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Nonstructural carbohydrate (NSC) concentrations might reflect the strategies described in the leaf economic spectrum (LES) due to their dependence on photosynthesis and respiration. We examined if NSC concentrations correlate with leaf structure, chemistry, and physiology traits for 114 species from 19 sites and 5 biomes around the globe. Total leaf NSC concentrations varied greatly from 16 to 199 mg g -1 dry mass and were mostly independent of leaf gas exchange and the LES traits. By contrast, leaf NSC residence time was shorter in species with higher rates of photosynthesis, following the fast-slow strategies in the LES. An average leaf held an amount of NSCs that could sustain one night of leaf respiration and could be replenished in just a few hours of photosynthesis under saturating light, indicating that most daily carbon gain is exported. Our results suggest that NSC export is clearly linked to the economics of return on resource investment., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

An approach to improving radiation use efficiency (RUE) in wheat is to screen for variability in rates of leaf respiration in darkness (Rdark). We used a high-throughput system to quantify variation in Rdark among a diverse range of spring wheat genotypes (301 lines) grown in two countries (Mexico and Australia) and two seasons (2017 and 2018), and in doing so quantify the relative importance of genotype (G) and environment (E) in influencing variations in leaf Rdark. Through careful design, residual (unexplained) variation represented less than 10% of the total observed. Up to a third of the variation in Rdark (and related traits) was under genetic control. This suggests opportunities for breeders to use Rdark as a novel selection tool. In addition, E accounted for more than half of the total variation in area-based rates of Rdark. Here, the day of measurement was crucial, suggesting that day-to-day variations in the environment influence rates of Rdark measured at a common temperature. Overall, this study provides new insights into the role G and E play in determining variation in rates of leaf Rdark of one of the most important cereal crops, with implications for future improvements in carbon use efficiency and yield., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

A significant warming effect on arctic tundra is greening. Although this increase in predominantly woody vegetation has been linked to increases in gross primary productivity, increasing temperatures also stimulate ecosystem respiration. We present a novel analysis from small-scale plot measurements showing that the shape of the temperature- and light-dependent sink-to-source threshold (where net ecosystem exchange (NEE) equals zero) differs between two tussock tundra ecosystems differing in leaf area index (LAI). At the higher LAI site, the threshold is exceeded (i.e the ecosystem becomes a source) at relatively higher temperatures under low light but at lower temperatures under high light. At the lower LAI site, the threshold is exceeded at relatively lower temperatures under low light but at higher temperatures under high light. We confirmed this response at a single site where LAI was experimentally increased. This suggests the carbon balance of the tundra may be sensitive to small increases in temperature under low light, but that this effect may be significantly offset by increases in LAI. Importantly, we found that this LAI effect is reversed under high light, and so in a warming tundra, greater vegetation cover could have a progressively negative effect on net carbon uptake., (© 2024. The Author(s).)

Masting, the synchronous, highly variable flowering across years by a population of perennial plants, has been reported to be precipitated by various factors including nitrogen levels, drought conditions, and spring and summer temperatures. However, the molecular mechanism leading to the initiation of flowering in masting plants in particular years remains largely unknown, despite the potential impact of climate change on masting phenology. We studied genes controlling flowering in the alpine snow tussock Chionochloa pallens (Poaceae), a strongly masting perennial grass. We used a range of in situ and manipulated plants to obtain leaf samples from tillers (shoots) which subsequently remained vegetative or flowered. Here, we show that a novel orthologue of TERMINAL FLOWER 1 (TFL1; normally a repressor of flowering in other species) promotes the induction of flowering in C. pallens (hence Anti-TFL1), a conclusion supported by structural, functional and expression analyses. Global transcriptomic analysis indicated differential expression of CpTPS1, CpGA20ox1, CpREF6 and CpHDA6, emphasizing the role of endogenous cues and epigenetic regulation in terms of responsiveness of plants to initiate flowering. Our molecular-based study provides insights into the cellular mechanism of flowering in masting plants and will supplement ecological and statistical models to predict how masting will respond to global climate change., (© 2021 John Wiley & Sons Ltd.)

Mast flowering (or masting) is synchronous, highly variable flowering among years in populations of perennial plants. Despite having widespread consequences for seed consumers, endangered fauna and human health, masting is hard to predict. While observational studies show links to various weather patterns in different plant species, the mechanism(s) underpinning the regulation of masting is still not fully explained. We studied floral induction in Celmisia lyallii (Asteraceae), a mast flowering herbaceous alpine perennial, comparing gene expression in flowering and nonflowering plants. We performed translocation experiments to induce the floral transition in C. lyallii plants followed by both global and targeted expression analysis of flowering-pathway genes. Differential expression analysis showed elevated expression of ClSOC1 and ClmiR172 (promoters of flowering) in leaves of plants that subsequently flowered, in contrast to elevated expression of ClAFT and ClTOE1 (repressors of flowering) in leaves of plants that did not flower. The warm summer conditions that promoted flowering led to differential regulation of age and hormonal pathway genes, including ClmiR172 and ClGA20ox2, known to repress the expression of floral repressors and permit flowering. Upregulated expression of epigenetic modifiers of floral promoters also suggests that plants may maintain a novel "summer memory" across years to induce flowering. These results provide a basic mechanistic understanding of floral induction in masting plants and evidence of their ability to imprint various environmental cues to synchronize flowering, allowing us to better predict masting events under climate change., (© 2021 John Wiley & Sons Ltd.)

Short-term temperature response curves of leaf dark respiration (R-T) provide insights into a critical process that influences plant net carbon exchange. This includes how respiratory traits acclimate to sustained changes in the environment. Our study analysed 860 high-resolution R-T (10-70°C range) curves for: (a) 62 evergreen species measured in two contrasting seasons across several field sites/biomes; and (b) 21 species (subset of those sampled in the field) grown in glasshouses at 20°C : 15°C, 25°C : 20°C and 30°C : 25°C, day : night. In the field, across all sites/seasons, variations in R 25 (measured at 25°C) and the leaf T where R reached its maximum (T max ) were explained by growth T (mean air-T of 30-d before measurement), solar irradiance and vapour pressure deficit, with growth T having the strongest influence. R 25 decreased and T max increased with rising growth T across all sites and seasons with the single exception of winter at the cool-temperate rainforest site where irradiance was low. The glasshouse study confirmed that R 25 and T max thermally acclimated. Collectively, the results suggest: (1) thermal acclimation of leaf R is common in most biomes; and (2) the high T threshold of respiration dynamically adjusts upward when plants are challenged with warmer and hotter climates., (© 2020 The Authors New Phytologist © 2020 New Phytologist Trust.)

Background: Mast flowering ('masting') is characterized by mass synchronized flowering at irregular intervals in populations of perennial plants over a wide geographical area, resulting in irregular high seed production. While masting is a global phenomenon, it is particularly prevalent in the alpine flora of New Zealand. Increases in global temperature may alter the masting pattern, affecting wider communities with a potential impact on plant-pollinator interactions, seed set and food availability for seed-consuming species., Scope: This review summarizes an ecological temperature model (ΔT) that is being used to predict the intensity of a masting season. We introduce current molecular studies on flowering and the concept of an 'epigenetic summer memory' as a driver of mast flowering. We propose a hypothetical model based on temperature-associated epigenetic modifications of the floral integrator genes FLOWERING LOCUS T, FLOWERING LOCUS C and SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1., Conclusions: Genome-wide transcriptomic and targeted gene expression analyses are needed to establish the developmental and physiological processes associated with masting. Such analyses may identify changes in gene expression that can be used to predict the intensity of a forthcoming masting season, as well as to determine the extent to which climate change will influence the mass synchronized flowering of masting species, with downstream impacts on their associated communities., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Variation in traits within a plant species contributes to differences in soil physicochemistry and rhizosphere microbial communities. However, how intraspecific variation in plant responses to nitrogen (N) shapes these communities remains unclear. We studied whether plant responses to organic and inorganic N forms vary among genotypes, and if these responses were associated with variation in root-associated communities. We investigated how the root microbiomes of two Pinus radiata D. Don genotypes were altered by two years of N-fertilisation in field conditions. We characterised rhizosphere bacterial and fungal communities, as well as root-associated fungal communities, of trees receiving yearly additions of NH4NO3 or L-arginine, and control trees. We also measured plant traits and rhizosphere soil physicochemical properties. Two main findings emerged: (i) N form and tree genotype affected soil physicochemical properties as well as plant measures, and these responses were associated with variation in microbial communities, and (ii) rhizosphere and root-associated communities differed in their responses to N form and host genotype. Our results suggest that N forms have different influences on N and carbon dynamics at the plant-soil interface by inducing root-mediated responses that are associated with shifts in the root microbiome such that communities more closely associated with roots are more sensitive to genotype-specific responses.

Enhanced pond systems (EPS) consist of a series of ponds that have been designed to work in synergy to provide both cost-effective enhanced wastewater treatment and resource recovery, in the form of algal biomass, for beneficial reuse. Due to the limited number of full-scale EPS systems worldwide, our understanding of factors governing both enhanced wastewater treatment and resource recovery is limited. This paper investigates the seasonal performance of a full-scale municipal wastewater EPS with respect to nutrient removal from the liquid fraction, microalgal biomass production and subsequent removal through the system. In the high rate algal pond both microalgal productivity (determined as organic matter and chlorophyll a biomass) and NH 4 -N removal varied seasonally, with significantly higher biomass and removal rates in summer than in spring (p < 0.05) or winter (p < 0.01). Microalgal biomass was not successfully harvested in the algal harvester pond (AHP), most likely due to poor flocc formation coupled with relatively short hydraulic residence time (HRT). High percentage removal rates, from sedimentation and zooplankton grazing, were achieved in the maturation pond (MP) series, particularly in winter and spring. However, in summer decreased efficiency of biomass removal and the growth of new microalgal species suggests that summer-time HRT in the MPs could be shortened. Further modifications to the operation of the AHP, seasonal changes in the HRT of the MPs and potential harvesting of zooplankton grazers are all potential strategies for improving resource recovery and producing a higher quality final discharge effluent., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

A central challenge in community ecology is understanding the role that phenotypic variation among genotypes plays in structuring host-associated communities. While recent studies have investigated the relationship between plant genotype and the composition of soil microbial communities, the effect of genotype-by-environment interactions on the plant microbiome remains unclear. In this study, we assessed the influence of tree genetics (G), nitrogen (N) form and genotype-by-environment interaction (G x N) on the composition of the root microbiome. Rhizosphere communities (bacteria and fungi) and root-associated fungi (including ectomycorrhizal and saprotrophic guilds) were characterised in two genotypes of Pinus radiata with contrasting physiological responses to exogenous organic or inorganic N supply. Genotype-specific responses to N form influenced the composition of the root microbiome. Specifically, (1) diversity and composition of rhizosphere bacterial and root-associated fungal communities differed between genotypes that had distinct responses to N form, (2) shifts in the relative abundance of individual taxa were driven by the main effects of N form or host genotype and (3) the root microbiome of the P. radiata genotype with the most divergent growth responses to organic and inorganic N was most sensitive to differences in N form. Our results show that intraspecific variation in tree response to N form has significant consequences for the root microbiome of P. radiata, demonstrating the importance of genotype-by-environment interactions in shaping host-associated communities.

Land-atmosphere exchanges influence atmospheric CO 2 . Emphasis has been on describing photosynthetic CO 2 uptake, but less on respiration losses. New global datasets describe upper canopy dark respiration (R d ) and temperature dependencies. This allows characterisation of baseline R d , instantaneous temperature responses and longer-term thermal acclimation effects. Here we show the global implications of these parameterisations with a global gridded land model. This model aggregates R d to whole-plant respiration R p , driven with meteorological forcings spanning uncertainty across climate change models. For pre-industrial estimates, new baseline R d increases R p and especially in the tropics. Compared to new baseline, revised instantaneous response decreases R p for mid-latitudes, while acclimation lowers this for the tropics with increases elsewhere. Under global warming, new R d estimates amplify modelled respiration increases, although partially lowered by acclimation. Future measurements will refine how R d aggregates to whole-plant respiration. Our analysis suggests R p could be around 30% higher than existing estimates.

In the present study we investigated variations in leaf respiration in darkness (RD) and light (RL), and associated traits in response to season, and along a gradient of soil moisture, in Mediterranean woodland dominated by holm oak (Quercus ilex L.) in central and north-eastern Spain respectively. On seven occasions during the year in the central Spain site, and along the soil moisture gradient in north-eastern Spain, we measured rates of leaf RD, RL (using the Kok method), light-saturated photosynthesis (A) and related light response characteristics, leaf mass per unit area (MA) and leaf nitrogen (N) content. At the central Spain site, significant seasonal changes in soil water content and ambient temperature (T) were associated with changes in MA, foliar N, A and stomatal conductance. RD measured at the prevailing daily T and in instantaneous R-T responses, displayed signs of partial acclimation and was not significantly affected by time of year. RL was always less than, and strongly related to, RD, and RL/RD did not vary significantly or systematically with seasonal changes in T or soil water content. Averaged over the year, RL/RD was 0.66±0.05s.e. (n=14) at the central Spain site. At the north-eastern Spain site, the soil moisture gradient was characterised by increasing MA and RD, and reduced foliar N, A, and stomatal conductance as soil water availability decreased. Light inhibition of R occurred across all sites (mean RL/RD=0.69±0.01s.e. (n=18)), resulting in ratios of RL/A being lower than for RD/A. Importantly, the degree of light inhibition was largely insensitive to changes in soil water content. Our findings provide evidence for a relatively constrained degree of light inhibition of R (RL/RD ~ 0.7, or inhibition of ~30%) across gradients of water availability, although the combined impacts of seasonal changes in both T and soil water content increase the range of values expressed. The findings thus have implications in terms of the assumptions made by predictive models that seek to account for light inhibition of R, and for our understanding of how environmental gradients impact on leaf trait relationships in Mediterranean plant communities.

In the last decade, studies have focused on identifying the most suitable microalgal species for coupled high rate algal pond (HRAP) wastewater treatment and resource recovery. However, one of the challenges facing outdoor HRAP systems is maintaining microalgal species dominance. By increasing our understanding of the environmental drivers of microalgal community composition within the HRAP environment, it may be possible to manipulate the system in such a way to favour the growth of desirable species. In this paper, we investigate the microalgal community composition in two full-scale HRAPs over a 23-month period. We compare wastewater treatment performance between dominant species and identify the environmental drivers that trigger change in community composition. A total of 33 microalgal species were identified over the 23-month period but species richness (the number of species present at any given time) was low and was not related to either productivity or nutrient removal efficiency. Species turnover of the dominant microalgae happened rapidly, typically <1 week. Changes in the influent NH 4 -N concentration and zooplankton grazer numbers were significantly associated with species turnover, accounting for 80% of the changes in dominant species throughout the 23-month study period. Both nutrient removal and biomass production did not differ between the two HRAPs when the dominant species was the same or differed in the two ponds. These results suggest that microalgal functional groups are more important than individual species for full-scale HRAP performance. This study has increased our understanding of some of the environmental drivers of the microalgae within the HRAP environment, which may assist with improving wastewater treatment and resource recovery., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Nitrogen (N) and phosphorus (P) have key roles in leaf metabolism, resulting in a strong coupling of chemical composition traits to metabolic rates in field-based studies. However, in such studies, it is difficult to disentangle the effects of nutrient supply per se on trait-trait relationships. Our study assessed how high and low N (5 mM and 0.4 mM, respectively) and P (1 mM and 2 μM, respectively) supply in 37 species from six plant functional types (PTFs) affected photosynthesis (A) and respiration (R) (in darkness and light) in a controlled environment. Low P supply increased scaling exponents (slopes) of area-based log-log A-N or R-N relationships when N supply was not limiting, whereas there was no P effect under low N supply. By contrast, scaling exponents of A-P and R-P relationships were altered by P and N supply. Neither R : A nor light inhibition of leaf R was affected by nutrient supply. Light inhibition was 26% across nutrient treatments; herbaceous species exhibited a lower degree of light inhibition than woody species. Because N and P supply modulates leaf trait-trait relationships, the next generation of terrestrial biosphere models may need to consider how limitations in N and P availability affect trait-trait relationships when predicting carbon exchange., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

Root system architecture is the spatial arrangement of roots that impacts the capacity of plants to access nutrients and water. We employed pharmacologically generated morphological and molecular phenotypes and used in situ 15N isotope labelling, to investigate whether contrasting root traits are of functional interest in relation to nitrate acquisition. Brassica napus L. were grown in solidified phytogel culture media containing 1mM KNO3 and treated with the cytokinin, 6-benzylaminopurine, the cytokinin antagonist, PI-55, or both in combination. The pharmacological treatments inhibited root elongation relative to the control. The contrasting root traits induced by PI-55 and 6-benzylaminopurine were strongly related to 15N uptake rate. Large root proliferation led to greater 15N cumulative uptake rather than greater 15N uptake efficiency per unit root length, due to a systemic response in the plant. This relationship was associated with changes in C and N resource distribution between the shoot and root, and in expression of BnNRT2.1, a nitrate transporter. The root:shoot biomass ratio was positively correlated with 15N cumulative uptake, suggesting the functional utility of root investment for nutrient acquisition. These results demonstrate that root proliferation in response to external nitrate is a behaviour which integrates local N availability and the systemic N status of the plant.

Background and Aims: The efficiency of N assimilation in response to defoliation is a critical component of plant regrowth and forage production. The aim of this research was to test the effect of the internal C/N balance on NO3- assimilation and to estimate the associated cytokinin signals following defoliation of perennial ryegrass ( Lolium perenne L. 'Grasslands Nui') plants., Methods: Plants, manipulated to have contrasting internal N content and contrasting availability of water soluble carbohydrates (WSCs), were obtained by exposure to either continuous light or short days (8:16 h light-dark), and watered with modified N-free Hoagland medium containing either high (5 m m ) or low (50 μ m ) NO3- as sole N source. Half of the plants were defoliated and the root, sheath and leaf tissue were harvested at 8, 24 and 168 h after cutting. The spatiotemporal changes in WSCs, synthesis of amino acids and associated cytokinin content were recorded after cutting., Key Results: Leaf regrowth following defoliation involved changes in the low- and high-molecular weight WSCs. The extent of the changes and the partitioning of the WSC following defoliation were dependant on the initial WSC levels and the C and N availability. Cytokinin levels varied in the sheath and root as early as 8 h following defoliation and preceded an overall increase in amino acids at 24 h. Subsequently, negative feedback brought the amino acid response back towards pre-defoliation levels within 168 h after cutting, a response that was under control of the C/N ratio., Conclusions: WSC remobilization in the leaf is coordinated with N availability to the root, potentially via a systemic cytokinin signal, leading to efficient N assimilation in the leaf and the sheath tissues and to early leaf regrowth following defoliation., (© The Author 2017. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com)

Leaf dark respiration (R dark ) represents an important component controlling the carbon balance in tropical forests. Here, we test how nitrogen (N) and phosphorus (P) affect R dark and its relationship with photosynthesis using three widely separated tropical forests which differ in soil fertility. R dark was measured on 431 rainforest canopy trees, from 182 species, in French Guiana, Peru and Australia. The variation in R dark was examined in relation to leaf N and P content, leaf structure and maximum photosynthetic rates at ambient and saturating atmospheric CO 2 concentration. We found that the site with the lowest fertility (French Guiana) exhibited greater rates of R dark per unit leaf N, P and photosynthesis. The data from Australia, for which there were no phylogenetic overlaps with the samples from the South American sites, yielded the most distinct relationships of R dark with the measured leaf traits. Our data indicate that no single universal scaling relationship accounts for variation in R dark across this large biogeographical space. Variability between sites in the absolute rates of R dark and the R dark  : photosynthesis ratio were driven by variations in N- and P-use efficiency, which were related to both taxonomic and environmental variability., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

The mechanisms linking C/N balance to N uptake and assimilation are central to plant responses to changing soil nutrient levels. Defoliation and subsequent regrowth of grasses both impact C partitioning, thereby creating a significant point of interaction with soil N availability. Using defoliation as an experimental treatment, we investigated the dynamic relationships between plant carbohydrate status and NO3--responsive uptake systems, transporter gene expression, and nitrate assimilation in Lolium perenne L. High- and low-affinity NO3- uptake was reduced in an N-dependent manner in response to a rapid and large shift in carbohydrate remobilization triggered by defoliation. This reduction in NO3- uptake was rescued by an exogenous glucose supplement, confirming the carbohydrate dependence of NO3- uptake. The regulation of NO3- uptake in response to the perturbation of the plant C/N ratio was associated with changes in expression of putative high- and low-affinity NO3- transporters. Furthermore, NO3- assimilation appears to be regulated by the C-N status of the plant, implying a mechanism that signals the availability of C metabolites for NO3- uptake and assimilation at the whole-plant level. We also show that cytokinins may be involved in the regulation of N acquisition and assimilation in response to the changing plant C/N ratio., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

The ability to quantify the impacts of changing management practices on the components of net ecosystem carbon balance (N B ) is required to forecast future changes in soil carbon stocks and potential feedbacks on atmospheric CO 2 concentrations. In this study we investigated seasonal changes on the components of net ecosystem carbon balance resulting from the application of irrigation and nitrogen fertiliser to a temperate grassland in New Zealand where we simulated grazing events. We made seasonal measurements of the components of N B using chamber measurements in field plots with and without irrigation and addition of nitrogen fertiliser. We developed models to determine the physiological responses of gross canopy photosynthesis (A), leaf respiration (R L ) and soil respiration (R S ) to soil and air temperature, soil water content and irradiance and we estimated annual N B for the first year after treatments were applied. Overall, irrigation and nitrogen addition had a synergistic effect to increase annual estimates of above-ground components of carbon balance (A, R L and carbon exported through simulated grazing, F export ), but there was no effect from adding nitrogen alone. Annual R S remained unchanged between treatments. The treatments resulted in increases in above-ground biomass production, but, with the high intensity of simulated grazing, these were not sufficient to offset ecosystem carbon losses, so all treatments remained a net source of carbon. There were no significant differences between treatments and annual N B ranged from -540gCm -2 y -1 for the treatment with no irrigation and no nitrogen addition and -284gCm -2 y -1 for the treatment with irrigation and nitrogen addition. Our findings from the first year of the treatments quantify the net benefits of addition of irrigation and nitrogen on increasing above-ground production for animal feed but show that this did not lead to a net increase carbon input to the soil., (Copyright © 2016 Elsevier B.V. All rights reserved.)

High-temperature tolerance in plants is important in a warming world, with extreme heat waves predicted to increase in frequency and duration, potentially leading to lethal heating of leaves. Global patterns of high-temperature tolerance are documented in animals, but generally not in plants, limiting our ability to assess risks associated with climate warming. To assess whether there are global patterns in high-temperature tolerance of leaf metabolism, we quantified T crit (high temperature where minimal chlorophyll a fluorescence rises rapidly and thus photosystem II is disrupted) and T max (temperature where leaf respiration in darkness is maximal, beyond which respiratory function rapidly declines) in upper canopy leaves of 218 plant species spanning seven biomes. Mean site-based T crit values ranged from 41.5 °C in the Alaskan arctic to 50.8 °C in lowland tropical rainforests of Peruvian Amazon. For T max , the equivalent values were 51.0 and 60.6 °C in the Arctic and Amazon, respectively. T crit and T max followed similar biogeographic patterns, increasing linearly (˜8 °C) from polar to equatorial regions. Such increases in high-temperature tolerance are much less than expected based on the 20 °C span in high-temperature extremes across the globe. Moreover, with only modest high-temperature tolerance despite high summer temperature extremes, species in mid-latitude (~20-50°) regions have the narrowest thermal safety margins in upper canopy leaves; these regions are at the greatest risk of damage due to extreme heat-wave events, especially under conditions when leaf temperatures are further elevated by a lack of transpirational cooling. Using predicted heat-wave events for 2050 and accounting for possible thermal acclimation of T crit and T max , we also found that these safety margins could shrink in a warmer world, as rising temperatures are likely to exceed thermal tolerance limits. Thus, increasing numbers of species in many biomes may be at risk as heat-wave events become more severe with climate change., (© 2016 John Wiley & Sons Ltd.)

We measured a diverse range of foliar characteristics in shrub and tree species in temperate rainforest communities along a soil chronosequence (six sites from 8 to 120000 years) and used multilevel model analysis to attribute the proportion of variance for each trait into genetic (G, here meaning species-level), environmental (E) and residual error components. We hypothesised that differences in leaf traits would be driven primarily by changes in soil nutrient availability during ecosystem progression and retrogression. Several leaf structural, chemical and gas-exchange traits were more strongly driven by G than E effects. For leaf mass per unit area (MA), foliar [N], net CO2 assimilation and dark respiration rates and foliar carbohydrate concentration, the G component accounted for 60-87% of the total variance, with the variability associated with plot, the E effect, much less important. Other traits, such as foliar [P] and N:P, displayed strong E and residual effects. Analyses revealed significant reductions in the slopes of G-only bivariate relationships when compared with raw relationships, indicating that a large proportion of trait-trait relationships is species based, and not a response to environment per se. This should be accounted for when assessing the mechanistic basis for using such relationships in order to make predictions of responses of plants to short-term environmental change.

Plant respiration constitutes a massive carbon flux to the atmosphere, and a major control on the evolution of the global carbon cycle. It therefore has the potential to modulate levels of climate change due to the human burning of fossil fuels. Neither current physiological nor terrestrial biosphere models adequately describe its short-term temperature response, and even minor differences in the shape of the response curve can significantly impact estimates of ecosystem carbon release and/or storage. Given this, it is critical to establish whether there are predictable patterns in the shape of the respiration-temperature response curve, and thus in the intrinsic temperature sensitivity of respiration across the globe. Analyzing measurements in a comprehensive database for 231 species spanning 7 biomes, we demonstrate that temperature-dependent increases in leaf respiration do not follow a commonly used exponential function. Instead, we find a decelerating function as leaves warm, reflecting a declining sensitivity to higher temperatures that is remarkably uniform across all biomes and plant functional types. Such convergence in the temperature sensitivity of leaf respiration suggests that there are universally applicable controls on the temperature response of plant energy metabolism, such that a single new function can predict the temperature dependence of leaf respiration for global vegetation. This simple function enables straightforward description of plant respiration in the land-surface components of coupled earth system models. Our cross-biome analyses shows significant implications for such fluxes in cold climates, generally projecting lower values compared with previous estimates.

The efficiency of inorganic nitrogen (N) assimilation is a critical component of fertilizer use by plants and of forage production in Lolium perenne, an important pasture species worldwide. We present a spatiotemporal description of nitrate use efficiency in terms of metabolic responses and carbohydrate remobilization, together with components of cytokinin signal transduction following nitrate addition to N-impoverished plants. Perennial ryegrass (L. perenne cv. Grasslands Nui) plants were grown for 10 weeks in unfertilized soil and then treated with nitrate (5 mM) hydroponically. Metabolomic analysis by gas chromatography-mass spectrometry and liquid chromatography-tandem mass spectrometry revealed a dynamic interaction between N and carbon metabolism over a week-long time course represented by the relative abundance of amino acids, tricarboxylic acid intermediates and stored water-soluble carbohydrates (WSCs). The initial response to N addition was characterized by a rapid remobilization of carbon stores from the low-molecular weight WSC, along with an increase in N content and assimilation into free amino acids. Subsequently, the shoot became the main source of carbon through remobilization of a large pool of high-molecular weight WSC. Associated quantification of cytokinin levels and expression profiling of putative cytokinin response regulator genes by quantitative reverse transcription polymerase chain reaction support a role for cytokinin in the mediation of the response to N addition in perennial ryegrass. The presence of high levels of cis-zeatin-type cytokinins is discussed in the context of hormonal homeostasis under the stress of steady-state N deficiency., (© 2015 Scandinavian Plant Physiology Society.)

With microalgal biofuels currently receiving much attention, there has been renewed interest in the combined use of high rate algal ponds (HRAP) for wastewater treatment and biofuel production. This combined use of HRAPs is considered to be an economically feasible option for biofuel production, however, increased microalgal productivity and nutrient removal together with reduced capital costs are needed before it can be commercially viable. Despite HRAPs being an established technology, microalgal photosynthesis and productivity is still limited in these ponds and is well below the theoretical maximum. This paper critically evaluates the parameters that limit microalgal light absorption and photosynthesis in wastewater HRAPs and examines biological, chemical and physical options for improving light absorption and utilisation, with the view of enhancing biomass production and nutrient removal., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Leaf dark respiration (Rdark ) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of Rdark and associated leaf traits. Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in Rdark . Area-based Rdark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8-28°C). By contrast, Rdark at a standard T (25°C, Rdark (25) ) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher Rdark (25) at a given photosynthetic capacity (Vcmax (25) ) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark (25) values at any given Vcmax (25) or [N] were higher in herbs than in woody plants. The results highlight variation in Rdark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of Rdark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs)., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

The combined use of high rate algal ponds (HRAPs) for wastewater treatment and commercial algal production is considered to be an economically viable option. However, microalgal photosynthesis and biomass productivity is constrained in HRAPs due to light limitation. This paper investigates how the light climate in the HRAP can be modified through changes in pond depth, hydraulic retention time (HRT) and light/dark turnover rate and how this impacts light absorption and utilisation by the microalgae. Wastewater treatment HRAPs were operated at three different pond depth and HRT during autumn. Light absorption by the microalgae was most affected by HRT, significantly decreasing with increasing HRT, due to increased internal self-shading. Photosynthetic performance (as defined by Pmax, Ek and α), significantly increased with increasing pond depth and decreasing HRT. Despite this, increasing pond depth and/or HRT, resulted in decreased pond light climate and overall integrated water column net oxygen production. However, increased light/dark turnover was able to compensate for this decrease, bringing the net oxygen production in line with shallower ponds operated at shorter HRT. On overcast days, modelled daily net photosynthesis significantly increased with increased light/dark turnover, however, on clear days such increased turnover did not enhance photosynthesis. This study has showed that light absorption and photosynthetic performance of wastewater microalgae can be modified through changes to pond depth, HRT and light/dark turnover., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Carbon limitation in domestic wastewater high rate algal ponds is thought to constrain microalgal photo-physiology and productivity, particularly in summer. This paper investigates the effects of CO₂ addition along a pH gradient on the performance of wastewater microalgae in high rate algal mesocosms. Performance was measured in terms of light absorption, electron transport rate, photosynthetic efficiency, biomass production and nutrient removal efficiency. Light absorption by the microalgae increased by up to 128% with increasing CO₂ supply, while a reduction in the package effect meant that there was less internal self-shading thereby increasing the efficiency of light absorption. CO₂ augmentation increased the maximum rate of both electron transport and photosynthesis by up to 256%. This led to increased biomass, with the highest yield occurring at the highest dissolved inorganic carbon/lowest pH combination tested (pH 6.5), with a doubling of chlorophyll-a (Chl-a) biomass while total microalgal biovolume increased by 660% in Micractinium bornhemiense and by 260% in Pediastrum boryanum dominated cultures. Increased microalgal biomass did not off-set the reduction in ammonia volatilisation in the control and overall nutrient removal was lower with CO₂ than without. Microalgal nutrient removal efficiency decreased as pH decreased and may have been related to decreased Chl-a per cell. This experiment demonstrated that CO₂ augmentation increased microalgal biomass in two distinct communities, however, care must be taken when interpreting results from standard biomass measurements with respect to CO₂ augmentation., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

When wastewater treatment high rate algal ponds (HRAP) are coupled with resource recovery processes, such as biofuel production, short hydraulic retention times (HRTs) are often favoured to increase the microalgal biomass productivity. However, short HRT can result in increased nutrient load to the HRAP which may negatively impact on the performance of the microalgae. This paper investigate the effects of high (NH4-N mean concentration 39.7 ± 17.9 g m(-3)) and moderate ((NH4-N mean concentration 19.9 ± 8.9 g m(-3)) nutrient loads and short HRT on the performance of microalgae with respect to light absorption, photosynthesis, biomass production and nutrient removal in pilot-scale (total volume 8 m(3)) wastewater treatment HRAPs. Microalgal biomass productivity was significantly higher under high nutrient loads, with a 133% and 126% increase in the chlorophyll-a and VSS areal productivities, respectively. Microalgae were more efficient at assimilating NH4-N from the wastewater under higher nutrient loads compared to moderate loads. Higher microalgal biomass with increased nutrient load resulted in increased light attenuation in the HRAP and lower light absorption efficiency by the microalgae. High nutrient loads also resulted in improved photosynthetic performance with significantly higher maximum rates of electron transport, oxygen production and quantum yield. This experiment demonstrated that microalgal productivity and nutrient removal efficiency were not inhibited by high nutrient loads, however, higher loads resulted in lower water quality in effluent discharge., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Laminar flows are a common problem in high rate algal ponds (HRAP) due to their long channels and gentle mixing by a single paddlewheel. Sustained laminar flows may modify the amount of light microalgal cells are exposed to, increase the boundary layer between the cell and the environment and increase settling out of cells onto the pond bottom. To date, there has been little focus on the effects of the time between mixing events (frequency of mixing) on the performance of microalgae in wastewater treatment HRAPs. This paper investigates the performance of three morphologically distinct microalgae in wastewater treatment high rate algal mesocosms operated at four different mixing frequencies (continuous, mixed every 45 min, mixed every 90 min and no mixing). Microalgal performance was measured in terms of biomass concentration, nutrient removal efficiency, light utilisation and photosynthetic performance. Microalgal biomass increased significantly with increasing mixing frequency for the two colonial species but did not differ for the single celled species. All three species were more efficient at NH4-N uptake as the frequency of mixing increased. Increased frequency of mixing supported larger colonies with improved harvest-ability by gravity but at the expense of efficient light absorption and maximum rate of photosynthesis. However, maximum quantum yield was highest in the continuously mixed cultures due to higher efficiency of photosynthesis under light limited conditions. Based on these results, higher microalgal productivity, improved wastewater treatment and better gravity based harvest-ability can be achieved with the inclusion of more mixing points and reduced laminar flows in full-scale HRAP., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Despite concern about the status of carbon (C) in the Arctic tundra, there is currently little information on how plant respiration varies in response to environmental change in this region. We quantified the impact of long-term nitrogen (N) and phosphorus (P) treatments and greenhouse warming on the short-term temperature (T) response and sensitivity of leaf respiration (R), the high-T threshold of R, and associated traits in shoots of the Arctic shrub Betula nana in experimental plots at Toolik Lake, Alaska. Respiration only acclimated to greenhouse warming in plots provided with both N and P (resulting in a ~30% reduction in carbon efflux in shoots measured at 10 and 20 °C), suggesting a nutrient dependence of metabolic adjustment. Neither greenhouse nor N+P treatments impacted on the respiratory sensitivity to T (Q10 ); overall, Q10 values decreased with increasing measuring T, from ~3.0 at 5 °C to ~1.5 at 35 °C. New high-resolution measurements of R across a range of measuring Ts (25-70 °C) yielded insights into the T at which maximal rates of R occurred (Tmax ). Although growth temperature did not affect Tmax , N+P fertilization increased Tmax values ~5 °C, from 53 to 58 °C. N+P fertilized shoots exhibited greater rates of R than nonfertilized shoots, with this effect diminishing under greenhouse warming. Collectively, our results highlight the nutrient dependence of thermal acclimation of leaf R in B. nana, suggesting that the metabolic efficiency allowed via thermal acclimation may be impaired at current levels of soil nutrient availability. This finding has important implications for predicting carbon fluxes in Arctic ecosystems, particularly if soil N and P become more abundant in the future as the tundra warms., (© 2014 John Wiley & Sons Ltd.)

We explored the impact of canopy position on leaf respiration (R) and associated traits in tree and shrub species growing in a lowland tropical rainforest in Far North Queensland, Australia. The range of traits quantified included: leaf R in darkness (RD) and in the light (RL; estimated using the Kok method); the temperature (T)-sensitivity of RD; light-saturated photosynthesis (Asat); leaf dry mass per unit area (LMA); and concentrations of leaf nitrogen (N), phosphorus (P), soluble sugars and starch. We found that LMA, and area-based N, P, sugars and starch concentrations were all higher in sun-exposed/upper canopy leaves, compared with their shaded/lower canopy and deep-shade/understory counterparts; similarly, area-based rates of RD, RL and Asat (at 28 °C) were all higher in the upper canopy leaves, indicating higher metabolic capacity in the upper canopy. The extent to which light inhibited R did not differ significantly between upper and lower canopy leaves, with the overall average inhibition being 32% across both canopy levels. Log-log RD-Asat relationships differed between upper and lower canopy leaves, with upper canopy leaves exhibiting higher rates of RD for a given Asat (both on an area and mass basis), as well as higher mass-based rates of RD for a given [N] and [P]. Over the 25-45 °C range, the T-sensitivity of RD was similar in upper and lower canopy leaves, with both canopy positions exhibiting Q10 values near 2.0 (i.e., doubling for every 10 °C rise in T) and Tmax values near 60 °C (i.e., T where RD reached maximal values). Thus, while rates of RD at 28 °C decreased with increasing depth in the canopy, the T-dependence of RD remained constant; these findings have important implications for vegetation-climate models that seek to predict carbon fluxes between tropical lowland rainforests and the atmosphere., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Depth has been widely recognised as a crucial operational feature of a high rate algal pond (HRAP) as it modifies the amount of light and frequency at which microalgal cells are exposed to optimal light. To date, there has been little focus on the optimisation of microalgal performance in wastewater treatment HRAPs with respect to depth, with advice ranging from as shallow as possible to 100 cm deep. This paper investigates the seasonal performance of microalgae in wastewater treatment HRAPs operated at three different depths (200, 300 and 400 mm). Microalgal performance was measured in terms of biomass production and areal productivity, nutrient removal efficiency and photosynthetic performance. The overall areal productivity significantly increased with increasing depth. Areal productivity ranged from 134 to 200% higher in the 400 mm deep HRAP compared to the 200 mm deep HRAP. Microalgae in the 400 mm deep HRAP were more efficient at NH4-N uptake and were photosynthetically more efficient compared to microalgae in the 200 mm deep HRAP. A higher chlorophyll-a concentration in the 200 mm deep HRAP resulted in a decrease in photosynthetic performance, due to insufficient carbon supply, over the course of the day in summer (as indicated by lower α, Pmax and oxygen production) compared to the 300 and 400 mm deep HRAPs. Based on these results, improved areal productivity and more wastewater can be treated per land area in the 400 mm deep HRAPs compared to 200 mm deep HRAPs without compromising wastewater treatment quality, while lowering capital and operational costs., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Seedlings of the shrub kānuka (Kunzea ericoides var. ericoides (A. Rich) J. Thompson) and the pasture grass brown top (Agrostis capillarus L.) were grown in intact soil cores in climate-controlled cabinets to analyse the thermal response of leaf-level carbon exchange at four levels of volumetric soil water content (θ). The objective was to resolve the combined effects of relatively rapid and short-term changes in θ and temperature on the thermal responses of both photosynthesis and respiration in these two contrasting plant types. Results showed that θ had a greater effect on the short-term temperature response of photosynthesis than the temperature response of respiration. The optimum value of θ for net photosynthesis was around 30% for both plants. The photosynthetic capacity of kānuka and the grass declined significantly when θ fell below 20%. The temperature sensitivity of photosynthesis was low at low soil water content and increased at moderate to high soil water content in both plant types. Statistical analysis showed that the temperature sensitivity of photosynthetic parameters was similar for both plant types, but the sensitivity of respiratory parameters differed. Respiratory capacity increased with increasing soil water content in kānuka but declined significantly when θ fell below 15%. There was no significant influence of soil water content on respiratory capacity in the grass. Collectively, our results indicate that θ influenced the temperature sensitivity of photosynthesis and respiration, and altered the balance between foliar respiration and photosynthetic capacity in both plant types.

Direct measurements of foliar carbon exchange through the growing season in Arctic species are limited, despite the need for accurate estimates of photosynthesis and respiration to characterise carbon cycling in the tundra. We examined seasonal variation in foliar photosynthesis and respiration (measured at 20°C) in two field-grown tundra species, Betula nana L. and Eriophorum vaginatum L., under ambient and long-term warming (LTW) conditions (+5°C), and the relationship of these fluxes to intraseasonal temperature variability. Species and seasonal timing drove most of the variation in photosynthetic parameters (e.g. gross photosynthesis (Agross)), respiration in the dark (Rdark) and light (Rlight), and foliar nitrogen concentration. LTW did not consistently influence fluxes through the season but reduced respiration in both species. Alongside the flatter respiratory response to measurement temperature in LTW leaves, this provided evidence of thermal acclimation. The inhibition of respiration by light increased by ~40%, with Rlight : Rdark values of ~0.8 at leaf out decreasing to ~0.4 after 8 weeks. Though LTW had no effect on inhibition, the cross-taxa seasonal decline in Rlight : Rdark greatly reduced respiratory carbon loss. Values of Rlight : Agross decreased from ~0.3 in both species to ~0.15 (B. nana) and ~0.05 (E. vaginatum), driven by decreases in respiratory rates, as photosynthetic rates remained stable. The influence of short-term temperature variability did not exhibit predictive trends for leaf gas exchange at a common temperature. These results underscore the influence of temperature on foliar carbon cycling, and the importance of respiration in controlling seasonal carbon exchange.

High nocturnal transpiration rates (5-15% of total water loss in terrestrial plants) may be adaptive under limited fertility, by increasing nutrient uptake or transport via transpiration-induced mass flow, but the response of stomata in the dark to environmental variables is poorly understood. Here we tested the impact of soil phosphorous (P) concentration, atmospheric CO2 concentration and air temperature on stomatal conductance (gs) during early and late periods in the night, as well as at midday in naturally, sun-lit glasshouse-grown Eucalyptus tereticornis Sm. seedlings. Soil P was the main driver of nocturnal gs, which was consistently higher in low soil P (37.3-79.9 mmol m(-2) s(-1)) than in high soil P (17.7-49.3 mmol m(-2)(-1)). Elevated temperature had only a marginal (P = 0.07) effect on gs early in the night (gs decreased from 34.7 to 25.8 mmol m(-2) s(-1) with an increase in temperature of 4 °C). The effect of CO2 depended on its interaction with temperature. Stomatal conductance responses to soil P were apparently driven by indirect effects of soil P on plant anatomy, since gs was significantly and negatively correlated with wood density. However, the relationship of gs with environmental factors became weaker late in the night, relative to early in the night, likely due to apparent endogenous processes; gs late in the night was two times larger than gs observed early in the night. Time-dependent controls over nocturnal gs suggest that daytime stomatal models may not apply during the night, and that different types of regulation may occur even within a single night. We conclude that the enhancement of nocturnal gs under low soil P availability is unlikely to be adaptive in our species because of the relatively small amount of transpiration-induced mass flow that can be achieved through rates of nocturnal water loss (3-6% of daytime mass flow).

Background: Non-structural carbohydrate (NSC; glucose, fructose, sucrose and fructan) composition of onions (Allium cepa L.) varies widely and is a key determinant of market usage. To analyse the physiology and genetics of onion carbohydrate metabolism and to enable selective breeding, an inexpensive, reliable and practicable sugar assay is required to phenotype large numbers of samples., Results: A rapid, reliable and cost-effective microplate-based assay was developed for NSC analysis in onions and used to characterise variation in tissue hexose, sucrose and fructan content in open-pollinated breeding populations and in mapping populations developed from a wide onion cross. Sucrose measured in microplates employing maltase as a hydrolytic enzyme was in agreement with HPLC-PAD results. The method revealed significant variation in bulb fructan content within open-pollinated 'Pukekohe Longkeeper' breeding populations over a threefold range. Very wide segregation from 80 to 600 g kg(-1) in fructan content was observed in bulbs of F2 genetic mapping populations from the wide onion cross 'Nasik Red × CUDH2150'., Conclusion: The microplate enzymatic assay is a reliable and practicable method for onion sugar analysis for genetics, breeding and food technology. Open-pollinated onion populations may harbour extensive within-population variability in carbohydrate content, which may be quantified and exploited using this method. The phenotypic data obtained from genetic mapping populations show that the method is well suited to detailed genetic and physiological analysis., (© 2013 Society of Chemical Industry.)

Plant respiratory metabolism is complicated by the fact that the rate of non-photorespiratory mitochondrial CO2 release in the light (R light) may be lower than the rate of leaf respiration in the dark (R dark). A body of work on this topic implies a linkage between light inhibition of respiration and photorespiration, although the direction of effect and underlying mechanisms remain uncertain. In this study we used a variety of short- and long-term environmental manipulations to explicitly manipulate the rate of photorespiration (νo) and quantify the effect on the inhibition of mitochondrial respiration in the light (R light:R dark). We address the following three questions: (i) will the R light:R dark ratio increase or decrease with high CO2 or low O2 and at low temperatures; (ii) does νo correlate with R light:R dark, and if so, in what way; (iii) will suppression of respiration by light (the 'Kok effect') be seen to the same extent in Zea mays, a C4 plant, and in Triticum aestivum, a C3 plant? We found that Rlight :Rdark decreased under conditions that suppressed νo in wheat, and this resulted in a positive relationship between R light:R dark and νo. Inhibition of respiration by light in C4 maize did not respond to environmental treatment, and the fixed R light:R dark (0.46-0.72) was consistent with the wheat response, assuming a νo approaching zero. The most likely mechanism to explain this finding is that R light increases (or the inhibition of respiration by light decreases) when there is an increase in photorespiration and thus an increase in the demand for TCA cycle substrates associated with the recovery of photorespiratory cycle intermediates in the peroxisome. This work is significant because it combines a comparison of C3 and C4 metabolism with a range of environmental treatments to independently suppress νo., (© 2013 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Laboratory studies indicate that plant respiratory efficiency may decrease in response to low nutrient availability due to increased partitioning of electrons to the energy-wasteful alternative oxidase (AOX); however, field confirmation of this hypothesis is lacking. We therefore investigated plant respiratory changes associated with succession and retrogression in soils aged from 10 to 120 000 years along the Franz Josef soil chronosequence, New Zealand. Respiration rates and electron partitioning were determined based on oxygen isotopic fractionation. Leaf structural traits, foliar nutrient status, carbohydrates and species composition were measured as explanatory variables. Although soil nutrient levels and species composition varied by site along the chronosequence, foliar respiration across all sites and species corresponded strongly with leaf nitrogen concentration (r(2)  = 0.8). In contrast, electron partitioning declined with increasing nitrogen/phosphorus (r(2)  = 0.23) and AOX activity correlated with phosphorus (r(2)  = 0.64). Independently, total respiration was further associated with foliar Cu, possibly linked to its effect on AOX. Independent control of AOX and cytochrome pathway activities is also discussed. These responses of plant terminal respiratory oxidases - and therefore respiratory carbon efficiency - to multiple nutrient deficiencies demonstrate that modulation of respiratory metabolism may play an important role in plant responses to nutrient gradients., (© 2012 Blackwell Publishing Ltd.)

Direct and indirect effects of warming are increasingly modifying the carbon-rich vegetation and soils of the Arctic tundra, with important implications for the terrestrial carbon cycle. Understanding the biological and environmental influences on the processes that regulate foliar carbon cycling in tundra species is essential for predicting the future terrestrial carbon balance in this region. To determine the effect of climate change impacts on gas exchange in tundra, we quantified foliar photosynthesis (A net), respiration in the dark and light (R D and R L, determined using the Kok method), photorespiration (PR), carbon gain efficiency (CGE, the ratio of photosynthetic CO2 uptake to total CO2 exchange of photosynthesis, PR, and respiration), and leaf traits of three dominant species - Betula nana, a woody shrub; Eriophorum vaginatum, a graminoid; and Rubus chamaemorus, a forb - grown under long-term warming and fertilization treatments since 1989 at Toolik Lake, Alaska. Under warming, B. nana exhibited the highest rates of A net and strongest light inhibition of respiration, increasing CGE nearly 50% compared with leaves grown in ambient conditions, which corresponded to a 52% increase in relative abundance. Gas exchange did not shift under fertilization in B. nana despite increases in leaf N and P and near-complete dominance at the community scale, suggesting a morphological rather than physiological response. Rubus chamaemorus, exhibited minimal shifts in foliar gas exchange, and responded similarly to B. nana under treatment conditions. By contrast, E. vaginatum, did not significantly alter its gas exchange physiology under treatments and exhibited dramatic decreases in relative cover (warming: -19.7%; fertilization: -79.7%; warming with fertilization: -91.1%). Our findings suggest a foliar physiological advantage in the woody shrub B. nana that is further mediated by warming and increased soil nutrient availability, which may facilitate shrub expansion and in turn alter the terrestrial carbon cycle in future tundra environments.