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Invasive species can cause shifts in vegetation composition and fire regimes by initiating positive vegetation-fire feedbacks. To understand the mechanisms underpinning these shifts, we need to determine how invasive species interact with other species when burned in combination and thus how they may influence net flammability in the communities they invade. Previous studies using litter and ground fuels suggest that flammability of a species mixture is nonadditive and is driven largely by the more-flammable species. However, this nonadditivity has not been investigated in the context of plant invasions nor for canopy fuels. Using whole shoots, we measured the flammability of indigenous-invasive species pairs for six New Zealand indigenous and four globally invasive plant species, along with single-species control burns. Our integrated measure of flammability was clearly nonadditive, and the more-flammable species per pairing had the stronger influence on flammability in 83 % of combinations. The degree of nonadditivity was significantly positively correlated with the flammability difference between the species in a pairing. The strength of nonadditivity differed among individual flammability components. Ignitability and combustibility were strongly determined by the more-flammable species per pair, yet both species contributed more equally to consumability and sustainability. Our results suggest mechanisms by which invasive species entrain positive vegetation-fire feedbacks that alter ecosystem flammability, enhancing their invasion. Of the species tested, Hakea sericea and Ulex europaeus are those most likely to increase the flammability of New Zealand ecosystems and should be priorities for management.

Background and aims Hakea prostrata (Proteaceae) is a highly phosphorus-use-efficient plant native to southwest Australia. It maintains a high photosynthetic rate at low leaf phosphorus (P) and exhibits delayed leaf greening, a convergent adaptation that increases nutrient-use efficiency. This study aimed to provide broad physiological and gene expression profiles across leaf development, uncovering pathways leading from young leaves as nutrient sinks to mature leaves as low-nutrient, energy-transducing sources. Methods To explore gene expression underlying delayed greening, we analysed a de novo transcriptome for H. prostrata across five stages of leaf development. Photosynthesis and respiration rates, and foliar pigment, P and nitrogen (N) concentrations were determined, including the division of P into five biochemical fractions. Key results Transcripts encoding functions associated with leaf structure generally decreased in abundance across leaf development, concomitant with decreases in foliar concentrations of 85% for anthocyanins, 90% for P and 70% for N. The expression of genes associated with photosynthetic function increased during or after leaf expansion, in parallel with increases in photosynthetic pigments and activity, much later in leaf development than in species that do not have delayed greening. As leaves developed, transcript abundance for cytosolic and mitochondrial ribosomal proteins generally declined, whilst transcripts for chloroplast ribosomal proteins increased. Conclusions There was a much longer temporal separation of leaf cell growth from chloroplast development in H. prostrata than is found in species that lack delayed greening. Transcriptome-guided analysis of leaf development in H. prostrata provided insight into delayed greening as a nutrient-saving strategy in severely phosphorus-impoverished landscapes.

Background and aims Phosphorus (P) is an essential plant nutrient and integral for crop yield. However, plants adapted to P-impoverished environments, such as Hakea prostrata (Proteaceae), are often sensitive to P supplies that would be beneficial to other plants. The strategies for phosphate uptake and transport in P-sensitive species have received little attention. Methods Using a recently-assembled transcriptome of H. prostrata , we identified 10 putative members of the PHOSPHATE TRANSPORTER1 ( PHT1 ) gene family, which is responsible for inorganic phosphate (Pi) uptake and transport in plants. We examined plant growth, organ P concentrations and the transcript levels for the eight PHT1 members that were expressed in roots of H. prostrata at Pi supplies ranging from P-impoverished to P-excess. Key results Hakea prostrata  plants suppressed cluster root growth above ecologically-relevant Pi supplies, whilst non-cluster root mass ratios were constant. Root P concentrations increased with increasing Pi supply. Of the eight H. prostrata PHT1 genes tested, four had relatively high transcript amounts in young roots suggesting important roles in Pi uptake; however, a maximum five-fold difference in expression between P-impoverished and P-excess conditions indicated a low P-responsiveness for these genes. The HpPHT1;8 and HpPHT1;9 genes were paralogous to Pi-responsive Arabidopsis thaliana PHT1;8 and PHT1;9 orthologues involved in root-to-shoot translocation of P, but only HpPHT1;9 was P responsive. Conclusions An attenuated ability of H. prostrata to regulate PHT1 expression in response to Pi supply is likely responsible for its low capacity to control P uptake and contributes to its high P sensitivity.

In this review, a brief description of the invasive phenomena associated with plants and its consequences to the ecosystem is presented. Five worldwide invasive plants that are a threat to Portugal were selected as an example, and a brief description of each is presented. A full description of their secondary metabolites and biological activity is given, and a resume of the biological activity of extracts is also included. The chemical and pharmaceutical potential of invasive species sensu lato is thus acknowledged. With this paper, we hope to demonstrate that invasive species have potential positive attributes even though at the same time they might need to be controlled or eradicated. Positive attributes include chemical and pharmaceutical properties and developing these could help mitigate the costs of management and eradication.

PurposeRoot exudation of organic acids (OAs) facilitates plant P uptake from soil, playing a key role in rhizosphere nutrient availability. However, OA exudation responses to CO2 concentrations and water availability remain largely untested.MethodsWe examined the effects of CO2 and water on OA exudates in three Australian woodland species: Eucalyptus tereticornis, Hakea sericea and Microlaena stipoides. Seedlings were grown in a glasshouse in low P soil, exposed to CO2 (400 ppm [aCO2] or 540 ppm [eCO2]) and water treatments (100% water holding capacity [high-watered] or 25–50% water holding capacity [low-watered]). After six weeks, we collected OAs from rhizosphere soil (OArhizo) and trap solutions in which washed roots were immersed (OAexuded).ResultsFor E. tereticornis, the treatments changed OArhizo composition, driven by increased malic acid in plants exposed to eCO2 and increased oxalic acid in low-watered plants. For H. sericea, low-watered plants had higher OAexuded per plant (+ 116%) and lower OArhizo per unit root mass (–77%) associated with larger root mass but fewer cluster roots. For M. stipoides, eCO2 increased OAexuded per plant (+ 107%) and per unit root mass (+ 160%), while low-watered plants had higher citric and lower malic acids for OArhizo and OAexuded: changes in OA amounts and composition driven by malic acid were positively associated with soil P availability under eCO2.ConclusionWe conclude that eCO2 and altered water availability shifted OAs in root exudates, modifying plant–soil interactions and the associated carbon and nutrient economy.

Water and nitrogen (N) interact to influence soil N cycling and plant N acquisition. We studied indices of soil N availability and acquisition by woody plant taxa with distinct nutritional specialisations along a north Australian rainfall gradient from monsoonal savanna (1,600-1,300 mm annual rainfall) to semi-arid woodland (600-250 mm). Aridity resulted in increased 'openness' of N cycling, indicated by increasing delta super(15)N sub(soil) and nitrate:ammonium ratios, as plant communities transitioned from N to water limitation. In this context, we tested the hypothesis that delta super(15)N sub(root xylem sap) provides a more direct measure of plant N acquisition than delta super(15)N sub(foliage). We found highly variable offsets between delta super(15)N sub(foliage) and delta super(15)N sub(root xylem sap), both between taxa at a single site (1.3-3.4 ppt) and within taxa across sites (0.8-3.4 ppt). As a result, delta super(15)N sub(foliage) overlapped between N-fixing Acacia and non-fixing Eucalyptus/Corymbia and could not be used to reliably identify biological N fixation (BNF). However, Acacia delta super(15)N sub(root xylem sap) indicated a decline in BNF with aridity corroborated by absence of root nodules and increasing xylem sap nitrate concentrations and consistent with shifting resource limitation. Acacia dominance at arid sites may be attributed to flexibility in N acquisition rather than BNF capacity. delta super(15)N sub(root xylem sap) showed no evidence of shifting N acquisition in non-mycorrhizal Hakea/Grevillea and indicated only minor shifts in Eucalyptus/Corymbia consistent with enrichment of delta super(15)N sub(soil) and/or decreasing mycorrhizal colonisation with aridity. We propose that delta super(15)N sub(root xylem sap) is a more direct indicator of N source than delta super(15)N sub(foliage), with calibration required before it could be applied to quantify BNF.

Adaptation to drought involves complex interactions of traits that vary within and among species. To date, few data are available to quantify within-species variation in functional traits and they are rarely integrated into mechanistic models to improve predictions of species response to climate change.We quantified intraspecific variation in functional traits of two Hakea species growing along an aridity gradient in southeastern Australia. Measured traits were later used to parameterise the model SurEau to simulate a transplantation experiment to identify the limits of drought tolerance.Embolism resistance varied between species but not across populations. Instead, populations adjusted to drier conditions via contrasting sets of trait trade-offs that facilitated homeostasis of plant water status. The species from relatively mesic climate, Hakea dactyloides, relied on tight stomatal control whereas the species from xeric climate, Hakea leucoptera dramatically increased Huber value and leaf mass per area, while leaf area index (LAI) and epidermal conductance (g(min)) decreased. With trait variability, SurEau predicts the plasticity of LAI and g(min) buffers the impact of increasing aridity on population persistence.Knowledge of within-species variability in multiple drought tolerance traits will be crucial to accurately predict species distributional limits.

Nitrogen is quantitatively the most important nutrient that plants acquire from the soil. It is well established that plant roots take up nitrogen compounds of low molecular mass, including ammonium, nitrate, and amino acids. However, in the soil of natural ecosystems, nitrogen occurs predominantly as proteins. This complex organic form of nitrogen is considered to be not directly available to plants. We examined the long-held view that plants depend on specialized symbioses with fungi (mycorrhizas) to access soil protein and studied the woody heathland plant Hakea actites and the herbaceous model plant Arabidopsis thaliana, which do not form mycorrhizas. We show that both species can use protein as a nitrogen source for growth without assistance from other organisms. We identified two mechanisms by which roots access protein. Roots exude proteolytic enzymes that digest protein at the root surface and possibly in the apoplast of the root cortex. Intact protein also was taken up into root cells most likely via endocytosis. These findings change our view of the spectrum of nitrogen sources that plants can access and challenge the current paradigm that plants rely on microbes and soil fauna for the breakdown of organic matter.

Plants respond to soil phosphorus (P) availability by adjusting leaf P among inorganic P (Pi) and organic P fractions (nucleic acids, phospholipids, small metabolites and a residual fraction). We tested whether phylogenetically divergent plants in a biodiversity hotspot similarly adjust leaf P allocation in response to P limitation by sampling along a 2 Myr chronosequence in southwestern Australia where nitrogen (N) limitation transitions to P limitation with increasing soil age. Total P and N, and P allocated to five chemical fractions were determined for photosynthetic organs from Melaleuca systena (Myrtaceae), Acacia rostellifera (Fabaceae) and Hakea prostrata (Proteaceae). Soil characteristics were also determined. Acacia rostellifera maintained phyllode total P and N concentrations at c. 0.5 and 16 mg g−1 DW, respectively, with a constant P-allocation pattern along the chronosequence. H. prostrata leaves allocated less P to Pi, phospholipids and nucleic acids with increasing soil age, while leaf N concentration was constant. M. systena had the greatest variation in allocating leaf P, whereas leaf N concentration decreased 20% along the chronosequence. Variation in P-allocation patterns was only partially conserved among species along the chronosequence. Such variation could have an impact on species distribution and contribute to species richness in P-limited environments.

Despite its agronomic importance, the metabolic networks mediating phosphorus (P) remobilization during plant senescence are poorly understood. Highly efficient P remobilization (~85%) from senescing leaves and proteoid roots of harsh hakea (Hakea prostrata), a native ‘extremophile’ plant of south-western Australia, was linked with striking up-regulation of cell wall-localized and intracellular acid phosphatase (APase) and RNase activities. Non-denaturing PAGE followed by in-gel APase activity staining revealed senescence-inducible 120kDa and 60kDa intracellular APase isoforms, whereas only the 120kDa isoform was detected in corresponding cell wall fractions. Kinetic and immunological properties of the 120kDa and 60kDa APases partially purified from senescing leaves indicated that they are purple acid phosphatases (PAPs). Results obtained with cell wall-targeted hydrolases of harsh hakea were corroborated using Arabidopsis thaliana in which an ~200% increase in cell wall APase activity during leaf senescence was paralleled by accumulation of immunoreactive 55kDa AtPAP26 polypeptides. Senescing leaves of an atpap26 T-DNA insertion mutant displayed a >90% decrease in cell wall APase activity. Previous research established that senescing leaves of atpap26 plants exhibited a similar reduction in intracellular (vacuolar) APase activity, while displaying markedly impaired P remobilization efficiency and delayed senescence. It is hypothesized that up-regulation and dual targeting of PAPs and RNases to the cell wall and vacuolar compartments make a crucial contribution to highly efficient P remobilization that dominates the P metabolism of senescing tissues of harsh hakea and Arabidopsis. To the best of the authors’ knowledge, the apparent contribution of cell wall-targeted hydrolases to remobilizing key macronutrients such as P during senescence has not been previously suggested. Targeting of senescence-inducible acid phosphatases and RNases to the cell wall and vacuolar compartments appears to make a crucial contribution to efficient P remobilization networks of senescing tissues of Hakea prostrata and Arabidopsis.