Ecosystem assembly and terrestrial carbon balance under elevated CO(2)

Research aimed at understanding how the global carbon balance will change with elevated CO(2) has largely ignored the responses of individual species and genotypes. Yet, plant traits strongly influence the biogeochemical cycling of carbon. Here, we illustrate how differences in inter- and intraspecific responses to elevated CO(2) affect not only physiology and growth, but also higher order biotic interactions and lifetime fitness, ultimately leading to new ecosystem assemblages. We assert that the unique combination of inter- and intraspecific traits in these ecosystem assemblages ultimately determine how ecosystems respond to elevated atmospheric CO(2). Thus, the identity of species and genotypes in an ecosystem is a crucial element to consider in forecasts of global carbon balance.     

Cellular Growth Arrest and Persistence from Enzyme Saturation

Metabolic efficiency depends on the balance between supply and demand of metabolites, which is sensitive to environmental and physiological fluctuations, or noise, causing shortages or surpluses in the metabolic pipeline. How cells can reliably optimize biomass production in the presence of metabolic fluctuations is a fundamental question that has not been fully answered. Here we use mathematical models to predict that enzyme saturation creates distinct regimes of cellular growth, including a phase of growth arrest resulting from toxicity of the metabolic process. Noise can drive entry of single cells into growth arrest while a fast-growing majority sustains the population. We confirmed these predictions by measuring the growth dynamics of Escherichia coli utilizing lactose as a sole carbon source. The predicted heterogeneous growth emerged at high lactose concentrations, and was associated with cell death and production of antibiotic-tolerant persister cells. These results suggest how metabolic networks may balance costs and benefits, with important implications for drug tolerance.     

Differential Growth Responses to Water Balance of Coexisting Deciduous Tree Species Are Linked to Wood Density in a Bolivian Tropical Dry Forest

A seasonal period of water deficit characterizes tropical dry forests (TDFs). There, sympatric tree species exhibit a diversity of growth rates, functional traits, and responses to drought, suggesting that each species may possess different strategies to grow under different conditions of water availability. The evaluation of the long-term growth responses to changes in the soil water balance should provide an understanding of how and when coexisting tree species respond to water deficit in TDFs. Furthermore, such differential growth responses may be linked to functional traits related to water storage and conductance. We used dendrochronology and climate data to retrospectively assess how the radial growth of seven coexisting deciduous tree species responded to the seasonal soil water balance in a Bolivian TDF. Linear mixed-effects models were used to quantify the relationships between basal area increment and seasonal water balance. We related these relationships with wood density and sapwood production to assess if they affect the growth responses to climate. The growth of all species responded positively to water balance during the wet season, but such responses differed among species as a function of their wood density. For instance, species with a strong growth response to water availability averaged a low wood density which may facilitate the storage of water in the stem. By contrast, species with very dense wood were those whose growth was less sensitive to water availability. Coexisting tree species thus show differential growth responses to changes in soil water balance during the wet season. Our findings also provide a link between wood density, a trait related to the ability of trees to store water in the stem, and wood formation in response to water availability.     

Interactions between increasing CO2 concentration and temperature on plant growth

The global environment is changing with increasing temperature and atmospheric carbon dioxide concentration, [CO₂]. Because these two factors are concomitant, and the global [CO₂] rise will affect all biomes across the full global range of temperatures, it is essential to review the theory and observations on effects of temperature and [CO₂] interactions on plant carbon balance, growth, development, biomass accumulation and yield. Although there are sound theoretical reasons for expecting a larger stimulation of net CO₂ assimilation rates by increased [CO₂] at higher temperatures, this does not necessarily mean that the pattern of biomass and yield responses to increasing [CO₂] and temperature is determined by this response. This paper reviews the interactions between the effects of [CO₂] and temperature on plants. There is little unequivocal evidence for large differences in response to [CO₂] at different temperatures, as studies are confounded by the different responses of species adapted and acclimated to different temperatures, and the interspecific differences in growth form and development pattern. We conclude by stressing the importance of initiation and expansion of meristems and organs and the balance between assimilate supply and sink activity in determining the growth response to increasing [CO₂] and temperature.     

A Framework for Understanding Variation in Pelagic Gross Primary Production of Lake Ecosystems

Light and nutrient availability are key physiological constraints for primary production. Widespread environmental changes are causing variability in loads of terrestrial dissolved organic carbon (DOC) and nutrients from watersheds to lakes, contributing to simultaneous changes in both light and nutrient supply. Experimental evidence highlights the potential for these watershed loads to create complex and context-dependent responses of within-lake primary production; however, the field lacks a predictive model to investigate these responses. We embedded a well-established physiological model of phytoplankton growth within an ecosystem model of nutrient and DOC supply to assess how simultaneous changes in DOC and nutrient loads could impact pelagic primary production in lakes. The model generated a unimodal relationship between GPP and DOC concentration when loads of DOC and nutrients were tightly correlated across space or time. In this unimodal relationship, the magnitude of the peak GPP was primarily determined by the DOC-to-nutrient ratio of the load, and the location of the peak along the DOC axis was primarily determined by lake area. Greater nutrient supply relative to DOC load contributed to greater productivity, and larger lake area increased light limitation for primary producers at a given DOC concentration, owing to the positive relationship between lake area and epilimnion depth. When loads of DOC and nutrients were not tightly correlated in space or time, the model generated a wedge-shaped pattern between GPP and DOC, consistent with spatial surveys from a global set of lakes. Our model is thus capable of unifying the diversity of empirically observed spatial and temporal responses of lake productivity to DOC and mineral nutrient supply presented in the literature, and provides qualitative predictions for how lake pelagic primary productivity may respond to widespread environmental changes.     

Control of carbon flux to acetate excretion during growth of Escherichia coli in batch and continuous cultures

During growth of Escherichia coli ML308 on pyruvate in a continuous culture (turbidostat) or batch culture, flux of carbon into the cells exceeds the amphibolic capacity of the central pathways. This is balanced by diversion of carbon flux to acetate excretion which in turn diminishes the efficiency of carbon conversion to biomass [g] dry wt (mol substrate)-1]. However, restriction of carbon supply in a chemostat diminishes flux to acetate excretion and at a dilution rate (D = mu) of 0.35 h-1 or less, no flux to acetate excretion was sustained thus permitting perfect balance between carbon input on the one hand, and the output to biosynthesis and energy generation on the other. This, in turn, improves the efficiency of carbon conversion to biomass. Inclusion of 3-bromopyruvate (an inhibitor of pyruvate dehydrogenase) at a concentration which diminishes growth rate (mu) to 0.35 h-1 or less also prevented flux to acetate excretion. Furthermore, in a family of fluoroacetate-resistant strains, excessive flux of pyruvate was balanced by diversion of carbon flux to lactate excretion rather than acetate and a higher growth rate (mu = 0.63 h-1) was sustained.     

Nutrient demand and fungal access to resources control the carbon allocation to the symbiotic partners in tripartite interactions of Medicago truncatula

Legumes form tripartite interactions with arbuscular mycorrhizal fungi and rhizobia, and both root symbionts exchange nutrients against carbon from their host. The carbon costs of these interactions are substantial, but our current understanding of how the host controls its carbon allocation to individual root symbionts is limited. We examined nutrient uptake and carbon allocation in tripartite interactions of Medicago truncatula under different nutrient supply conditions, and when the fungal partner had access to nitrogen, and followed the gene expression of several plant transporters of the Sucrose Uptake Transporter (SUT) and Sugars Will Eventually be Exported Transporter (SWEET) family. Tripartite interactions led to synergistic growth responses and stimulated the phosphate and nitrogen uptake of the plant. Plant nutrient demand but also fungal access to nutrients played an important role for the carbon transport to different root symbionts, and the plant allocated more carbon to rhizobia under nitrogen demand, but more carbon to the fungal partner when nitrogen was available. These changes in carbon allocation were consistent with changes in the SUT and SWEET expression. Our study provides important insights into how the host plant controls its carbon allocation under different nutrient supply conditions and changes its carbon allocation to different root symbionts to maximize its symbiotic benefits.     

大连金普新区碳固持生态服务供需格局时空差异

碳固持服务供需匹配状况对于优化区域生态系统管理、提升生态系统碳汇能力具有重要意义。以大连金普新区为研究区,利用InVEST模型、ArcGIS模型,研究固碳项生态系统服务供需匹配及其变化特征。结果表明：(1)研究期内,2010年至2018年,碳固持服务供给总量呈现出U字型变化趋势;需求总量呈现出持续增长的变化趋势,表现为低等级需求量地区向高等级需求量地区不断转化;碳固持服务供需求不匹配,供需错配现象突出,总体上呈现出供不应求。(2)碳固持服务供需赤字缺口正持续扩大。赤字类型总面积有持续增加的趋势;盈余类型总面积表现为先增后减;供需相对平衡类型总面积不断缩减,朝着低供给高需求方向转变。(3)金普新区碳汇地面积持续缩减,2018年仅为564km²;碳源地面积大量增加,2018年达到1340km²。(4)提出了控制城镇碳需求增长,增加绿地,提高碳供给能力,消减赤字,维持供需能力基本平衡的对策。     

A Race for Space? How Sphagnum fuscum stabilizes vegetation composition during long‐term climate manipulations

Strong climate warming is predicted at higher latitudes this century, with potentially major consequences for productivity and carbon sequestration. Although northern peatlands contain one‐third of the world's soil organic carbon, little is known about the long‐term responses to experimental climate change of vascular plant communities in these Sphagnum‐dominated ecosystems. We aimed to see how long‐term experimental climate manipulations, relevant to different predicted future climate scenarios, affect total vascular plant abundance and species composition when the community is dominated by mosses. During 8 years, we investigated how the vascular plant community of a Sphagnum fuscum‐dominated subarctic peat bog responded to six experimental climate regimes, including factorial combinations of summer as well as spring warming and a thicker snow cover. Vascular plant species composition in our peat bog was more stable than is typically observed in (sub)arctic experiments: neither changes in total vascular plant abundance, nor in individual species abundances, Shannon's diversity or evenness were found in response to the climate manipulations. For three key species (Empetrum hermaphroditum, Betula nana and S. fuscum) we also measured whether the treatments had a sustained effect on plant length growth responses and how these responses interacted. Contrasting with the stability at the community level, both key shrubs and the peatmoss showed sustained positive growth responses at the plant level to the climate treatments. However, a higher percentage of moss‐encroached E. hermaphroditum shoots and a lack of change in B. nana net shrub height indicated encroachment by S. fuscum, resulting in long‐term stability of the vascular community composition: in a warmer world, vascular species of subarctic peat bogs appear to just keep pace with growing Sphagnum in their race for space. Our findings contribute to general ecological theory by demonstrating that community resistance to environmental changes does not necessarily mean inertia in vegetation response.     

The significance of differences in the mechanisms of photosynthetic acclimation to light, nitrogen and CO2 for return on investment in leaves

1. We report changes in photosynthetic capacity of leaves developed in varying photon flux density (PFD), nitrogen supply and CO₂ concentration. We determined the relative effect of these environmental factors on photosynthetic capacity per unit leaf volume as well as the volume of tissue per unit leaf area. We calculated resource-use efficiencies from the photosynthetic capacities and measurements of leaf dry mass, carbohydrates and nitrogen content.
2. There were clear differences between the mechanisms of photosynthetic acclimation to PFD, nitrogen supply and CO₂. PFD primarily affected volume of tissue per unit area whereas nitrogen supply primarily affected photosynthetic capacity per unit volume. CO₂ concentration affected both of these parameters and interacted strongly with the PFD and nitrogen treatments.
3. Photosynthetic capacity per unit carbon invested in leaves increased in the low PFD, high nitrogen and low CO₂ treatments. Photosynthetic capacity per unit nitrogen was significantly affected only by nitrogen supply.
4. The responses to low PFD and low nitrogen appear to function to increase the efficiency of utilization of the limiting resource. However, the responses to elevated CO₂ in the high PFD and high nitrogen treatments suggest that high CO₂ can result in a situation where growth is not limited by either carbon or nitrogen supply. Limitation of growth at elevated CO₂ appears to result from internal plant factors that limit utilization of carbohydrates at sinks and/or transport of carbohydrates to sinks.     

Elevated CO2 increases nitrogen fixation and decreases soil nitrogen mineralization in Florida scrub oak

We report changes in nitrogen cycling in Florida scrub oak in response to elevated atmospheric CO₂ during the first 14 months of experimental treatment. Elevated CO₂ stimulated above-ground growth, nitrogen mass, and root nodule production of the nitrogen-fixing vine, Galactia elliottii Nuttall. During this period, elevated CO₂ reduced rates of gross nitrogen mineralization in soil, and resulted in lower recovery of nitrate on resin lysimeters. Elevated CO₂ did not alter nitrogen in the soil microbial biomass, but increased the specific rate of ammonium immobilization (NH₄⁺ immobilized per unit microbial N) measured over a 24-h period. Increased carbon input to soil through greater root growth combined with a decrease in the quality of that carbon in elevated CO₂ best explains these changes. These results demonstrate that atmospheric CO₂ concentration influences both the internal cycling of nitrogen (mineralization, immobilization, and nitrification) as well as the processes that regulate total ecosystem nitrogen mass (nitrogen fixation and nitrate leaching) in Florida coastal scrub oak. If these changes in nitrogen cycling are sustained, they could cause long-term feedbacks to the growth responses of plants to elevated CO₂. Greater nitrogen fixation and reduced leaching could stimulate nitrogen-limited plant growth by increasing the mass of labile nitrogen in the ecosystem. By contrast, reduced nitrogen mineralization and increased immobilization will restrict the supply rate of plant-available nitrogen, potentially reducing plant growth. Thus, the net feedback to plant growth will depend on the balance of these effects through time.     

The influence of cytokinin-auxin cross-regulation on cell-fate determination in Arabidopsis thaliana root development

Root growth and development in Arabidopsis thaliana are sustained by a specialised zone termed the meristem, which contains a population of dividing and differentiating cells that are functionally analogous to a stem cell niche in animals. The hormones auxin and cytokinin control meristem size antagonistically. Local accumulation of auxin promotes cell division and the initiation of a lateral root primordium. By contrast, high cytokinin concentrations disrupt the regular pattern of divisions that characterises lateral root development, and promote differentiation. The way in which the hormones interact is controlled by a genetic regulatory network. In this paper, we propose a deterministic mathematical model to describe this network and present model simulations that reproduce the experimentally observed effects of cytokinin on the expression of auxin regulated genes. We show how auxin response genes and auxin efflux transporters may be affected by the presence of cytokinin. We also analyse and compare the responses of the hormones auxin and cytokinin to changes in their supply with the responses obtained by genetic mutations of SHY2, which encodes a protein that plays a key role in balancing cytokinin and auxin regulation of meristem size. We show that although shy2 mutations can qualitatively reproduce the effect of varying auxin and cytokinin supply on their response genes, some elements of the network respond differently to changes in hormonal supply and to genetic mutations, implying a different, general response of the network. We conclude that an analysis based on the ratio between these two hormones may be misleading and that a mathematical model can serve as a useful tool for stimulate further experimental work by predicting the response of the network to changes in hormone levels and to other genetic mutations.     

Bromeliad growth and stoichiometry: responses to atmospheric nutrient supply in fog-dependent ecosystems of the hyper-arid Atacama Desert, Chile

Carbon, nitrogen, and phosphorus (C, N, P) stoichiometry influences the growth of plants and nutrient cycling within ecosystems. Indeed, elemental ratios are used as an index for functional differences between plants and their responses to natural or anthropogenic variations in nutrient supply. We investigated the variation in growth and elemental content of the rootless terrestrial bromeliad Tillandsia landbeckii, which obtains its moisture, and likely its nutrients, from coastal fogs in the Atacama Desert. We assessed (1) how fog nutrient supply influences plant growth and stoichiometry and (2) the response of plant growth and stoichiometry to variations in nutrient supply by using reciprocal transplants. We hypothesized that T. landbeckii should exhibit physiological and biochemical plastic responses commensurate with nutrient supply from atmospheric deposition. In the case of the Atacama Desert, nutrient supply from fog is variable over space and time, which suggests a relatively high variation in the growth and elemental content of atmospheric bromeliads. We found that the nutrient content of T. landbeckii showed high spatio-temporal variability, driven partially by fog nutrient deposition but also by plant growth rates. Reciprocal transplant experiments showed that transplanted individuals converged to similar nutrient content, growth rates, and leaf production of resident plants at each site, reflecting local nutrient availability. Although plant nutrient content did not exactly match the relative supply of N and P, our results suggest that atmospheric nutrient supply is a dominant driver of plant growth and stoichiometry. In fact, our results indicate that N uptake by T. landbeckii plants depends more on N supplied by fog, whereas P uptake is mainly regulated by within-plant nutrient demand for growth. Overall, these findings indicate that variation in fog nutrient supply exerts a strong control over growth and nutrient dynamics of atmospheric plants, which are ubiquitous across fog-dominated ecosystems.     

Stable Isotopes and Carbon Cycle Processes in Forests and Grasslands

Abstract: Scaling and partitioning are frequently two difficult challenges facing ecology today. With regard to ecosystem carbon balance studies, ecologists and atmospheric scientists are often interested in asking how fluxes of carbon dioxide scale across the landscape, region and continent. Yet at the same time, physiological ecologists and ecosystem ecologists are interested in dissecting the net ecosystem CO₂ exchange between the biosphere and the atmosphere to achieve a better understanding of the balance between photosynthesis and respiration within a forest. In both of these multiple-scale ecological questions, stable isotope analyses of carbon dioxide can play a central role in influencing our understanding of the extent to which terrestrial ecosystems are carbon sinks. In this synthesis, we review the theory and present field evidence to address isotopic scaling of CO₂ fluxes. We first show that the ¹³C isotopic signal which ecosystems impart to the atmosphere does not remain constant over time at either temporal or spatial scales. The relative balances of different biological activities and plant responses to stress result in dynamic changes in the ¹³C isotopic exchange between the biosphere and atmosphere, with both seasonal and stand-age factors playing major roles influencing the ¹³C biosphere-atmosphere exchange. We then examine how stable isotopes are used to partition net ecosystem exchange fluxes in order to calculate shifts in the balance of photosynthesis and respiration. Lastly, we explore how fundamental differences in the ¹⁸O isotopic gas exchange of forest and grassland ecosystems can be used to further partition terrestrial fluxes.     

A Coordination Model of Whole-plant Carbon Allocation in Relation to Water Stress

Although water is an important determinant of the allocationof material between roots and shoots during growth, and oftenparallels the effects of nitrogen, few models have explicitlyconsidered allocation in relation to water supply. We use coordinationtheory to develop a simple exponential model that considersallocation of dry matter between shoots and roots during growthin relation to carbon and watersupplies, and accounts for theeffects of water stress on growth. We compare coordinationvs.optimization(global and local) versions of the exponential model by examiningsimilarities and differences in model behaviour obtained underconstant and variable environmental conditions, and with drasticallychanging conditions (mild, moderate and severe water stress).The greatest differences between coordination and optimizationexist in the drastically changing conditions. In a second versionof the model, we remove the restriction of exponential growthand show how coordination principles can be extended to a morecomplicated structure. The non-exponential model is used toanalyse experimental data on the effects of different pot sizes(and hence water availability) on root restriction and plantgrowth as reported by Thomas and Strain (Plant Physiology96:627–634, 1991). With further refinements, the coordinationmodel has potential as a tool to model plant growth in relationto water supply under various environmental conditions. Optimization; functional balance; root: shoot ratio; root restriction     

Influence of atmospheric carbon dioxide enrichment on induced response and growth compensation after herbivore damage in Lotus corniculatus

Abstract 1. Plant growth and chemical defence compounds in four Lotus corniculatus genotypes exposed to factorial combinations of ambient and elevated carbon dioxide, and herbivory by caterpillars of Polyommatus icarus were measured to test the predictions of the carbon/nutrient balance hypothesis.
2. Shoot and root biomass, allocation to shoots versus roots, and carbon-based defence compounds were greater under elevated carbon dioxide. Pupal weight of P. icarus was greater and development time shorter under elevated carbon dioxide.
3. Herbivory decreased shoot growth relative to root growth and production of nitrogen-based defence (cyanide). Young leaves contained more defence compounds than old leaves, and this response depended on carbon dioxide and herbivory treatments (significant interactions).
4. Genotype-specific responses of plants to carbon dioxide and herbivory were found for the production of cyanide. Furthermore, maternal butterfly-specific responses of caterpillars to carbon dioxide were found for development time. This suggests the existence of genetic variation for important defence and life-history traits in plants and herbivores in response to rising carbon dioxide levels.     

Growing dendrites and axons differ in their reliance on the secretory pathway

Little is known about how the distinct architectures of dendrites and axons are established. From a genetic screen, we isolated dendritic arbor reduction (dar) mutants with reduced dendritic arbors but normal axons of Drosophila neurons. We identified dar2, dar3, and dar6 genes as the homologs of Sec23, Sar1, and Rab1 of the secretory pathway. In both Drosophila and rodent neurons, defects in Sar1 expression preferentially affected dendritic growth, revealing evolutionarily conserved difference between dendritic and axonal development in the sensitivity to limiting membrane supply from the secretory pathway. Whereas limiting ER-to-Golgi transport resulted in decreased membrane supply from soma to dendrites, membrane supply to axons remained sustained. We also show that dendritic growth is contributed by Golgi outposts, which are found predominantly in dendrites. The distinct dependence between dendritic and axonal growth on the secretory pathway helps to establish different morphology of dendrites and axons.     

Quantifying and Mapping the Supply of and Demand for Carbon Storage and Sequestration Service from Urban Trees

Studies that assess the distribution of benefits provided by ecosystem services across urban areas are increasingly common. Nevertheless, current knowledge of both the supply and demand sides of ecosystem services remains limited, leaving a gap in our understanding of balance between ecosystem service supply and demand that restricts our ability to assess and manage these services. The present study seeks to fill this gap by developing and applying an integrated approach to quantifying the supply and demand of a key ecosystem service, carbon storage and sequestration, at the local level. This approach follows three basic steps: (1) quantifying and mapping service supply based upon Light Detection and Ranging (LiDAR) processing and allometric models, (2) quantifying and mapping demand for carbon sequestration using an indicator based on local anthropogenic CO₂ emissions, and (3) mapping a supply-to-demand ratio. We illustrate this approach using a portion of the Twin Cities Metropolitan Area of Minnesota, USA. Our results indicate that 1735.69 million kg carbon are stored by urban trees in our study area. Annually, 33.43 million kg carbon are sequestered by trees, whereas 3087.60 million kg carbon are emitted by human sources. Thus, carbon sequestration service provided by urban trees in the study location play a minor role in combating climate change, offsetting approximately 1% of local anthropogenic carbon emissions per year, although avoided emissions via storage in trees are substantial. Our supply-to-demand ratio map provides insight into the balance between carbon sequestration supply in urban trees and demand for such sequestration at the local level, pinpointing critical locations where higher levels of supply and demand exist. Such a ratio map could help planners and policy makers to assess and manage the supply of and demand for carbon sequestration.     

Thermal acclimation in widespread heterotrophic soil microbes

Respiration by plants and microorganisms is primarily responsible for mediating carbon exchanges between the biosphere and atmosphere. Climate warming has the potential to influence the activity of these organisms, regulating exchanges between carbon pools. Physiological ‘down‐regulation’ of warm‐adapted species (acclimation) could ameliorate the predicted respiratory losses of soil carbon under climate change scenarios, but unlike plants and symbiotic microbes, the existence of this phenomenon in heterotrophic soil microbes remains controversial. Previous studies using complex soil microbial communities are unable to distinguish physiological acclimation from other community‐scale adjustments. We explored the temperature‐sensitivity of individual saprotrophic basidiomycete fungi growing in agar, showing definitively that these widespread heterotrophic fungi can acclimate to temperature. In almost all cases, the warm‐acclimated individuals had lower growth and respiration rates at intermediate temperatures than cold‐acclimated isolates. Inclusion of such microbial physiological responses to warming is essential to enhance the robustness of global climate‐ecosystem carbon models.