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Abstract Theory predicts that rising CO₂increases global photosynthesis, a process known as CO₂fertilization, and that this is responsible for much of the current terrestrial carbon sink. The estimated magnitude of the historic CO₂fertilization, however, differs by an order of magnitude between long-term proxies, remote sensing-based estimates and terrestrial biosphere models. Here we constrain the likely historic effect of CO₂on global photosynthesis by combining terrestrial biosphere models, ecological optimality theory, remote sensing approaches and an emergent constraint based on global carbon budget estimates. Our analysis suggests that CO₂fertilization increased global annual terrestrial photosynthesis by 13.5 ± 3.5% or 15.9 ± 2.9 PgC (mean ± s.d.) between 1981 and 2020. Our results help resolve conflicting estimates of the historic sensitivity of global terrestrial photosynthesis to CO₂and highlight the large impact anthropogenic emissions have had on ecosystems worldwide. 

Abstract Plants with the C₄photosynthesis pathway typically respond to climate change differently from more common C₃-type plants, due to their distinct anatomical and biochemical characteristics. These different responses are expected to drive changes in global C₄and C₃vegetation distributions. However, current C₄vegetation distribution models may not predict this response as they do not capture multiple interacting factors and often lack observational constraints. Here, we used global observations of plant photosynthetic pathways, satellite remote sensing, and photosynthetic optimality theory to produce an observation-constrained global map of C₄vegetation. We find that global C₄vegetation coverage decreased from 17.7% to 17.1% of the land surface during 2001 to 2019. This was the net result of a reduction in C₄natural grass cover due to elevated CO₂favoring C₃-type photosynthesis, and an increase in C₄crop cover, mainly from corn (maize) expansion. Using an emergent constraint approach, we estimated that C₄vegetation contributed 19.5% of global photosynthetic carbon assimilation, a value within the range of previous estimates (18–23%) but higher than the ensemble mean of dynamic global vegetation models (14 ± 13%; mean ± one standard deviation). Our study sheds insight on the critical and underappreciated role of C₄plants in the contemporary global carbon cycle. 

Abstract The connection between soil nitrogen availability, leaf nitrogen, and photosynthetic capacity is not perfectly understood. Because these three components tend to be positively related over large spatial scales, some posit that soil nitrogen positively drives leaf nitrogen, which positively drives photosynthetic capacity. Alternatively, others posit that photosynthetic capacity is primarily driven by above-ground conditions. Here, we examined the physiological responses of a non-nitrogen-fixing plant (Gossypium hirsutum) and a nitrogen-fixing plant (Glycine max) in a fully factorial combination of light by soil nitrogen availability to help reconcile these competing hypotheses. Soil nitrogen stimulated leaf nitrogen in both species, but the relative proportion of leaf nitrogen used for photosynthetic processes was reduced under elevated soil nitrogen in all light availability treatments due to greater increases in leaf nitrogen content than chlorophyll and leaf biochemical process rates. Leaf nitrogen content and biochemical process rates in G. hirsutum were more responsive to changes in soil nitrogen than those in G. max, probably due to strong G. max investments in root nodulation under low soil nitrogen. Nonetheless, whole-plant growth was significantly enhanced by increased soil nitrogen in both species. Light availability consistently increased relative leaf nitrogen allocation to leaf photosynthesis and whole-plant growth, a pattern that was similar between species. These results suggest that the leaf nitrogen–photosynthesis relationship varies under different soil nitrogen levels and that these species preferentially allocated more nitrogen to plant growth and non-photosynthetic leaf processes, rather than photosynthesis, as soil nitrogen increased. 

Summary Leaf dark respiration (R_d) acclimates to environmental changes. However, the magnitude, controls and time scales of acclimation remain unclear and are inconsistently treated in ecosystem models.We hypothesized thatR_dand Rubisco carboxylation capacity (V_cmax) at 25°C (R_d,25,V_cmax,25) are coordinated so thatR_d,25variations supportV_cmax,25at a level allowing full light use, withV_cmax,25reflecting daytime conditions (for photosynthesis), andR_d,25/V_cmax,25reflecting 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:R_d,25declines linearly with daily average prior temperature;R_dat average prior night temperatures tends towards a constant value; andR_d,25/V_cmax,25is constant.Our hypothesis accounted for more variation in observedR_d,25over time (R² = 0.74) and space (R² = 0.68) than the alternatives. Night‐time temperature dominated the seasonal time‐course ofR_d, with an apparent response time scale ofc.2 wk.V_cmaxdominated the spatial patterns.Our acclimation hypothesis results in a smaller increase in globalR_din response to rising CO₂and warming than is projected by the two of three alternative hypotheses, and by current models. 

Abstract Dominance often indicates one or a few species being best suited for resource capture and retention in a given environment. Press perturbations that change availability of limiting resources can restructure competitive hierarchies, allowing new species to capture or retain resources and leaving once dominant species fated to decline. However, dominant species may maintain high abundances even when their new environments no longer favour them due to stochastic processes associated with their high abundance, impeding deterministic processes that would otherwise diminish them.Here, we quantify the persistence of dominance by tracking the rate of decline in dominant species at 90 globally distributed grassland sites under experimentally elevated soil nutrient supply and reduced vertebrate consumer pressure.We found that chronic experimental nutrient addition and vertebrate exclusion caused certain subsets of species to lose dominance more quickly than in control plots. In control plots, perennial species and species with high initial cover maintained dominance for longer than annual species and those with low initial cover respectively. In fertilized plots, species with high initial cover maintained dominance at similar rates to control plots, while those with lower initial cover lost dominance even faster than similar species in controls. High initial cover increased the estimated time to dominance loss more strongly in plots with vertebrate exclosures than in controls. Vertebrate exclosures caused a slight decrease in the persistence of dominance for perennials, while fertilization brought perennials' rate of dominance loss in line with those of annuals. Annual species lost dominance at similar rates regardless of treatments.Synthesis.Collectively, these results point to a strong role of a species' historical abundance in maintaining dominance following environmental perturbations. Because dominant species play an outsized role in driving ecosystem processes, their ability to remain dominant—regardless of environmental conditions—is critical to anticipating expected rates of change in the structure and function of grasslands. Species that maintain dominance while no longer competitively favoured following press perturbations due to their historical abundances may result in community compositions that do not maximize resource capture, a key process of system responses to global change. 

Summary Nitrogen (N) limitation has been considered as a constraint on terrestrial carbon uptake in response to rising CO₂and climate change. By extension, it has been suggested that declining carboxylation capacity (V_cmax) and leaf N content in enhanced‐CO₂experiments and satellite records signify increasing N limitation of primary production. We predictedV_cmaxusing the coordination hypothesis and estimated changes in leaf‐level photosynthetic N for 1982–2016 assuming proportionality with leaf‐levelV_cmaxat 25°C. The whole‐canopy photosynthetic N was derived using satellite‐based leaf area index (LAI) data and an empirical extinction coefficient forV_cmax, and converted to annual N demand using estimated leaf turnover times. The predicted spatial pattern ofV_cmaxshares key features with an independent reconstruction from remotely sensed leaf chlorophyll content. Predicted leaf photosynthetic N declined by 0.27% yr⁻¹, while observed leaf (total) N declined by 0.2–0.25% yr⁻¹. Predicted global canopy N (and N demand) declined from 1996 onwards, despite increasing LAI. Leaf‐level responses to rising CO₂, and to a lesser extent temperature, may have reduced the canopy requirement for N by more than rising LAI has increased it. This finding provides an alternative explanation for declining leaf N that does not depend on increasing N limitation. 

Abstract Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions. 

Abstract Aim Understanding the considerable variability and drivers of global leaf photosynthetic capacity [indicated by the maximum carboxylation rate standardized to 25°C ( V c,max25 )] is an essential step for accurate modelling of terrestrial plant photosynthesis and carbon uptake under climate change. Although current environmental conditions have often been connected with empirical and theoretical models to explain global V c,max25 variability through acclimatization and adaptation, long‐term evolutionary history has largely been neglected, but might also explicitly play a role in shaping the V c,max25 variability. Location Global. Time period Contemporary. Major taxa studied Terrestrial plants. Methods We compiled a geographically comprehensive global dataset of V c,max25 for C 3 plants ( n  = 6917 observations from 2157 species and 425 sites covering all major biomes world‐wide), explored the biogeographical and phylogenetic patterns of V c,max25 , and quantified the relative importance of current environmental factors and evolutionary history in driving global V c,max25 variability. Results We found that V c,max25 differed across different biomes, with higher mean values in relatively drier regions, and across different life‐forms, with higher mean values in non‐woody relative to woody plants and in legumes relative to non‐leguminous plants. The values of V c,max25 displayed a significant phylogenetic signal and diverged in a contrasting manner across phylogenetic groups, with a significant trend along the evolutionary axis towards a higher V c,max25 in more modern clades. A Bayesian phylogenetic linear mixed model revealed that evolutionary history (indicated by phylogeny and species) explained nearly 3‐fold more of the variation in global V c,max25 than present‐day environment (53 vs. 18%). Main conclusions These findings contribute to a comprehensive assessment of the patterns and drivers of global V c,max25 variability, highlighting the importance of evolutionary history in driving global V c,max25 variability, hence terrestrial plant photosynthesis.