Variation in leaf dark respiration among C3 and C4 grasses is associated with use of different substrates.

Measurements of respiratory properties have often been made at a single time point either during daytime using dark-adapted leaves or during nighttime. The influence of the day-night cycle on respiratory metabolism has received less attention but is crucial to understand photosynthesis and photorespiration. Here, we examined how CO2- and O2-based rates of leaf dark respiration (Rdark) differed between midday (after 30-min dark adaptation) and midnight in 8 C3 and C4 grasses. We used these data to calculate the respiratory quotient (RQ; ratio of CO2 release to O2 uptake), and assessed relationships between Rdark and leaf metabolome. Rdark was higher at midday than midnight, especially in C4 species. The day-night difference in Rdark was more evident when expressed on a CO2 than O2 basis, with the RQ being higher at midday than midnight in all species, except in rice (Oryza sativa). Metabolomic analyses showed little correlation of Rdark or RQ with leaf carbohydrates (sucrose, glucose, fructose, or starch) but strong multivariate relationships with other metabolites. The results suggest that rates of Rdark and differences in RQ were determined by several concurrent CO2-producing and O2-consuming metabolic pathways, not only the tricarboxylic acid cycle (organic acids utilization) but also the pentose phosphate pathway, galactose metabolism, and secondary metabolism. As such, Rdark was time-, type- (C3/C4) and species-dependent, due to the use of different substrates.

Heat tolerance of a tropical-subtropical rainforest tree species Polyscias elegans: time-dependent dynamic responses of physiological thermostability and biochemistry.

Heat stress interrupts physiological thermostability and triggers biochemical responses that are essential for plant survival. However, there is limited knowledge on the speed plants adjust to heat in hours and days, and which adjustments are crucial. Tropical-subtropical rainforest tree species (Polyscias elegans) were heated at 40°C for 5 d, before returning to 25°C for 13 d of recovery. Leaf heat tolerance was quantified using the temperature at which minimal chl a fluorescence sharply rose (Tcrit ). Tcrit , metabolites, heat shock protein (HSP) abundance and membrane lipid fatty acid (FA) composition were quantified. Tcrit increased by 4°C (48-52°C) within 2 h of 40°C exposure, along with rapid accumulation of metabolites and HSPs. By contrast, it took > 2 d for FA composition to change. At least 2 d were required for Tcrit , HSP90, HSP70 and FAs to return to prestress levels. The results highlight the multi-faceted response of P. elegans to heat stress, and how this response varies over the scale of hours to days, culminating in an increased level of photosynthetic heat tolerance. These responses are important for survival of plants when confronted with heat waves amidst ongoing global climate change.

Reduced global plant respiration due to the acclimation of leaf dark respiration coupled with photosynthesis.

Leaf dark respiration (Rd ) acclimates to environmental changes. However, the magnitude, controls and time scales of acclimation remain unclear and are inconsistently treated in ecosystem models. We hypothesized that Rd and Rubisco carboxylation capacity (Vcmax ) at 25°C (Rd,25 , Vcmax,25 ) are coordinated so that Rd,25 variations support Vcmax,25 at a level allowing full light use, with Vcmax,25 reflecting daytime conditions (for photosynthesis), and Rd,25 /Vcmax,25 reflecting night-time conditions (for starch degradation and sucrose export). We tested this hypothesis temporally using a 5-yr warming experiment, and spatially using an extensive field-measurement data set. We compared the results to three published alternatives: Rd,25 declines linearly with daily average prior temperature; Rd at average prior night temperatures tends towards a constant value; and Rd,25 /Vcmax,25 is constant. Our hypothesis accounted for more variation in observed Rd,25 over time (R2 = 0.74) and space (R2 = 0.68) than the alternatives. Night-time temperature dominated the seasonal time-course of Rd , with an apparent response time scale of c. 2 wk. Vcmax dominated the spatial patterns. Our acclimation hypothesis results in a smaller increase in global Rd in response to rising CO2 and warming than is projected by the two of three alternative hypotheses, and by current models.

Photosynthesis in newly developed leaves of heat-tolerant wheat acclimates to long-term nocturnal warming.

We examined photosynthetic traits of pre-existing and newly developed flag leaves of four wheat genotypes grown in controlled-environment experiments. In newly developed leaves, acclimation of the maximum rate of net CO2 assimilation (An) to warm nights (i.e. increased An) was associated with increased capacity of Rubisco carboxylation and photosynthetic electron transport, with Rubisco activation state probably contributing to increased Rubisco activity. Metabolite profiling linked acclimation of An to greater accumulation of monosaccharides and saturated fatty acids in leaves; these changes suggest roles for osmotic adjustment of leaf turgor pressure and maintenance of cell membrane integrity. By contrast, where An decreased under warm nights, the decline was related to lower stomatal conductance and rates of photosynthetic electron transport. Decreases in An occurred despite higher basal PSII thermal stability in all genotypes exposed to warm nights: Tcrit of 45-46.5 °C in non-acclimated versus 43.8-45 °C in acclimated leaves. Pre-existing leaves showed no change in An-temperature response curves, except for an elite heat-tolerant genotype. These findings illustrate the impact of night-time warming on the ability of wheat plants to photosynthesize during the day, thereby contributing to explain the impact of global warming on crop productivity.

Reduced tillering and dwarfing genes alter root traits and rhizo-economics in wheat.

The tiller inhibition (tin) and Reduced height (Rht) genes strongly influence the carbon partitioning and architecture of wheat shoots, but their effects on the energy economy of roots have not been examined in detail. We examined multiple root traits in three sets of near-isogenic wheat lines (NILs) that differ in the tin gene or various dwarfing gene alleles (Rht-B1b, Rht-D1b, Rht-B1c and Rht-B1b + Rht-D1b) to determine their effects on root structure, anatomy and carbon allocation. The tin gene resulted in fewer tillers but more costly roots in an extreme tin phenotype with a Banks genetic background due to increases in root-to-shoot ratio, total root length, and whole root respiration. However, this effect depended on the genetic background as tin caused both smaller shoots and roots in a different genetic background. The semi-dwarf gene Rht-B1b caused few changes to the root structure, whereas Rht-D1b, Rht-B1c and the double dwarf (Rht-B1b + Rht-D1b) decreased the root biomass. Rht-B1c reduced the energy cost of roots by increasing specific root length, increasing the volume of cortical aerenchyma and by reducing root length, number, and biomass without affecting the root-to-shoot ratio. This work informs researchers using tin and Rht genes how to modify root system architecture to suit specific environments.

Wheat photosystem II heat tolerance: evidence for genotype-by-environment interactions.

High temperature stress inhibits photosynthesis and threatens wheat production. One measure of photosynthetic heat tolerance is Tcrit - the critical temperature at which incipient damage to photosystem II (PSII) occurs. This trait could be improved in wheat by exploiting genetic variation and genotype-by-environment interactions (GEI). Flag leaf Tcrit of 54 wheat genotypes was evaluated in 12 thermal environments over 3 years in Australia, and analysed using linear mixed models to assess GEI effects. Nine of the 12 environments had significant genetic effects and highly variable broad-sense heritability (H2 ranged from 0.15 to 0.75). Tcrit GEI was variable, with 55.6% of the genetic variance across environments accounted for by the factor analytic model. Mean daily growth temperature in the month preceding anthesis was the most influential environmental driver of Tcrit GEI, suggesting biochemical, physiological and structural adjustments to temperature requiring different durations to manifest. These changes help protect or repair PSII upon exposure to heat stress, and may improve carbon assimilation under high temperature. To support breeding efforts to improve wheat performance under high temperature, we identified genotypes superior to commercial cultivars commonly grown by farmers, and demonstrated potential for developing genotypes with greater photosynthetic heat tolerance.

Enhancing crop yields through improvements in the efficiency of photosynthesis and respiration.

The rate with which crop yields per hectare increase each year is plateauing at the same time that human population growth and other factors increase food demand. Increasing yield potential ( Y p ) of crops is vital to address these challenges. In this review, we explore a component of Y p that has yet to be optimised - that being improvements in the efficiency with which light energy is converted into biomass ( ε c ) via modifications to CO2 fixed per unit quantum of light (α), efficiency of respiratory ATP production ( ε prod ) and efficiency of ATP use ( ε use ). For α, targets include changes in photoprotective machinery, ribulose bisphosphate carboxylase/oxygenase kinetics and photorespiratory pathways. There is also potential for ε prod to be increased via targeted changes to the expression of the alternative oxidase and mitochondrial uncoupling pathways. Similarly, there are possibilities to improve ε use via changes to the ATP costs of phloem loading, nutrient uptake, futile cycles and/or protein/membrane turnover. Recently developed high-throughput measurements of respiration can serve as a proxy for the cumulative energy cost of these processes. There are thus exciting opportunities to use our growing knowledge of factors influencing the efficiency of photosynthesis and respiration to create a step-change in yield potential of globally important crops.

Coordination of photosynthetic traits across soil and climate gradients.

"Least-cost theory" posits that C3 plants should balance rates of photosynthetic water loss and carboxylation in relation to the relative acquisition and maintenance costs of resources required for these activities. Here we investigated the dependency of photosynthetic traits on climate and soil properties using a new Australia-wide trait dataset spanning 528 species from 67 sites. We tested the hypotheses that plants on relatively cold or dry sites, or on relatively more fertile sites, would typically operate at greater CO2 drawdown (lower ratio of leaf internal to ambient CO2 , Ci :Ca ) during light-saturated photosynthesis, and at higher leaf N per area (Narea ) and higher carboxylation capacity (Vcmax 25 ) for a given rate of stomatal conductance to water vapour, gsw . These results would be indicative of plants having relatively higher water costs than nutrient costs. In general, our hypotheses were supported. Soil total phosphorus (P) concentration and (more weakly) soil pH exerted positive effects on the Narea -gsw and Vcmax 25 -gsw slopes, and negative effects on Ci :Ca . The P effect strengthened when the effect of climate was removed via partial regression. We observed similar trends with increasing soil cation exchange capacity and clay content, which affect soil nutrient availability, and found that soil properties explained similar amounts of variation in the focal traits as climate did. Although climate typically explained more trait variation than soil did, together they explained up to 52% of variation in the slope relationships and soil properties explained up to 30% of the variation in individual traits. Soils influenced photosynthetic traits as well as their coordination. In particular, the influence of soil P likely reflects the Australia's geologically ancient low-relief landscapes with highly leached soils. Least-cost theory provides a valuable framework for understanding trade-offs between resource costs and use in plants, including limiting soil nutrients.

The crucial roles of mitochondria in supporting C4 photosynthesis.

C4 photosynthesis involves a series of biochemical and anatomical traits that significantly improve plant productivity under conditions that reduce the efficiency of C3 photosynthesis. We explore how evolution of the three classical biochemical types of C4 photosynthesis (NADP-ME, NAD-ME and PCK types) has affected the functions and properties of mitochondria. Mitochondria in C4 NAD-ME and PCK types play a direct role in decarboxylation of metabolites for C4 photosynthesis. Mitochondria in C4 PCK type also provide ATP for C4 metabolism, although this role for ATP provision is not seen in NAD-ME type. Such involvement has increased mitochondrial abundance/size and associated enzymatic capacity, led to changes in mitochondrial location and ultrastructure, and altered the role of mitochondria in cellular carbon metabolism in the NAD-ME and PCK types. By contrast, these changes in mitochondrial properties are absent in the C4 NADP-ME type and C3 leaves, where mitochondria play no direct role in photosynthesis. From an eco-physiological perspective, rates of leaf respiration in darkness vary considerably among C4 species but does not differ systematically among the three C4 types. This review outlines further mitochondrial research in key areas central to the engineering of the C4 pathway into C3 plants and to the understanding of variation in rates of C4 dark respiration.

Dark respiration rates are not determined by differences in mitochondrial capacity, abundance and ultrastructure in C4 leaves.

Our understanding of the regulation of respiration in C4 plants, where mitochondria play different roles in the different types of C4 photosynthetic pathway, remains limited. We examined how leaf dark respiration rates (Rdark ), in the presence and absence of added malate, vary in monocots representing the three classical biochemical types of C4 photosynthesis (NADP-ME, NAD-ME and PCK) using intact leaves and extracted bundle sheath strands. In particular, we explored to what extent rates of Rdark are associated with mitochondrial number, volume and ultrastructure. Based on examination of a single species per C4 type, we found that the respiratory response of NAD-ME and PCK type bundle sheath strands to added malate was associated with differences in mitochondrial number, volume, and/or ultrastructure, while NADP-ME type bundle sheath strands did not respond to malate addition. In general, mitochondrial traits reflected the contributions mitochondria make to photosynthesis in the three C4 types. However, despite the obvious differences in mitochondrial traits, no clear correlation was observed between these traits and Rdark . We suggest that Rdark is primarily driven by cellular maintenance demands and not mitochondrial composition per se, in a manner that is somewhat independent of mitochondrial organic acid cycling in the light.

Wheat photosystem II heat tolerance responds dynamically to short- and long-term warming.

Wheat photosynthetic heat tolerance can be characterized using minimal chlorophyll fluorescence to quantify the critical temperature (Tcrit) above which incipient damage to the photosynthetic machinery occurs. We investigated intraspecies variation and plasticity of wheat Tcrit under elevated temperature in field and controlled-environment experiments, and assessed whether intraspecies variation mirrored interspecific patterns of global heat tolerance. In the field, wheat Tcrit varied diurnally-declining from noon through to sunrise-and increased with phenological development. Under controlled conditions, heat stress (36 °C) drove a rapid (within 2 h) rise in Tcrit that peaked after 3-4 d. The peak in Tcrit indicated an upper limit to PSII heat tolerance. A global dataset [comprising 183 Triticum and wild wheat (Aegilops) species] generated from the current study and a systematic literature review showed that wheat leaf Tcrit varied by up to 20 °C (roughly two-thirds of reported global plant interspecies variation). However, unlike global patterns of interspecies Tcrit variation that have been linked to latitude of genotype origin, intraspecific variation in wheat Tcrit was unrelated to that. Overall, the observed genotypic variation and plasticity of wheat Tcrit suggest that this trait could be useful in high-throughput phenotyping of wheat photosynthetic heat tolerance.

Oxygen uptake rates have contrasting responses to temperature in the root meristem and elongation zone.

Growing at either 15 or 25°C, roots of Arabidopsis thaliana, Columbia accession, produce cells at the same rate and have growth zones of the same length. To determine whether this constancy is related to energetics, we measured oxygen uptake by means of a vibrating oxygen-selective electrode. Concomitantly, the spatial distribution of elongation was measured kinematically, delineating meristem and elongation zone. All seedlings were germinated, grown, and measured at a given temperature (15 or 25°C). Columbia was compared to lines where cell production rate roughly doubles between 15 and 25°C: Landsberg and two Columbia mutants, er-105 and ahk3-3. For all genotypes and temperatures, oxygen uptake rate at any position was highest at the root cap, where mitochondrial density was maximal, based on the fluorescence of a reporter. Uptake rate declined through the meristem to plateau within the elongation zone. For oxygen uptake rate integrated over a zone, the meristem had steady-state Q10 values ranging from 0.7 to 2.1; by contrast, the elongation zone had values ranging from 2.6 to 3.3, implying that this zone exerts a greater respiratory demand. These results highlight a substantial energy consumption by the root cap, perhaps helpful for maintaining hypoxia in stem cells, and suggest that rapid elongation is metabolically more costly than is cell division.

Acclimation of leaf respiration temperature responses across thermally contrasting biomes.

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 R25 (measured at 25°C) and the leaf T where R reached its maximum (Tmax ) 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. R25 decreased and Tmax 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 R25 and Tmax 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.

Responses of leaf respiration to heatwaves.

Mitochondrial respiration (R) is central to plant physiology and responds dynamically to daily short-term temperature changes. In the longer-term, changes in energy demand and membrane fluidity can decrease leaf R at a common temperature and increase the temperature at which leaf R peaks (Tmax ). However, leaf R functionality is more susceptible to short-term heatwaves. Catalysis increases with rising leaf temperature, driving faster metabolism and leaf R demand, despite declines in photosynthesis restricting assimilate supply and growth. Proteins denature as temperatures increase further, adding to maintenance costs. Excessive heat also inactivates respiratory enzymes, with a concomitant limitation on the capacity of the R system. These competing push-and-pull factors are responsible for the diminishing acceleration in leaf R rate as temperature rises. Under extreme heat, membranes become overly fluid, and enzymes such as the cytochrome c oxidase are impaired. Such changes can lead to over-reduction of the energy system culminating in reactive oxygen species production. This ultimately leads to the total breakdown of leaf R, setting the limit of leaf survival. Understanding the heat stress responses of leaf R is imperative, given the continued rise in frequency and intensity of heatwaves and the importance of R for plant fitness and survival.

Acclimation of leaf photosynthesis and respiration to warming in field-grown wheat.

Climate change and future warming will significantly affect crop yield. The capacity of crops to dynamically adjust physiological processes (i.e., acclimate) to warming might improve overall performance. Understanding and quantifying the degree of acclimation in field crops could ensure better parameterization of crop and Earth System models and predictions of crop performance. We hypothesized that for field-grown wheat, when measured at a common temperature (25°C), crops grown under warmer conditions would exhibit acclimation, leading to enhanced crop performance and yield. Acclimation was defined as (a) decreased rates of net photosynthesis at 25°C (A25 ) coupled with lower maximum carboxylation capacity (Vcmax25 ), (b) reduced leaf dark respiration at 25°C (both in terms of O2 consumption Rdark _O225 and CO2 efflux Rdark _CO225 ) and (c) lower Rdark _CO225 to Vcmax25 ratio. Field experiments were conducted over two seasons with 20 wheat genotypes, sown at three different planting dates, to test these hypotheses. Leaf-level CO2 -based traits (A25 , Rdark _CO225 and Vcmax25 ) did not show the classic acclimation responses that we hypothesized; by contrast, the hypothesized changes in Rdark_ O2 were observed. These findings have implications for predictive crop models that assume similar temperature response among these physiological processes and for predictions of crop performance in a future warmer world.

Diel- and temperature-driven variation of leaf dark respiration rates and metabolite levels in rice.

Leaf respiration in the dark (Rdark ) is often measured at a single time during the day, with hot-acclimation lowering Rdark at a common measuring temperature. However, it is unclear whether the diel cycle influences the extent of thermal acclimation of Rdark , or how temperature and time of day interact to influence respiratory metabolites. To examine these issues, we grew rice under 25°C : 20°C, 30°C : 25°C and 40°C : 35°C day : night cycles, measuring Rdark and changes in metabolites at five time points spanning a single 24-h period. Rdark differed among the treatments and with time of day. However, there was no significant interaction between time and growth temperature, indicating that the diel cycle does not alter thermal acclimation of Rdark . Amino acids were highly responsive to the diel cycle and growth temperature, and many were negatively correlated with carbohydrates and with organic acids of the tricarboxylic acid (TCA) cycle. Organic TCA intermediates were significantly altered by the diel cycle irrespective of growth temperature, which we attributed to light-dependent regulatory control of TCA enzyme activities. Collectively, our study shows that environmental disruption of the balance between respiratory substrate supply and demand is corrected for by shifts in TCA-dependent metabolites.

Leaf trait variation is similar among genotypes of Eucalyptus camaldulensis from differing climates and arises in plastic responses to the seasons rather than water availability.

We used a widely distributed tree Eucalyptus camaldulensis subsp. camaldulensis to partition intraspecific variation in leaf functional traits to genotypic variation and phenotypic plasticity. We examined if genotypic variation is related to the climate of genotype provenance and whether phenotypic plasticity maintains performance in a changing environment. Ten genotypes from different climates were grown in a common garden under watering treatments reproducing the wettest and driest edges of the subspecies' distribution. We measured functional traits reflecting leaf metabolism and associated with growth (respiration rate, nitrogen and phosphorus concentrations, and leaf mass per area) and performance proxies (aboveground biomass and growth rate) each season over a year. Genotypic variation contributed substantially to the variation in aboveground biomass but much less in growth rate and leaf traits. Phenotypic plasticity was a large source of the variation in leaf traits and performance proxies and was greater among sampling dates than between watering treatments. The variation in leaf traits was weakly correlated to performance proxies, and both were unrelated to the climate of genotype provenance. Intraspecific variation in leaf traits arises similarly among genotypes in response to seasonal environmental variation, instead of long-term water availability or climate of genotype provenance.

Molecular and physiological responses during thermal acclimation of leaf photosynthesis and respiration in rice.

To further our understanding of how sustained changes in temperature affect the carbon economy of rice (Oryza sativa), hydroponically grown plants of the IR64 cultivar were developed at 30°C/25°C (day/night) before being shifted to 25/20°C or 40/35°C. Leaf messenger RNA and protein abundance, sugar and starch concentrations, and gas-exchange and elongation rates were measured on preexisting leaves (PE) already developed at 30/25°C or leaves newly developed (ND) subsequent to temperature transfer. Following a shift in growth temperature, there was a transient adjustment in metabolic gene transcript abundance of PE leaves before homoeostasis was reached within 24 hr, aligning with Rdark (leaf dark respiratory CO2 release) and An (net CO2 assimilation) changes. With longer exposure, the central respiratory protein cytochrome c oxidase (COX) declined in abundance at 40/35°C. In contrast to Rdark , An was maintained across the three growth temperatures in ND leaves. Soluble sugars did not differ significantly with growth temperature, and growth was fastest with extended exposure at 40/35°C. The results highlight that acclimation of photosynthesis and respiration is asynchronous in rice, with heat-acclimated plants exhibiting a striking ability to maintain net carbon gain and growth when exposed to heat-wave temperatures, even while reducing investment in energy-conserving respiratory pathways.

Acclimation of leaf respiration consistent with optimal photosynthetic capacity.

Plant respiration is an important contributor to the proposed positive global carbon-cycle feedback to climate change. However, as a major component, leaf mitochondrial ('dark') respiration (Rd ) differs among species adapted to contrasting environments and is known to acclimate to sustained changes in temperature. No accepted theory explains these phenomena or predicts its magnitude. Here we propose that the acclimation of Rd follows an optimal behaviour related to the need to maintain long-term average photosynthetic capacity (Vcmax ) so that available environmental resources can be most efficiently used for photosynthesis. To test this hypothesis, we extend photosynthetic co-ordination theory to predict the acclimation of Rd to growth temperature via a link to Vcmax , and compare predictions to a global set of measurements from 112 sites spanning all terrestrial biomes. This extended co-ordination theory predicts that field-measured Rd and Vcmax accessed at growth temperature (Rd,tg and Vcmax,tg ) should increase by 3.7% and 5.5% per degree increase in growth temperature. These acclimated responses to growth temperature are less steep than the corresponding instantaneous responses, which increase 8.1% and 9.9% per degree of measurement temperature for Rd and Vcmax respectively. Data-fitted responses proof indistinguishable from the values predicted by our theory, and smaller than the instantaneous responses. Theory and data are also shown to agree that the basal rates of both Rd and Vcmax assessed at 25°C (Rd,25 and Vcmax,25 ) decline by ~4.4% per degree increase in growth temperature. These results provide a parsimonious general theory for Rd acclimation to temperature that is simpler-and potentially more reliable-than the plant functional type-based leaf respiration schemes currently employed in most ecosystem and land-surface models.

TRY plant trait database - enhanced coverage and open access.

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives.