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

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.

Leaf phosphorus (P) comprises four major fractions: inorganic phosphate (P

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. Methods 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. Results 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. Conclusions 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.

Plantago is a major genus belonging to the Plantaginaceae family and is used in herbal medicine, functional food, and pastures. Several Plantago species are also characterized by their global distribution, but the mechanism underpinning this is not known. Here, we present a high-quality, chromosome-level genome assembly of Plantago major L., a species of Plantago, by incorporating Oxford Nanopore sequencing and Hi-C technologies. The genome assembly size was approximately 671.27 Mb with a contig N50 length of 31.30 Mb. 31,654 protein-coding genes were identified from the genome. Evolutionary analysis showed that P. major diverged from other Lamiales species at ~62.18 Mya and experienced two rounds of WGD events. Notably, many gene families related to plant acclimation and adaptation expanded. We also found that many polyphenol biosynthesis genes showed high expression patterns in roots. Some amino acid biosynthesis genes, such as those involved in histidine synthesis, were highly induced under metal (Ni) stress which led to the accumulation of corresponding metabolites. These results suggest persuasive arguments for the global distribution of P. major through multiscale analysis. Decoding the P. major genome provides a valuable genomic resource for research on dissecting biological function, molecular evolution, taxonomy, and breeding

Background and Aims The leaf economic spectrum (LES) is related to dry mass and nutrient investments towards photosynthetic processes and leaf structures, and to the duration of returns on those investments (leaf lifespan, LL). Phosphorus (P) is a key limiting nutrient for plant growth, yet it is unclear how the allocation of leaf P among different functions is coordinated with the LES. We addressed this question among 12 evergreen woody species co-occurring on P-impoverished soils in south-eastern Australia. Methods Leaf ‘economic’ traits, including LL, leaf mass per area (LMA), light-saturated net photosynthetic rate per mass (Amass), dark respiration rate, P concentration ([Ptotal]), nitrogen concentration, and P resorption, were measured for three pioneer and nine non-pioneer species. Leaf P was separated into five functional fractions: orthophosphate P (Pi), metabolite P (PM), nucleic acid P (PN), lipid P (PL), and residual P (PR; phosphorylated proteins and unidentified compounds that contain P). Results LL was negatively correlated with Amass and positively correlated with LMA, representing the LES. Pioneers occurred towards the short-LL end of the spectrum and exhibited higher [Ptotal] than non-pioneer species, primarily associated with higher concentrations of Pi, PN and PL. There were no significant correlations between leaf P fractions and LL or LMA, while Amass was positively correlated with the concentration of PR. Conclusions Allocation of leaf P to different fractions varied substantially among species. This variation was partially associated with the LES, which may provide a mechanism underlying co-occurrence of species with different ecological strategies under P limitation.

Background and aims This study investigated whether root traits at the seedling stage are maintained at the flowering stage in two chickpea (Cicer arietinum) genotypes with contrasting root morphology and physiology; and whether the genotype with greater rhizosheath carboxylates mobilises more poorly-available phosphorus (P) pools to increase shoot P at flowering/podding and seed yield at maturity. Methods Two chickpea genotypes were grown in a low P soil with or without P addition (0 and 40 µg P g−1 soil as KH2PO4) under controlled glasshouse conditions and harvested at seedling, flowering/podding, physiological maturity. Results At the seedling and flowering/podding stages, ICC2884 had thinner roots and greater root mass ratio, specific root length and rhizosheath carboxylates per root dry weight (DW) than ICC456. Both genotypes had smaller root diameter, higher carboxylates and acid phosphatase activity in rhizosheath soil at flowering/podding than at seedling. In the rhizosheath soil of both genotypes, NaHCO3-Pi concentration was depleted under P0 only; under both P0 and P40, NaHCO3-Po concentration increased while NaOH-Pi and NaOH-Po concentrations decreased at the seedling stage but accumulated at the flowering/podding stage, relative to the bulk soil. ICC2884 did not mobilise more poorly available soil P or acquire more P at the seedling or flowering/podding stages, or produce higher seed yields than ICC456. Conclusion ICC2884 and ICC456 maintained the difference in root morphological and physiological characteristics from the seedling stage to the flowering/podding stage. The genotype with greater rhizosheath carboxylates (root DW basis) did not produce higher yield than genotype with less rhizosheath carboxylates.

Aims South-western Australia has some of the most phosphorus (P)-impoverished soils in the world. Hakea prostrata (Proteaceae) has nitrate (NO3−)-uptake restraint, associated with its high P-use efficiency. This research explores how common this trait is in other Proteaceae and in co-occurring plant species in south-western Australia. Methods Banksia attenuata (Proteaceae), B. telmatiaea (Proteaceae), Melaleuca seriata (Myrtaceae) and M. rhaphiophylla (Myrtaceae) were grown with no NO3−, Medium (500 µM) NO3− or High (2500 µM) NO3− treatments with no other source of nitrogen (N). Plants were harvested after treatments. Leaf nitrate and phosphate concentrations were determined, as well as biomass, total N and total P concentrations of leaves, stems and roots separately. Results Banksia attenuata, B. telmatiaea and M. seriata had similar total N content when supplied with High NO3− as they did when supplied with Medium NO3−. All four species had the same low leaf NO3−-N concentration in High and Medium NO3− treatments, no more than 6% of leaf total N. All species had similar leaf NO3− concentrations as those of plants growing in their natural habitat where the available soil nitrate concentration was much lower. These results are in sharp contrast with plants that are typically studied, for example Arabidopsis thaliana. Conclusions All four species exhibited NO3−-uptake restraint to different degrees. Nitrate-uptake restraint appears a convergent trait common to many species that evolved in severely P-impoverished landscapes, allowing them to be more P-efficient.

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). Scope 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. Conclusions 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.

Background and aims Waterlilies (Nymphaea spp) are ancient iconic plants. Scientific knowledge of their nutrient requirements is scarce. We investigated plant responses to phosphorus (P) and potassium (K) nutrition in a cultivar of tropical waterlilies used for commercial flower production. Methods Two studies with waterlilies were conducted simultaneously over 24 weeks. In Experiment 1, three amounts of fertiliser differing in P content were supplied either four, six or 12 times. Experiment 2 was similar, but fertiliser varied in K content. Flower production was recorded every two weeks, detailed plant measurements were made every six weeks and leaves were collected for nutrient analyses at week 20. At week 24, shoot biomass was harvested and weighed. Results Total flower production increased with increasing P supply but decreased with increasing K supply. With increasing P supply, leaf P concentration increased from 1.3 to 2.0 mg g−1 dry weight. Increasing the K supply decreased leaf P concentration but had no effect on K concentration. In the P experiment, leaf calcium and magnesium concentrations were generally low as was the leaf zinc concentration. Final plant size increased with increasing P supply but declined with increasing K supply. Conclusion Waterlily growth and flowering declined with increasing K supply and increased with increasing P supply. Fertiliser-P requirement was very high, and it is possible that plants would have responded to greater amounts of P than we used. This was partly due to the very high P-sorbing capacity of the soil we used. Inefficient nutrient uptake owing to the low capacity for P acquisition of waterlily roots at the near-neutral pH of flooded soil was also a likely factor.

Background and aims Understanding the contributions of winter cover crops (CCs) to nitrogen (N) uptake of succeeding spring maize will help achieve more efficient soil N use. Methods A single-factor field experiment was conducted to quantify residual N effects of contrasting CC tops on spring maize N utilization and environmental benefits. 15N-labeled fertilizers were applied at the time of CC establishment in a microplot experiment to determine the contribution of aboveground residue N of winter cover crops (hairy vetch (HV), February orchid (OV), hairy vetch/ February orchid mixture (HO)) to the following spring maize. Results Compared with the winter fallow, HV, OV and HO treatments significantly increased the partial factor productivity of N fertilizer by 43%, 16%, and 23%, respectively. The spring maize N derived from the residues was 27 (HV), 6 (OV) and 22 (HO) kg ha−1 and the total contribution to the N uptake of spring maize from the aboveground residues was 2.2~10.3%. Net N surplus was defined as total N input minus N output, and the value of OV (112 kg N ha−1) was significantly lower than that of OV (182 kg N ha−1) and HO (197 kg N ha−1). The HO was the most environmental-friendly cropping system with the lowest carbon footprint, which decreased greenhouse gas (GHG) emissions by 30%. Conclusions The introduction of winter mixed CCs in the rotation system of three crops across two years on the North China Plain had a higher N-use efficiency, maintained yields with higher net profits, and lowered emissions.

Background and aims In extremely low-phosphorus (P) environments, most Proteaceae exude carboxylates from cluster roots. These carboxylates mobilise inorganic P which leads to a relatively high leaf manganese concentration ([Mn]). However, we found that Adenanthos cygnorum (Proteaceae) in a low-P habitat did not invariably have a high leaf [Mn] in south-western Australia. We aimed to explore how A. cygnorum acquires P in severely P-impoverished habitats. Methods We determined soil P concentrations and leaf [Mn] of A. cygnorum growing within 1 m and more than 10 m away from other large Proteaceae. We also grew plants in a glasshouse to determine its root carboxylate exudation and rhizosheath phosphatase activity. Results Adenanthos cygnorum did not produce functional cluster roots. It depended on carboxylates released by a P-mobilising neighbour, Banksia attenuata (Proteaceae), to acquire P when growing in severely P-impoverished soil (− 1 dry soil). In slightly less P-impoverished soil (> 11 mg P kg− 1 dry soil), phosphatases released by A. cygnorum hydrolysed sufficient organic P that was relatively mobile. Conclusion The reliance on facilitation of P acquisition in A cygnorum depended strongly on location. We demonstrated the exudation of phosphatases, which mobilise inorganic P; this P was adequate for growth when there was sufficient organic P in soil. Facilitation of P acquisition by B. attenuata allowed A. cygnorum to extend its range into severely P-impoverished habitats where it cannot exist without facilitation. This knowledge provides a better understanding of the diversity of P-acquisition strategies in severely P-impoverished environments.

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

Background and aims The characterisation of plant-available phosphorus (P) pools and the assessment of the microbial community in the rhizosheath of cover crops can improve our understanding of plant–microbe interactions and P availability. Methods Mustard (Sinapis alba), phacelia (Phacelia tanacetifolia) and buckwheat (Fagopyrum esculentum) were grown as cover crops before soybean (Glycine max) in an on-farm experiment on a soil low in available P in southwest Germany. The cycling of P through the cover crop biomass and the enzyme-availability of organic P (Porg) pools in the cover crop rhizosheath were characterised. The soil microbial community (PLFA), activity (acid and alkaline phosphomonoesterase, as well as phosphodiesterase), and microbial P were assessed. The abundance of 16S-rRNA and phoD, coding for alkaline phosphomonoesterase in bacteria, were quantified using real-time qPCR. Results Mustard contained the greatest amount of P in its large biomass. In the rhizosheath of all cover crops, the concentration of enzyme-labile Porg was higher than that in the control bulk soil, along with substantial increases of microbial abundance and activity. There were little differences among cover crop species, few changes in the bulk soil and only a limited carryover effect to soybean, except for fungi. Conclusions Turnover of microbial biomass, especially saprotrophic fungi, increased by rhizodeposition of cover crop roots; this was likely responsible for the observed increases in enzyme-available Porg. Microbial function was correlated linearly with microbial biomass, and the data of enzyme activity and phoD did not suggest a difference of their specific activity between bulk and rhizosheath soil.

Belowground plant traits play important roles in plant diversity loss driven by atmospheric nitrogen (N) deposition. However, the way N enrichment shapes plant microhabitats by patterning belowground traits and finally determines aboveground responses is poorly understood. Here, we investigated the rhizosheath trait of 74 plant species in seven N-addition-simulation experiments across multiple grassland ecosystems in China. We found that rhizosheath formation differed among plant functional groups and contributed to changes in plant community composition induced by N enrichment. Compared with forb species, grass and sedge species exhibited more distinct rhizosheaths; moreover, grasses and sedges expanded their rhizosheaths with increasing N-addition rate which allowed them to colonize belowground habitats. Grasses also shaped a different microenvironment around their roots compared with forbs by affecting the physicochemical, biological and stress-avoiding properties of their rhizosphere soil. Rhizosheaths act as a “biofilm-like shield” by accumulating of protective compounds, carboxylic anions and polysaccharides, determined by both plants and microorganisms. This enhanced the tolerance of grasses and sedges to stresses induced by N enrichment. Conversely, forbs lacked the protective rhizosheaths which renders their roots sensitive to stresses induced by N enrichment, thus contributing to their disappearance under N-enriched conditions. This study uncovers the processes by which belowground facilitation and trait matching affects aboveground responses under conditions of N enrichment, which advances our mechanistic understanding of the contribution of competitive exclusion and environmental tolerance to plant diversity loss caused by N deposition.

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.