In this study, we evaluated the effectiveness of indigenous and introduced AM fungi and rhizobia in a soybean/maize intercropping system under unsterilized soil conditions. The results show that introduced rhizobial inoculation promoted soybean nodulation and growth. Single AM fungal inoculation promoted indigenous rhizobial nodulation and increased maize AM root colonization, but had little effect on maize growth. Co-inoculation strongly increased plant dry weight, and nitrogen (N) and phosphorus (P) content of the intercropped soybean compared to those with only indigenous rhizobial and AM fungal communities, but had no effect on those of maize in the same intercropped system. These results indicate that AM fungal inoculation promoted nodulation by indigenous rhizobia. Effective rhizobial inoculation combined with introduced AM fungi may not only promote symbiotic N2 fixation, but also enhance rhizosphere processes, which has the potential to improve nutrient acquisition in the field. Soybean is a weak competitor compared with maize in intercropping systems. Inoculation with introduced AM and rhizobia ameliorated the growth suppression of intercropped soybean, but inhibited growth of intercropped maize, indicating that inoculation of introduced AM fungi and/or rhizobia partly shifted the competitive advantage from maize to soybean. In conclusion, introduced AM fungi and rhizobia contributed more than indigenous populations to promoting growth and nutrient acquisition of the intercropped soybean in unsterilized soils.

We sought to describe the species and functional composition of Brazilian campos rupestres plant communities on severely nutrient-impoverished white sands, to test hypotheses relating plant communities and physiological adaptations to infertile soils. Based on recently-published information on a south-western Australian dune chronosequence, we hypothesised that campos rupestres plant communities would similarly contain a relatively large proportion of non-mycorrhizal species, because of the phosphorus-(P) impoverished nature of the soils. We also sought to test the hypothesis that many of these non-mycorrhizal species have high leaf manganese (Mn) concentrations as a consequence of carboxylate exudation to mobilise soil P. We conducted flora surveys and quantified mycorrhizal status and foliar Mn concentrations in field sites with strongly-weathered sandy soils. Rhizosphere carboxylates were collected from glasshouse-grown plants to assess a potential correlation of carboxylates and leaf Mn concentrations. Soils were depleted of all major plant nutrients. Non-mycorrhizal plants were abundant in most field sites (mean relative cover = 48%). Vellozia species were dominant aboveground; belowground, roots were colonised more by dark septate endophytic fungi than by mycorrhizal fungi. From the field sites, foliar Mn concentrations in non-mycorrhizal species increased with decreasing soil P concentrations, but only when soil Mn concentrations were above a minimum threshold (exchangeable [Mn] above detection limit). Across all species, however, there was no relationship of foliar Mn concentrations with soil P concentrations. Our hypothesis that white-sand campos rupestres communities contain a relatively large proportion of non-mycorrhizal plants was supported. Comparison with similar ecosystems in south-western Australia suggests that plant communities on severely P-impoverished sandy soils, despite differing evolutionary histories and little overlap in plant families, follow convergent evolutionary paths towards increasing abundance of non-mycorrhizal species.

Desiccation tolerance (DT) is the ability of plants to tolerate low leaf water content from which they can recover when water becomes available. DT species mostly occur in seasonally-dry environments, tropical rocky outcrops such as the campos rupestres. Velloziaceae, a prominent family in the campos rupestres, comprises both DT and non-DT species. We tested if DT Vellozia nivea was associated with low productivity under mild drought conditions and survival under extreme drought. We measured traits related to drought resistance (turgor loss point – ΨTLP), plant functioning and productivity (e.g., gas exchange, biomass), non-structural carbohydrate dynamics and mortality in two species, one DT and one non-DT Vellozia species in two water-stress experiments simulating mild and extreme drought. Both species had similar survival rates under mild drought conditions, but the DT species had a lower productivity. Under extreme drought, the dynamics of non-structural carbohydrates was similar in both species but only the non-DT species died, and the cause of mortality was likely hydraulic dysfunction, as they died after reaching ΨTLP. All individuals of the DT species desiccated when they reached ΨTLP but rehydrated after re-watering. We show that the DT strategy in V. nivea implies a costs in terms of productivity, specially under mild drought conditions, but it allows survival under extreme drought, indicating a trade-off between productivity and survival in the DT species. Climate change predictions with more frequent and severe droughts will likely affect the abundance and distribution of DT and non-DT taxa.

Phosphorus (P) deficiency is a major problem for alfalfa (Medicago sativa) productivity on alkaline soils on the Loess Plateau, China. Our aim was to investigate growth, morphological and physiological responses of alfalfa to P supply in two alkaline soils when water supply is limited. A pot experiment was carried out to grow alfalfa in two alkaline soils supplied with different rates of P. Parameters of plant growth and root morphology, rhizosphere pH and carboxylates, and plant concentrations of mineral nutrients were measured. Plant growth and nutrient uptake were enhanced by supplying P, but shoot growth was not further increased when P supply was >20 μg P g−1 soil. Specific root length was only responsive to changes in soil P when P supply was low in the loessial soil. The rhizosphere carboxylate amount was significantly greater when no P was supplied than when P was supplied to the loessial soil. The rhizosphere pH was lower than the bulk soil pH, but did not vary with soil P. A P supply of 20 μg P g−1 soil was optimal for alfalfa growth. The responses of specific root length and rhizosphere carboxylates depended on soil type.

Nitrogen (N) and phosphorus (P) are essential nutrients for plant metabolism, and their availability often limits primary productivity. Whereas the effects of N availability on photosynthetic capacity are well established, we still know relatively little about the effects of P availability at a foliar level, especially in P‐limited tropical forests. We examined photosynthetic capacity, leaf mass per area (LMA) and foliar P fractions in five woody plant species after 6 years of N and P fertilization in a lowland tropical forest. Foliar N:P ratios indicated P limitation of the unfertilized plants; accordingly, photosynthetic P‐use efficiency (PPUE) and LMA decreased with P addition, and foliar N and P concentrations increased, whereas N addition had little effect on measured foliar traits. However, P addition enhanced photosynthetic capacity only in one species and not in other four species. We then assessed plant acclimation to low P availability by quantifying four fractions of foliar P representing different functional pools: structural P, metabolic P (including inorganic P), nucleic acid P, and residual P. We found that P addition enhanced the concentrations of metabolic, structural, and nucleic acid P fractions in all species, but the magnitude of the effect was species‐specific. Our findings indicate that tropical species acclimate to low P availability by altering allocation of foliar P to meet the demand of P for photosynthesis. Importantly, species typical of lowland tropical forests in East Asia maintained their photosynthetic rate under low P availability. We conclude that P limitation of leaf photosynthetic capacity may not be as common as previously assumed due to plant acclimation mechanisms in low‐P tropical forests. Species‐specific strategies to allocate P to different foliar fractions represent a potentially important adaptive mechanism for plants in P‐limited systems.