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Summary: In severely phosphorus (P)‐impoverished environments, plants have evolved to use P very efficiently. Yet, it is unclear how P allocation in leaves contributes to their photosynthetic P‐use efficiency (PPUE) and position along the leaf economics spectrum (LES). We address this question in 10 species of Banksia and Hakea, two highly P‐efficient Proteaceae genera.We characterised traits in leaves of Banksia and Hakea associated with the LES: leaf mass per area, light‐saturated photosynthetic rates, P and nitrogen concentrations, and PPUE. We also determined leaf P partitioning to five biochemical fractions (lipid, nucleic acid, metabolite, inorganic and residual P) and their possible association with the LES.For both genera, PPUE was negatively correlated with fractional allocation of P to lipids, but positively correlated with that to metabolites. For Banksia only, PPUE was negatively correlated with residual P, highlighting a strategy contrasting to that of Hakea. Phosphorus‐allocation patterns significantly explained PPUE but were not linked to the resource acquisition vs resource conservation gradient defined by the LES.We conclude that distinct P‐allocation patterns enable species from different genera to achieve high PPUE and discuss the implications of different P investments. We surmise that different LES axes representing different ecological strategies coexist in extremely P‐impoverished environments. [ABSTRACT FROM AUTHOR]

Background and aims: Proteaceae are a prominent plant family in south-western Australia. Most Proteaceae are 'calcifuge', occurring exclusively on old phosphorus (P)-impoverished acidic soils, with a few 'soil-indifferent' species also found on young P-richer calcareous soils. Calcium (Ca)-enhanced P toxicity explains the calcifuge habit of Proteaceae. However, previous research has so far been focused exclusively on the roles of Ca and P in determining Proteaceae distribution, and consequently there is little knowledge on how other soil-based strategies influence this distribution. We aimed to study the effects of young calcareous soils on four soil-grown Proteaceae and assess differences between calcifuge and soil-indifferent Proteaceae to better understand their natural distribution. Methods: Two calcifuge and two soil-indifferent Proteaceae from south-western Australia were grown in six contrasting soils, including young calcareous, and old acidic soils. Results: When grown in calcareous soils all species showed root growth inhibition, micronutrient deficiency, Ca-enhanced P toxicity, and negative impacts on physiology. Calcifuge species were more sensitive to calcareous soils than soil-indifferent ones, although this varied between genera. Soil-indifferent species tended to produce more cluster roots, release more carboxylates per root mass, and allocate less Ca to their leaves, compared with calcifuges; they also had smaller seeds and were less sensitive to Ca-enhanced P toxicity. Conclusion: We surmise that a combination of these traits allows soil-indifferent species to tolerate calcareous soils. This study provides insight into how Proteaceae respond to young calcareous soils and how this influences their distribution. [ABSTRACT FROM AUTHOR]

Background and Aims Phosphorus (P) availability is often limiting for rice (Oryza sativa) production. Improving internal P-use efficiency (PUE) is crucial to sustainable food production, particularly in low-input systems. A critical aspect of PUE in plants, and one that remains poorly understood, is the investment of leaf P in different chemical P fractions (nucleic acid-P, lipid-P, inorganic-P, metabolite-P and residual-P). The overarching objective of this study was to understand how these key P fractions influence PUE. Methods Three high-PUE and two low-PUE rice genotypes were grown in hydroponics with contrasting P supplies. We measured PUE, total P, P fractions, photosynthesis and biomass. Key Results Low investment in lipid-P was strongly associated with increased photosynthetic PUE (PPUE), achieved by reducing total leaf P concentration while maintaining rapid photosynthetic rates. All low-P plants exhibited a low investment in inorganic-P and lipid-P, but not nucleic acid-P. In addition, whole-plant PUE was strongly associated with reduced total P concentration, increased biomass and increased preferential allocation of resources to the youngest mature leaves. Conclusions Lipid remodelling has been shown in rice before, but we show for the first time that reduced lipid-P investment improves PUE in rice without reducing photosynthesis. This presents a novel pathway for increasing PUE by targeting varieties with reduced lipid-P investment. This will benefit rice production in low-P soils and in areas where fertilizer use is limited, improving global food security by reducing P fertilizer demands and food production costs. [ABSTRACT FROM AUTHOR]

The calcifuge habit of plants is commonly explained in terms of high soil pH and its effects on nutrient availability, particularly that of phosphorus (P). However, most Proteaceae that occur on nutrient‐impoverished soils in south‐western Australia are calcifuge, despite their ability to produce cluster‐roots, which effectively mobilize soil P and micronutrients. We hypothesize that the mechanism explaining the calcifuge habit in Proteaceae is their sensitivity to P and calcium (Ca), and that soil‐indifferent species are less sensitive to the interaction of these nutrients. In this study, we analyzed growth, gas‐exchange rate, and chlorophyll fluorescence of two soil‐indifferent and four calcifuge Hakea and Banksia (Proteaceae) species from south‐western Australia, across a range of P and Ca concentrations in hydroponic solution. We observed Ca‐enhanced P toxicity in all analyzed species, but to different extents depending on distribution type and genus. Increasing P supply enhanced plant growth, leaf biomass, and photosynthetic rates of soil‐indifferent species in a pattern largely independent of Ca supply. In contrast, positive physiological responses to increasing [P] in calcifuges were either absent or limited to low Ca supply, indicating that calcifuges were far more sensitive to Ca‐enhanced P toxicity. In calcifuge Hakeas, we attributed this to higher leaf [P], and in calcifuge Banksias to lower leaf zinc concentration. These differences help to explain these species' contrasting sensitivity to Ca‐enhanced P toxicity and account for the exclusion of most Proteaceae from calcareous habitats. We surmise that Ca‐enhanced P toxicity is a major factor explaining the calcifuge habit of Proteaceae, and, possibly, other P‐sensitive plants. [ABSTRACT FROM AUTHOR]

Background and aims: Plant species richness increases with declining soil phosphorus (P) availability, especially for Proteaceae in old infertile landscapes. This difference in richness might be attributed to faster diversification in lineages adapted to P-impoverished soils, i.e. species that possess specialised P-acquisition strategies, and have lower leaf P concentration ([P]) and higher seed [P]. Alternatively, a longer time for species accumulation might contribute to high species richness in low-P environments due to the geological stability of the landscapes in which they evolved. Methods: We assessed differences in diversification of Proteaceae in P-impoverished vs. nutrient-rich environments and whether these were linked to adaptations to P-impoverished soils. We explored mature leaf and seed [P] and investigated how these traits changed over the evolutionary history of the family, and within two species-rich genera (Banksia, Hakea). Results: Faster diversification was correlated with lower leaf and higher seed [P] for species-rich genera across the Proteaceae. For Banksia and Hakea, diversification rates peaked at relatively low leaf [P], but not at the lowest leaf [P]. Ancestral state reconstructions indicated that low leaf [P] is a trait that was likely present in the early evolution of the Proteaceae, with recent transitions to higher leaf [P] across several species-poor rainforest genera. Conclusions: Diversification of Proteaceae correlated strongly with P-related traits. In an evolutionary context, functional cluster roots, low leaf [P] and high seed [P] were likely key innovations allowing diversification. Selection for low leaf [P] early in the evolutionary history of Proteaceae pre-adapted ancestors of this family to diversify into oligotrophic environments. We discuss how our findings are likely relevant for understanding diversification dynamics of other plant families that occur in P-impoverished environments. [ABSTRACT FROM AUTHOR]

Over 650 Proteaceae occur in south-western Australia, contributing to the region's exceptionally high biodiversity. Most Proteaceae occur exclusively on severely nutrient-impoverished, acidic soils (calcifuge), whilst only few also occur on young, calcareous soils (soil-indifferent), higher in calcium (Ca) and phosphorus (P). The calcifuge habit of Proteaceae is explained by Ca-enhanced P toxicity, putatively linked to the leaf cell-specific allocation of Ca and P. Separation of these elements is essential to avoid the deleterious precipitation of Ca-phosphate. We used quantitative X-ray microanalysis to determine leaf cell-specific nutrient concentrations of two calcifuge and two soil-indifferent Proteaceae grown in hydroponics at a range of Ca and P concentrations. Calcium enhanced the preferential allocation of P to palisade mesophyll (PM) cells under high P conditions, without a significant change in whole leaf [P]. Calcifuges showed a greater PM [P] compared with soil-indifferent species, corresponding to their greater sensitivity. This study advances our mechanistic understanding of Ca-enhanced P toxicity, supporting the proposed model, and demonstrating its role in the calcifuge distribution of Proteaceae. This furthers our understanding of nutrient interactions at the cellular level and highlights its importance to plant functioning. [ABSTRACT FROM AUTHOR]

The Jurien Bay dune chronosequence in south‐western Australia's biodiversity hotspot comprises sites differing in nutrient availability, with phosphorus (P) availability declining strongly with increasing soil age. We have explored the exceptionally high photosynthetic P‐use efficiency (PPUE) of Proteaceae in this region, triggering the question what the PPUE of co‐occurring species in other families might be along the Jurien Bay chronosequence.We explored how traits associated with PPUE, photosynthetic nitrogen (N)‐use efficiency (PNUE) and leaf respiration might converge along the chronosequence, and whether Proteaceae and non‐Proteaceae species differ in leaf traits associated with nutrient use.Seven to 10 species were sampled at three sites differing in nutrient availability (ranging from N‐ to P‐limited). Measurements of leaf light‐saturated photosynthesis and dark respiration were integrated with measurements of total N and P concentration in both mature and senesced leaves, and leaf mass per unit area (LMA).Contrary to what is known for other chronosequences, rates of photosynthesis and respiration did not decrease with increasing soil age and LMA along the Jurien Bay chronosequence. However, they increased when expressed per unit leaf P. Both N and P were used much more efficiently for photosynthesis on nutrient‐poor sites, in both Proteaceae and non‐Proteaceae species. Proteaceae had the fastest rate of photosynthesis per unit leaf P, followed by species that preferentially allocate P to mesophyll cells, rather than epidermal cells.Synthesis. Our results show that with declining soil P availability, photosynthetic P‐use efficiency of all investigated species from different families increased. Plants growing on the oldest, most nutrient‐impoverished soils exhibited similar rates of CO2 exchange as plants growing on more nutrient‐rich younger soils, and extraordinarily high photosynthetic P‐use efficiency. This indicates convergence in leaf traits related to photosynthetic nutrient use on severely P‐impoverished sites. [ABSTRACT FROM AUTHOR]

Summary: Many Proteaceae are highly phosphorus (P)‐sensitive and occur exclusively on old nutrient‐impoverished acidic soils (calcifuge), whilst a few also occur on young calcareous soils (soil‐indifferent) that are higher in available calcium (Ca) and P. Calcium increases the severity of P‐toxicity symptoms, but its underlying mechanisms are unknown. We propose that Ca‐enhanced P toxicity explains the calcifuge habit of most Proteaceae.Four calcifuge and four soil‐indifferent Proteaceae from South‐Western Australia were grown in hydroponics, at a range of P and Ca concentrations.Calcium increased the severity of P‐toxicity symptoms in all species. Calcifuge Proteaceae were more sensitive to Ca‐enhanced P toxicity than soil‐indifferent ones. Calcifuges shared these traits: low leaf zinc concentration ([Zn]), low Zn allocation to leaves, low leaf [Zn]:[P], low root : shoot ratio, and high seed P content, compared with soil‐indifferent species.This is the first demonstration of Ca‐enhanced P toxicity across multiple species. Calcium‐enhanced P toxicity provides an explanation for the calcifuge habit of most Proteaceae and is critical for the management of this iconic Australian family. This study represents a major advance towards an understanding of the physiological mechanisms of P toxicity and its role in the distribution of Proteaceae. [ABSTRACT FROM AUTHOR]

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

Abstract: Plants allocate nutrients to specific leaf cell types; eudicots are thought to predominantly allocate phosphorus (P) to epidermal/bundle sheath cells. However, three Proteaceae species have been shown to preferentially allocate P to mesophyll cells instead. These Proteaceae species are highly adapted to P‐impoverished habitats, with exceptionally high photosynthetic P‐use efficiencies (PPUE). We hypothesized that preferential allocation of P to photosynthetic mesophyll cells is an important trait in species adapted to extremely P‐impoverished habitats, contributing to their high PPUE. We used elemental X‐ray mapping to determine leaf cell‐specific nutrient concentrations for 12 Proteaceae species, from habitats of strongly contrasting soil P concentrations, in Australia, Brazil, and Chile. We found that only species from extremely P‐impoverished habitats preferentially allocated P to photosynthetic mesophyll cells, suggesting it has evolved as an adaptation to their extremely P‐impoverished habitat and that it is not a family‐wide trait. Our results highlight the possible role of soil P in driving the evolution of ecologically relevant nutrient allocation patterns and that these patterns cannot be generalized across families. Furthermore, preferential allocation of P to photosynthetic cells may provide new and exciting strategies to improve PPUE in crop species. [ABSTRACT FROM AUTHOR]

Plants that deploy a phosphorus (P)-mobilising strategy based on the release of carboxylates tend to have high leaf manganese concentrations ([Mn]). This occurs because the carboxylates mobilise not only soil inorganic and organic P, but also a range of micronutrients, including Mn. Concentrations of most other micronutrients increase to a small extent, but Mn accumulates to significant levels, even when plants grow in soil with low concentrations of exchangeable Mn availability. Here, we propose that leaf [Mn] can be used to select for genotypes that are more efficient at acquiring P when soil P availability is low. Likewise, leaf [Mn] can be used to screen for belowground functional traits related to nutrient-acquisition strategies among species in low-P habitats. [ABSTRACT FROM AUTHOR]

The article describes the assemblage of minerals that were identified from a hedyphane-bearing tunnel and a fornacite-bearing pit in Singer mine located in the Goodsprings mining district of Clark County, Nevada. The identities of the minerals describes in this article were confirmed by semiquantitative EDS and/or powder x-ray diffraction (XRD) analysis. Minerals identified include Aragonite, Barite and Calcite.

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

Phosphorus (P) is an essential plant nutrient. Most rice growing lands lack adequate P, requiring multiple P fertiliser applications to obtain expected yields. However, P fertiliser is environmentally damaging, and already unaffordable to the marginal farmers. This warrants developing P-efficient rice varieties that require less P to produce the expected yield. However, genetic factors underlying P-use efficiency (PUE) in rice remain elusive. Here, we conducted comparative transcriptome analysis using two rice varieties with contrasting PUE; a P-efficient landrace DJ123 and a P-inefficient modern cultivar IR64. We aimed to understand the transcriptomic responses in DJ123 that allow it to achieve a high PUE under low P conditions. Our results showed that both DJ123 and IR64 had replete tissue P concentrations after 48 h of P deprivation. Yet, DJ123 strongly responded to the external low P availability by inducing P starvation-inducible genes that included SPX2, PHO1, PAPs and SQDs, while these genes were not significantly induced in IR64. We envisage that the ability of DJ123 to rapidly respond to low P conditions might be the key to its high PUE. Our findings lay a valuable foundation in elucidating PUE mechanism in rice, thus will potentially contribute to developing P-efficient modern rice variety. [ABSTRACT FROM AUTHOR]

Phosphorus (P) is an essential plant nutrient. Most rice growing lands lack adequate P, requiring multiple P fertiliser applications to obtain expected yields. However, P fertiliser is environmentally damaging, and already unaffordable to the marginal farmers. This warrants developing P-efficient rice varieties that require less P to produce the expected yield. However, genetic factors underlying P-use efficiency (PUE) in rice remain elusive. Here, we conducted comparative transcriptome analysis using two rice varieties with contrasting PUE; a P-efficient landrace DJ123 and a P-inefficient modern cultivar IR64. We aimed to understand the transcriptomic responses in DJ123 that allow it to achieve a high PUE under low P conditions. Our results showed that both DJ123 and IR64 had replete tissue P concentrations after 48 h of P deprivation. Yet, DJ123 strongly responded to the external low P availability by inducing P starvation-inducible genes that included SPX2, PHO1, PAPs and SQDs, while these genes were not significantly induced in IR64. We envisage that the ability of DJ123 to rapidly respond to low P conditions might be the key to its high PUE. Our findings lay a valuable foundation in elucidating PUE mechanism in rice, thus will potentially contribute to developing P-efficient modern rice variety. [ABSTRACT FROM AUTHOR]

Background and aims: 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. Methods: 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. 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. Conclusions: 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. [ABSTRACT FROM AUTHOR]

Belowground organisms play critical roles in maintaining multiple ecosystem processes, including plant productivity, decomposition, and nutrient cycling. Despite their importance, however, we have a limited understanding of how and why belowground biodiversity (bacteria, fungi, protists, and invertebrates) may change as soils develop over centuries to millennia (pedogenesis). Moreover, it is unclear whether belowground biodiversity changes during pedogenesis are similar to the patterns observed for aboveground plant diversity. Here we evaluated the roles of resource availability, nutrient stoichiometry, and soil abiotic factors in driving belowground biodiversity across 16 soil chronosequences (from centuries to millennia) spanning a wide range of globally distributed ecosystem types. Changes in belowground biodiversity during pedogenesis followed two main patterns. In lower-productivity ecosystems (i.e., drier and colder), increases in belowground biodiversity tracked increases in plant cover. In more productive ecosystems (i.e., wetter and warmer), increased acidification during pedogenesis was associated with declines in belowground biodiversity. Changes in the diversity of bacteria, fungi, protists, and invertebrates with pedogenesis were strongly and positively correlated worldwide, highlighting that belowground biodiversity shares similar ecological drivers as soils and ecosystems develop. In general, temporal changes in aboveground plant diversity and belowground biodiversity were not correlated, challenging the common perception that belowground biodiversity should follow similar patterns to those of plant diversity during ecosystem development. Taken together, our findings provide evidence that ecological patterns in belowground biodiversity are predictable across major globally distributed ecosystem types and suggest that shifts in plant cover and soil acidification during ecosystem development are associated with changes in belowground biodiversity over centuries to millennia. [ABSTRACT FROM AUTHOR]

The abundance of nitrogen (N)-fixing plants in ecosystems where phosphorus (P) limits plant productivity poses a paradox because N fixation entails a high P cost. One explanation for this paradox is that the N-fixing strategy allows greater root phosphatase activity to enhance P acquisition from organic sources, but evidence to support this contention is limited., We measured root phosphomonoesterase ( PME) activity of 10 N-fixing species, including rhizobial legumes and actinorhizal Allocasuarina species, and eight non-N-fixing species across a retrogressive soil chronosequence showing a clear shift from N to P limitation of plant growth and representing a strong natural gradient in P availability., Legumes showed greater root PME activity than non-legumes, with the difference between these two groups increasing markedly as soil P availability declined. By contrast, root PME activity of actinorhizal species was always lower than that of co-occurring legumes and not different from non-N-fixing plants., The difference in root PME activity between legumes and actinorhizal plants was not reflected in a greater or similar reliance on N fixation for N acquisition by actinorhizal species compared to co-occurring legumes., Synthesis. Our results support the idea that N-fixing legumes show high root phosphatase activity, especially at low soil P availability, but suggest that this is a phylogenetically conserved trait rather than one directly linked to their N-fixation capacity. [ABSTRACT FROM AUTHOR]

Background and aims: Southwest Australia is a biodiversity hotspot, with greatest plant species diversity on the most severely phosphorus (P)-impoverished soils. Here, non-mycorrhizal species with highly-effective carboxylate-releasing P-acquisition strategies coexist with mycorrhizal species that are less effective at accessing P on these soils. Non-mycorrhizal carboxylate-releasing species facilitate P acquisition of mycorrhizal neighbours that are better defended against pathogens. In the Southwest Australian Biodiversity Hotspot, there are also ‘cool spots’ of low-diversity tall mycorrhizal Eucalyptus communities on P-impoverished soils. These Eucalyptus trees obviously do not require facilitation of their P acquisition by carboxylate-releasing neighbours, because these are only a minor component of the low-diversity communities. We hypothesised that in low-diversity tall Eucalyptus forests, mycorrhizal species release carboxylates to acquire P. Thus, they would not depend on facilitation, and must be strong competitors. However, because they would not depend on external mycorrhizal hyphae to acquire P, they would also not be able to access soil organic nitrogen (N), for which they would need external hyphae.Since carboxylates not only mobilise P, but also manganese (Mn), we used leaf Mn concentrations ([Mn]) in the natural habitat to proxy rhizosphere carboxylates. To verify this proxy, we also measured carboxylate exudation of targeted species with high leaf [Mn] using seedlings grown in low-P nutrient solutions.Using these complementary approaches, we confirmed our hypothesis that dominant Eucalyptus species in ‘cool spots’ release carboxylates. Since mineralisation of organic N is associated with fractionation of N, enriching organic N with 15N while nitrate is depleted in 15N, we measured the stable N isotope composition of leaf material. The results show that dominant Eucalyptus species did not access organic N, despite being ectomycorrhizal.The low diversity of tall Eucalyptus forests in southwest Australia can be explained by dominant mycorrhizal species exhibiting a carboxylate-releasing strategy. The tall eucalypts are therefore strong competitors that do not require facilitation, but also do not access organic N.Methods: Southwest Australia is a biodiversity hotspot, with greatest plant species diversity on the most severely phosphorus (P)-impoverished soils. Here, non-mycorrhizal species with highly-effective carboxylate-releasing P-acquisition strategies coexist with mycorrhizal species that are less effective at accessing P on these soils. Non-mycorrhizal carboxylate-releasing species facilitate P acquisition of mycorrhizal neighbours that are better defended against pathogens. In the Southwest Australian Biodiversity Hotspot, there are also ‘cool spots’ of low-diversity tall mycorrhizal Eucalyptus communities on P-impoverished soils. These Eucalyptus trees obviously do not require facilitation of their P acquisition by carboxylate-releasing neighbours, because these are only a minor component of the low-diversity communities. We hypothesised that in low-diversity tall Eucalyptus forests, mycorrhizal species release carboxylates to acquire P. Thus, they would not depend on facilitation, and must be strong competitors. However, because they would not depend on external mycorrhizal hyphae to acquire P, they would also not be able to access soil organic nitrogen (N), for which they would need external hyphae.Since carboxylates not only mobilise P, but also manganese (Mn), we used leaf Mn concentrations ([Mn]) in the natural habitat to proxy rhizosphere carboxylates. To verify this proxy, we also measured carboxylate exudation of targeted species with high leaf [Mn] using seedlings grown in low-P nutrient solutions.Using these complementary approaches, we confirmed our hypothesis that dominant Eucalyptus species in ‘cool spots’ release carboxylates. Since mineralisation of organic N is associated with fractionation of N, enriching organic N with 15N while nitrate is depleted in 15N, we measured the stable N isotope composition of leaf material. The results show that dominant Eucalyptus species did not access organic N, despite being ectomycorrhizal.The low diversity of tall Eucalyptus forests in southwest Australia can be explained by dominant mycorrhizal species exhibiting a carboxylate-releasing strategy. The tall eucalypts are therefore strong competitors that do not require facilitation, but also do not access organic N.Results: Southwest Australia is a biodiversity hotspot, with greatest plant species diversity on the most severely phosphorus (P)-impoverished soils. Here, non-mycorrhizal species with highly-effective carboxylate-releasing P-acquisition strategies coexist with mycorrhizal species that are less effective at accessing P on these soils. Non-mycorrhizal carboxylate-releasing species facilitate P acquisition of mycorrhizal neighbours that are better defended against pathogens. In the Southwest Australian Biodiversity Hotspot, there are also ‘cool spots’ of low-diversity tall mycorrhizal Eucalyptus communities on P-impoverished soils. These Eucalyptus trees obviously do not require facilitation of their P acquisition by carboxylate-releasing neighbours, because these are only a minor component of the low-diversity communities. We hypothesised that in low-diversity tall Eucalyptus forests, mycorrhizal species release carboxylates to acquire P. Thus, they would not depend on facilitation, and must be strong competitors. However, because they would not depend on external mycorrhizal hyphae to acquire P, they would also not be able to access soil organic nitrogen (N), for which they would need external hyphae.Since carboxylates not only mobilise P, but also manganese (Mn), we used leaf Mn concentrations ([Mn]) in the natural habitat to proxy rhizosphere carboxylates. To verify this proxy, we also measured carboxylate exudation of targeted species with high leaf [Mn] using seedlings grown in low-P nutrient solutions.Using these complementary approaches, we confirmed our hypothesis that dominant Eucalyptus species in ‘cool spots’ release carboxylates. Since mineralisation of organic N is associated with fractionation of N, enriching organic N with 15N while nitrate is depleted in 15N, we measured the stable N isotope composition of leaf material. The results show that dominant Eucalyptus species did not access organic N, despite being ectomycorrhizal.The low diversity of tall Eucalyptus forests in southwest Australia can be explained by dominant mycorrhizal species exhibiting a carboxylate-releasing strategy. The tall eucalypts are therefore strong competitors that do not require facilitation, but also do not access organic N.Conclusions: Southwest Australia is a biodiversity hotspot, with greatest plant species diversity on the most severely phosphorus (P)-impoverished soils. Here, non-mycorrhizal species with highly-effective carboxylate-releasing P-acquisition strategies coexist with mycorrhizal species that are less effective at accessing P on these soils. Non-mycorrhizal carboxylate-releasing species facilitate P acquisition of mycorrhizal neighbours that are better defended against pathogens. In the Southwest Australian Biodiversity Hotspot, there are also ‘cool spots’ of low-diversity tall mycorrhizal Eucalyptus communities on P-impoverished soils. These Eucalyptus trees obviously do not require facilitation of their P acquisition by carboxylate-releasing neighbours, because these are only a minor component of the low-diversity communities. We hypothesised that in low-diversity tall Eucalyptus forests, mycorrhizal species release carboxylates to acquire P. Thus, they would not depend on facilitation, and must be strong competitors. However, because they would not depend on external mycorrhizal hyphae to acquire P, they would also not be able to access soil organic nitrogen (N), for which they would need external hyphae.Since carboxylates not only mobilise P, but also manganese (Mn), we used leaf Mn concentrations ([Mn]) in the natural habitat to proxy rhizosphere carboxylates. To verify this proxy, we also measured carboxylate exudation of targeted species with high leaf [Mn] using seedlings grown in low-P nutrient solutions.Using these complementary approaches, we confirmed our hypothesis that dominant Eucalyptus species in ‘cool spots’ release carboxylates. Since mineralisation of organic N is associated with fractionation of N, enriching organic N with 15N while nitrate is depleted in 15N, we measured the stable N isotope composition of leaf material. The results show that dominant Eucalyptus species did not access organic N, despite being ectomycorrhizal.The low diversity of tall Eucalyptus forests in southwest Australia can be explained by dominant mycorrhizal species exhibiting a carboxylate-releasing strategy. The tall eucalypts are therefore strong competitors that do not require facilitation, but also do not access organic N. [ABSTRACT FROM AUTHOR]

Background: Unveiling the diversity of plant strategies to acquire and use phosphorus (P) is crucial to understand factors promoting their coexistence in hyperdiverse P-impoverished communities within fire-prone landscapes such as in cerrado (South America), fynbos (South Africa) and kwongan (Australia).We explore the diversity of P-acquisition strategies, highlighting one that has received little attention: acquisition of P following fires that temporarily enrich soil with P. This strategy is expressed by fire ephemerals as well as fast-resprouting perennial shrubs. A plant’s leaf manganese concentration ([Mn]) provides significant clues on P-acquisition strategies. High leaf [Mn] indicates carboxylate-releasing P-acquisition strategies, but other exudates may play the same role as carboxylates in P acquisition. Intermediate leaf [Mn] suggests facilitation of P acquisition by P-mobilising neighbours, through release of carboxylates or functionally similar compounds. Very low leaf [Mn] indicates that carboxylates play no immediate role in P acquisition. Release of phosphatases also represents a P-mining strategy, mobilising organic P. Some species may express multiple strategies, depending on time since germination or since fire, or on position in the landscape. In severely P-impoverished landscapes, photosynthetic P-use efficiency converges among species. Efficient species exhibit rapid rates of photosynthesis at low leaf P concentrations. A high P-remobilisation efficiency from senescing organs is another way to use P efficiently, as is extended longevity of plant organs.Many P-acquisition strategies coexist in P-impoverished landscapes, but P-use strategies tend to converge. Common strategies of which we know little are those expressed by ephemeral or perennial species that are the first to respond after a fire. We surmise that carboxylate-releasing P-mobilising strategies are far more widespread than envisaged so far, and likely expressed by species that accumulate metals, exemplified by Mn, metalloids, such as selenium, fluorine, in the form of fluoroacetate, or silicon. Some carboxylate-releasing strategies are likely important to consider when restoring sites in biodiverse regions as well as in cropping systems on P-impoverished or strongly P-sorbing soils, because some species may only be able to establish themselves next to neighbours that mobilise P.Scope: Unveiling the diversity of plant strategies to acquire and use phosphorus (P) is crucial to understand factors promoting their coexistence in hyperdiverse P-impoverished communities within fire-prone landscapes such as in cerrado (South America), fynbos (South Africa) and kwongan (Australia).We explore the diversity of P-acquisition strategies, highlighting one that has received little attention: acquisition of P following fires that temporarily enrich soil with P. This strategy is expressed by fire ephemerals as well as fast-resprouting perennial shrubs. A plant’s leaf manganese concentration ([Mn]) provides significant clues on P-acquisition strategies. High leaf [Mn] indicates carboxylate-releasing P-acquisition strategies, but other exudates may play the same role as carboxylates in P acquisition. Intermediate leaf [Mn] suggests facilitation of P acquisition by P-mobilising neighbours, through release of carboxylates or functionally similar compounds. Very low leaf [Mn] indicates that carboxylates play no immediate role in P acquisition. Release of phosphatases also represents a P-mining strategy, mobilising organic P. Some species may express multiple strategies, depending on time since germination or since fire, or on position in the landscape. In severely P-impoverished landscapes, photosynthetic P-use efficiency converges among species. Efficient species exhibit rapid rates of photosynthesis at low leaf P concentrations. A high P-remobilisation efficiency from senescing organs is another way to use P efficiently, as is extended longevity of plant organs.Many P-acquisition strategies coexist in P-impoverished landscapes, but P-use strategies tend to converge. Common strategies of which we know little are those expressed by ephemeral or perennial species that are the first to respond after a fire. We surmise that carboxylate-releasing P-mobilising strategies are far more widespread than envisaged so far, and likely expressed by species that accumulate metals, exemplified by Mn, metalloids, such as selenium, fluorine, in the form of fluoroacetate, or silicon. Some carboxylate-releasing strategies are likely important to consider when restoring sites in biodiverse regions as well as in cropping systems on P-impoverished or strongly P-sorbing soils, because some species may only be able to establish themselves next to neighbours that mobilise P.Conclusions: Unveiling the diversity of plant strategies to acquire and use phosphorus (P) is crucial to understand factors promoting their coexistence in hyperdiverse P-impoverished communities within fire-prone landscapes such as in cerrado (South America), fynbos (South Africa) and kwongan (Australia).We explore the diversity of P-acquisition strategies, highlighting one that has received little attention: acquisition of P following fires that temporarily enrich soil with P. This strategy is expressed by fire ephemerals as well as fast-resprouting perennial shrubs. A plant’s leaf manganese concentration ([Mn]) provides significant clues on P-acquisition strategies. High leaf [Mn] indicates carboxylate-releasing P-acquisition strategies, but other exudates may play the same role as carboxylates in P acquisition. Intermediate leaf [Mn] suggests facilitation of P acquisition by P-mobilising neighbours, through release of carboxylates or functionally similar compounds. Very low leaf [Mn] indicates that carboxylates play no immediate role in P acquisition. Release of phosphatases also represents a P-mining strategy, mobilising organic P. Some species may express multiple strategies, depending on time since germination or since fire, or on position in the landscape. In severely P-impoverished landscapes, photosynthetic P-use efficiency converges among species. Efficient species exhibit rapid rates of photosynthesis at low leaf P concentrations. A high P-remobilisation efficiency from senescing organs is another way to use P efficiently, as is extended longevity of plant organs.Many P-acquisition strategies coexist in P-impoverished landscapes, but P-use strategies tend to converge. Common strategies of which we know little are those expressed by ephemeral or perennial species that are the first to respond after a fire. We surmise that carboxylate-releasing P-mobilising strategies are far more widespread than envisaged so far, and likely expressed by species that accumulate metals, exemplified by Mn, metalloids, such as selenium, fluorine, in the form of fluoroacetate, or silicon. Some carboxylate-releasing strategies are likely important to consider when restoring sites in biodiverse regions as well as in cropping systems on P-impoverished or strongly P-sorbing soils, because some species may only be able to establish themselves next to neighbours that mobilise P. [ABSTRACT FROM AUTHOR]