Net primary and ecosystem production and carbon stocks of terrestrial ecosystems and their responses to climate change

Evaluating the role of terrestrial ecosystems in the global carbon cycle requires a detailed understanding of carbon exchange between vegetation, soil, and the atmosphere. Global climatic change may modify the net carbon balance of terrestrial ecosystems, causing feedbacks on atmospheric CO₂ and climate. We describe a model for investigating terrestrial carbon exchange and its response to climatic variation based on the processes of plant photosynthesis, carbon allocation, litter production, and soil organic carbon decomposition. The model is used to produce geographical patterns of net primary production (NPP), carbon stocks in vegetation and soils, and the seasonal variations in net ecosystem production (NEP) under both contemporary and future climates. For contemporary climate, the estimated global NPP is 57.0 Gt C y^–1, carbon stocks in vegetation and soils are 640 Gt C and 1358 Gt C, respectively, and NEP varies from –0.5 Gt C in October to 1.6 Gt C in July. For a doubled atmospheric CO₂ concentration and the corresponding climate, we predict that global NPP will rise to 69.6 Gt C y^–1, carbon stocks in vegetation and soils will increase by, respectively, 133 Gt C and 160 Gt C, and the seasonal amplitude of NEP will increase by 76%. A doubling of atmospheric CO₂ without climate change may enhance NPP by 25% and result in a substantial increase in carbon stocks in vegetation and soils. Climate change without CO₂ elevation will reduce the global NPP and soil carbon stocks, but leads to an increase in vegetation carbon because of a forest extension and NPP enhancement in the north. By combining the effects of CO₂ doubling, climate change, and the consequent redistribution of vegetation, we predict a strong enhancement in NPP and carbon stocks of terrestrial ecosystems. This study simulates the possible variation in the carbon exchange at equilibrium state. We anticipate to investigate the dynamic responses in the carbon exchange to atmospheric CO₂ elevation and climate change in the past and future.     

Root growth and functioning under atmospheric CO2 enrichment

This paper examines the extent to which atmospheric CO₂ enrichment may influence growth of plant roots and function in terms of uptake of water and nutrients, and carbon allocation towards symbionts. It is concluded that changes in dry matter allocation greatly depend on the experimental conditions during the experiment, the growth phase of the plant, and its morphological characteristics. Under non-limiting conditions of water and nutrients for growth, dry matter partitioning to the root is not changed by CO₂ enrichment. The increase in root/shoot ratio, frequently observed under limiting conditions of water and/or nutrients, enables the plant to explore a greater soil volume, and hence acquire more water and nutrients. However, more data on changes in dry matter allocation within the root due to atmospheric CO₂ are needed. It is concluded that nitrogen fixation is favored by CO₂ enrichment since nodule mass is increased, concomitant with an increase in root length. The papers available so far on the influence of CO₂ enrichment on mycorrhizal functioning suggest that carbon allocation to the roots might be increased, but also here more experiments are needed.Abbreviations LAR leaf area ratio - LWR leaf weight ratio - SWR stem weight ratio - RGR relative growth rate - R/S root/shoot - RWR root weight ratio     

Transport Exchange of Carbon,Nitrogen and Water in the Context of Whole Plant Growth and Functioning — Case History of a Nodulated Annual Legume

The economy of carbon, nitrogen and water during growth of nodulated, nitrogen-fixing plants of white lupin (Lupinus albus L.) was studied by measuring C, N and H₂O content of plant parts, concentrations of C and N in bleeding sap of xylem and phloem, transpirational losses of whole shoots and shoot parts, and daily exchanges of CO₂ between shoot and root parts and the surrounding atmosphere. Relationships were studied between water use and dry matter accumulation of shoot and fruits, and between net photosynthesis rate and leaf area, transpiration rate and nitrogen fixation. Conversion efficiencies were computed for utilization of net photosynthate for nitrogen fixation and for production of dry matter and protein in seeds. Partitioning of the plant's intake of C, N and H₂O was described in terms of growth, transpiration, and respiration of plant parts. An empirically-based model was developed to describe transport exchanges in xylem and phloem for a 10-day interval of growth. The model depicted quantitatively the mixtures of xylem and phloem streams which matched precisely the recorded amounts of C, N and H₂O assimilated, absorbed or consumed by the various parts of the plant. The model provided information on phloem translocation of carbon and nitrogen to roots from shoots, the cycling of carbon and nitrogen through leaves, the relationship between transpiration and nitrogen partitioning to shoot organs through the xylem, the relative amount of the plant's water budget committed to phloem translocation, and the significance of xylem to phloem transfer of nitrogen in stems as a means of supplying nitrogen to apical regions of the shoot.     

Root to shoot ratio of crops as influenced by CO2

Crops of tomorrow are likely to grow under higher levels of atmospheric CO₂. Fundamental crop growth processes will be affected and chief among these is carbon allocation. The root to shoot ratio (R:S, defined as dry weight of root biomass divided by dry weight of shoot biomass) depends upon the partitioning of photosynthate which may be influenced by environmental stimuli. Exposure of plant canopies to high CO₂ concentration often stimulates the growth of both shoot and root, but the question remains whether elevated atmospheric CO₂ concentration will affect roots and shoots of crop plants proportionally. Since elevated CO₂ can induce changes in plant structure and function, there may be differences in allocation between root and shoot, at least under some conditions. The effect of elevated atmospheric CO₂ on carbon allocation has yet to be fully elucidated, especially in the context of changing resource availability. Herein we review root to shoot allocation as affected by increased concentrations of atmospheric CO₂ and provide recommendations for further research. Review of the available literature shows substantial variation in R:S response for crop plants. In many cases (59.5%) R:S increased, in a very few (3.0%) remained unchanged, and in others (37.5%) decreased. The explanation for these differences probably resides in crop type, resource supply, and other experimental factors. Efforts to understand allocation under CO₂ enrichment will add substantially to the global change response data base.Abbreviations R:S root to shoot ratio, dry weight basis     

Carbon allocation to shoots and roots in relation to nitrogen supply is mediated by cytokinins and sucrose: Opinion

In this paper we firstly show some general responses of biomass partitioning upon nitrogen deprivation. Secondly, these responses are explained in terms of allocation of carbon and nitrogen, photosynthesis and respiration, using a simulation model. Thirdly, we present a hypothesis for the regulation of biomass partitioning to shoots and roots.Shortly after nitrogen deprivation, the relative growth rate (RGR) of the roots generally increases and thereafter decreases, whereas that of the shoot decreases immediately. The increased RGR of the root and decreased RGR of the shoot shortly after a reduction in the nitrogen supply, cause the root weight ratio (root weight per unit plant weight) to increase rapidly.We showed previously that allocation of carbon and nitrogen to shoots and roots can satisfactorily be described as a function of the internal organic plant nitrogen concentration. Using these functions in a simulation model, we analyzed why the relative growth rate of the roots increases shortly after a reduction in nitrogen supply. The model predicts that upon nitrogen deprivation, the plant nitrogen concentration and the rate of photosynthesis per unit plant weight rapidly decrease, and the allocation of recently assimilated carbon and nitrogen to roots rapidly increases. Simulations show that the increased relative growth rate of the root upon nitrogen deprivation is explained by decreased use of carbon for root respiration, due to decreased carbon costs for nitrogen uptake. The stimulation of the relative growth rate of the root is further amplified by the increased allocation of carbon and nitrogen to roots. Using the simple relation between the plant nitrogen concentration and allocation, the model describes plant responses quite realistically.Based on information in the literature and on our own experiments we hypothesize that allocation of carbon is mediated by sucrose and cytokinins. We propose that nitrogen deprivation leads to a reduced cytokinin production, a decreased rate of cytokinin export from the roots to the shoot, and decreased cytokinin concentrations. A reduced cytokinin concentration in the shoot represses cell division in leaves, whereas a low cytokinin concentration in roots neutralizes the inhibitory effect of cytokinins on cell division. A reduced rate of cell division in the leaves leads to a reduced unloading of sucrose from the phloem into the expanding cells. Consequently, the sucrose concentration in the phloem nearby the expanding cells increases, leading to an increase in turgor pressure in the phloem nearby the leaf's division zone. In the roots, cell division continues and no accumulation of sugars occurs in dividing cells, leading to only marginal changes in osmotic potential and turgor pressure in the phloem nearby the root's cell division zone. These changes in turgor pressure in the phloem of roots and sink leaves affect the turgor pressure gradients between source leaf-sink leaf and source leaf-root in such a way that relatively more carbohydrates are exported to the roots. As a consequence RWR increases after nitrogen deprivation. This hypothesis also explains the strong relationship between allocation and the plant nitrogen status.     

The Influence of Carbon Dioxide and Daily Photon-flux Density on Optimal Leaf Nitrogen Concentration and Root: Shoot Ratio

Using a cost-benefit model, the leaf nitrogen concentrationand root : shoot ratio that maximize whole-plant relative growthrate are determined as a function of the above-ground environment(integrated daily photon flux density and the concentrationof carbon dioxide at the site of fixation within the leaf).The major advantage of this approach is that it determines theadaptive significance of leaf physiology by considering thefunctional integration of leaves and roots. The predicted responseto increasing daily photon flux densities is an increase inoptimal leaf N concentration (N_opt) and a concomitant increasein root: shoot ratio. Increased carbon dioxide concentrations,on the other hand, reduce N_opt and only slightly change root:shoot ratio. The observed increase in leaf nitrogen concentrationfound in plants growing at high altitudes (low CO₂ partial pressure)is also predicted. Since these responses to light and CO₂ maximizethe whole-plant relative growth rate, the observed adjustmentsthat plants make to light and carbon dioxide concentration appearto be adaptive. We show that the relationship between photosynthesis and leafnitrogen concentration is complex and depends on the light andCO₂ levels at which photosynthesis is measured. The shape ofthis function is important in determining N_opt and the oppositeresponse of leaf nitrogen to light and carbon dioxide is shownto be the result of the different effects of light and CO₂ onthe photosynthesis-leaf nitrogen curve. Plant growth, photosynthesis, leaf nitrogen, biomass allocation, optimization, carbon dioxide light     

The interaction between elevated carbon dioxide and nitrogen nutrition: the physiological and molecular background

AGPase, ADP glucose pyrophosphorylase
GS, glutamine synthetase
GOGAT, glutamate : oxoglutarate amino transferase
NADP-ICDH, NADP-dependent isocitrate dehydrogenase
NR, nitrate reductase
OPPP, oxidative pentose phosphate pathway
3PGA, glycerate-3-phosphate
PEPCase, phosphoenolpyruvate carboxylase
Rubisco, ribulose-1,5-bisphosphate carboxylase/oxygenase
SPS, sucrose phosphate-synthase

This review first summarizes the numerous studies that have described the interaction between the nitrogen supply and the response of photosynthesis, metabolism and growth to elevated [CO₂]. The initial stimulation of photosynthesis in elevated [CO₂] is often followed by a decline of photosynthesis, that is typically accompanied by a decrease of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), an accumulation of carbohydrate especially starch, and a decrease of the nitrogen concentration in the plant. These changes are particularly marked when the nitrogen supply is low, whereas when the nitrogen supply is adequate there is no acclimation of photosynthesis, no major decrease in the internal concentration of nitrogen or the levels of nitrogen metabolites, and growth is stimulated markedly. Second, emerging evidence is discussed that signals derived from nitrate and nitrogen metabolites such as glutamine act to regulate the expression of genes involved in nitrate and ammonium uptake and assimilation, organic acid synthesis and starch accumulation, to modulate the sugar-mediated repression of the expression of genes involved in photosynthesis, and to modulate whole plant events including shoot–root allocation, root architecture and flowering. Third, increased rates of growth in elevated [CO₂] will require higher rates of inorganic nitrogen uptake and assimilation. Recent evidence is discussed that an increased supply of sugars can increase the rates of nitrate and ammonium uptake and assimilation, the synthesis of organic acid acceptors, and the synthesis of amino acids. Fourth, interpretation of experiments in elevated [CO₂] requires that the nitrogen status of the plants is monitored. The suitability of different criteria to assess the plant nitrogen status is critically discussed. Finally the review returns to experiments with elevated [CO₂] and discusses the following topics: is, and if so how, are nitrate and ammonium uptake and metabolism stimulated in elevated [CO₂], and does the result depend on the nitrogen supply? Is acclimation of photosynthesis the result of sugar-mediated repression of gene expression, end-product feedback of photosynthesis, nitrogen-induced senescence, or ontogenetic drift? Is the accumulation of starch a passive response to increased carbohydrate formation, or is it triggered by changes in the nutrient status? How do changes in sugar production and inorganic nitrogen assimilation interact in different conditions and at different stages of the life history to determine the response of whole plant growth and allocation to elevated [CO₂]?     

Responses of grassland production to single and multiple global environmental changes

In this century, increasing concentrations of carbon dioxide (CO₂) and other greenhouse gases in the Earth's atmosphere are expected to cause warmer surface temperatures and changes in precipitation patterns. At the same time, reactive nitrogen is entering natural systems at unprecedented rates. These global environmental changes have consequences for the functioning of natural ecosystems, and responses of these systems may feed back to affect climate and atmospheric composition. Here, we report plant growth responses of an ecosystem exposed to factorial combinations of four expected global environmental changes. We exposed California grassland to elevated CO₂, temperature, precipitation, and nitrogen deposition for five years. Root and shoot production did not respond to elevated CO₂ or modest warming. Supplemental precipitation led to increases in shoot production and offsetting decreases in root production. Supplemental nitrate deposition increased total production by an average of 26%, primarily by stimulating shoot growth. Interactions among the main treatments were rare. Together, these results suggest that production in this grassland will respond minimally to changes in CO₂ and winter precipitation, and to small amounts of warming. Increased nitrate deposition would have stronger effects on the grassland. Aside from this nitrate response, expectations that a changing atmosphere and climate would promote carbon storage by increasing plant growth appear unlikely to be realized in this system.     

Growth and shoot: root ratio of seedlings in relation to nutrient availability

The influence of mineral nutrient availability, light intensity and CO₂ on growth and shoot:root ratio in young plants is reviewed. Special emphasis in this evaluation is given to data from laboratory experiments with small Betula pendula plants, in which the concept of steady-state nutrition has been applied.Three distinctly different dry matter allocation patterns were observed when growth was limited by the availability of mineral nutrients: 1, Root growth was favoured when N, P or S were the major growth constraints. 2, The opposite pattern obtained when K, Mg and Mn restricted growth. 3, Shortage of Ca, Fe and Zn had almost no effect on the shoot:root ratio. The light regime had no effect on dry matter allocation except at very low photon flux densities (< 6.5 mol m^-2 day^-1), in which a small decrease in the root fraction was observed. Shortage of CO₂, on the other hand, strongly decreased root development, while an increase of the atmospheric CO₂ concentration had no influence on dry matter partitioning. An increased allocation of dry matter to below-ground parts was associated with an increased amount of starch in the tissues. Depletion of the carbohydrate stores occurred under all conditions in which root development was inhibited. It is concluded that the internal balance between labile nitrogen and carbon in the root and the shoot system determines how dry matter is being partitioned in the plant. The consistency of this statement with literature data and existing models for shoot:root regulation is examined.     

Allocation responses to CO2 enrichment and defoliation by a native annual plant Heterotheca subaxillaris

Among plants grown under enriched atmospheric CO₂, root:shoot balance (RSB) theory predicts a proportionately greater allocation of assimilate to roots than among ambient‐grown plants. Conversely, defoliation, which decreases the plant's capacity to assimilate carbon, is predicted to increase allocation to shoot. We tested these RSB predictions, and whether responses to CO₂ enrichment were modified by defoliation, using Heterotheca subaxillaris, an annual plant native to south‐eastern USA. Plants were grown under near‐ambient (400 μmol mol^?1) and enriched (700 μmol mol^?1) levels of atmospheric CO₂. Defoliation consisted of the weekly removal of 25% of each new fully expanded, but not previously defoliated, leaf from either rosette or bolted plants. In addition to dry mass measurements of leaves, stems, and roots, Kjeldahl N, protein, starch and soluble sugars were analysed in these plant components to test the hypothesis that changes in C:N uptake ratio drive shifts in root:shoot ratio. Young, rapidly growing CO₂‐enriched plants conformed to the predictions of RSB, with higher root:shoot ratio than ambient‐grown plants (P < 0.02), whereas older, slower growing plants did not show a CO₂ effect on root:shoot ratio. Defoliation resulted in smaller plants, among which both root and shoot biomass were reduced, irrespective of CO₂ treatment (P < 0.03). However, H. subaxillaris plants were able to compensate for leaf area removal through flexible shoot allocation to more leaves vs. stem (P < 0.01). Increased carbon availability through CO₂ enrichment did not enhance the response to defoliation, apparently because of complete growth compensation for defoliation, even under ambient conditions. CO₂‐enriched plants had higher rates of photosynthesis (P < 0.0001), but this did not translate into increased final biomass accumulation. On the other hand, earlier and more abundant yield of flower biomass was an important consequence of growth under CO₂ enrichment.     

Is carbon within the global terrestrial biosphere becoming more oxidized? Implications for trends in atmospheric O2

Measurements of atmospheric O₂ and CO₂ concentrations serve as a widely used means to partition global land and ocean carbon sinks. Interpretation of these measurements has assumed that the terrestrial biosphere contributes to changing O₂ levels by either expanding or contracting in size, and thus serving as either a carbon sink or source (and conversely as either an oxygen source or sink). Here, we show how changes in atmospheric O₂ can also occur if carbon within the terrestrial biosphere becomes more reduced or more oxidized, even with a constant carbon pool. At a global scale, we hypothesize that increasing levels of disturbance within many biomes has favored plant functional types with lower oxidative ratios and that this has caused carbon within the terrestrial biosphere to become increasingly more oxidized over a period of decades. Accounting for this mechanism in the global atmospheric O₂ budget may require a small increase in the size of the land carbon sink. In a scenario based on the Carnegie–Ames–Stanford Approach model, a cumulative decrease in the oxidative ratio of net primary production (NPP) (moles of O₂ produced per mole of CO₂ fixed in NPP) by 0.01 over a period of 100 years would create an O₂ disequilibrium of 0.0017 and require an increased land carbon sink of 0.1 Pg C yr⁻¹ to balance global atmospheric O₂ and CO₂ budgets. At present, however, it is challenging to directly measure the oxidative ratio of terrestrial ecosystem exchange and even more difficult to detect a disequilibrium caused by a changing oxidative ratio of NPP. Information on plant and soil chemical composition complement gas exchange approaches for measuring the oxidative ratio, particularly for understanding how this quantity may respond to various global change processes over annual to decadal timescales.     

Modelling forest response to increasing CO2 concentration under nutrient-limited conditions

The growth rates of woody plants depend on both the rate of photosynthetic carbon gain and the availability of essential nutrients. Instantaneous carbon gain is known to increase in response to increasing atmospheric CO₂ concentration, but it is uncertain whether this will translate into increased growth in the longer term under nutrient-limited conditions. An analytical model to address this question was developed by Comins & McMurtrie (1993, Ecological Applications 3, 666–681). Their model was further tested and analysed. Manipulation of various assumptions in the model revealed its key assumptions and allowed a more confident prediction of expected growth responses to CO₂ enrichment under nutrient-limited conditions. The analysis indicated that conclusions about the CO₂ sensitivity of production were strongly influenced by assumptions about the relationship between foliar and heartwood nitrogen concentrations. With heartwood nitrogen concentration proportional to foliar nitrogen concentration, the model predicted a strong response of plant productivity to increasing CO₂ concentration, whereas with heartwood nitrogen concentration set constant, the model predicted only a very slight growth response to changing CO₂ concentration. On the other hand, predictions were only slightly affected by: (1) assumptions about the extent of nitrogen retranslocation out of senescing roots and foliage or wood during heartwood formation; (2) the effects of nitrogen status on specific leaf area or (3) leaf longevity; (4) carbon allocation between different plant parts; or (5) changes in the N:C ratio of organic matter sequestered in the passive pool of soil organic matter. Modification of the effect of foliar nitrogen concentration on the light utilization coefficient had only a small effect on the CO₂ sensitivity for pines. However, this conclusion was strongly dependent on the chosen relationship between single-leaf photosynthesis and leaf nitrogen concentration. Overall, the analysis suggested that trees growing under nitrogen-limited conditions can respond to increasing atmospheric CO₂ concentration with considerable increases in growth.     

Interactive effects of growth‐limiting N supply and elevated atmospheric CO2 concentration on growth and carbon balance of Plantago major

To assess the interactions between concentration of atmospheric CO₂ and N supply, the response of Plantago major ssp. pleiosperma Pilger to a doubling of the ambient CO₂ concentration of 350 µl l^?1 was investigated in a range of exponential rates of N addition. The relative growth rate (RGR) as a function of the internal plant nitrogen concentration (N_i), was increased by elevated CO₂ at optimal and intermediate N_i. The rate of photosynthesis, expressed per unit leaf area and plotted versus N_i. was increased by 20-30% at elevated CO₂ for N_i above 30 mg N g^?1 dry weight. However, the rate of photosynthesis, expressed on a leaf dry matter basis and plotted versus N_i, was not affected by the CO₂ concentration. The allocation of dry matter between shoot and root was not affected by the CO₂ concentration at any of the N addition rates. This is in good agreement with theoretical models. based on a balance between the rate of photosynthesis of the shoot and the acquisition of N by the roots. The concentration of total nonstructural carbohydrates (TNC) was increased at elevated CO₂ and at N limitation, resulting in a shift in the partitioning of photosynthates from structural to nonstructural and, in terms of carbon balance, unproductive dry matter. The increase in concentration of TNC led to a decrease in both specific leaf area (SLA) and N_i at all levels of nutrient supply, and was the cause of the increased rate of photosynthesis per unit leaf area. Correction of the relationship between RGR and Ni for the accumulation of TNC made the effect of elevated CO₂ on the relationship between RGR and N_i disappear. We conclude that the shift in the relationship between RGR and N_i was due to the accumulation of TNC and not due to differences in physiological variables such as photosynthesis and shoot and root respiration, changes in leaf morphology or allocation of dry matter.     

Carbon use in root respiration as affected by elevated atmospheric O2

The use of fossil fuel is predicted to cause an increase of the atmospheric CO₂ concentration, which will affect the global pattern of temperature and precipitation. It is therefore essential to incorporate effects of temperature and water supply on the carbon requirement for root respiration of plants to predict effects of elevated [CO₂] on the carbon budget of natural and managed systems.There is insufficient information to support the contentention that an increase in the concentration of CO₂ in the atmosphere will enhance the CO₂ concentration in the soil to an extent that is likely to affect root respiration. Moreover, there is no convincing evidence for a direct effect of elevated atmospheric [CO₂] on the rate of root respiration per unit root mass or the fraction of carbon required for root respiration. However, there are likely to be indirect effects of elevated [CO₂] on the carbon requirement of plants in natural systems.Firstly, it is very likely that the carbon requirement of root respiration relative to that fixed in photosynthesis will increase when elevated [CO₂] induces a decrease in nutrient status of the plants. Although earlier papers have emphasized that elevated [CO₂] favours investment of biomass in roots relative to that in leaves, these are in fact indirect effects. The increase in root weight ratio is due to the more rapid depletion of nutrients in the root environment as a consequence of enhanced growth. This will decrease the specific rate of root respiration, but increase the carbon requirement as a fraction of the carbon fixed in photosynthesis. It is likely that these effects will be minor in systems where the nutrient supply is very high, e.g. in many managed arable systems, and increase with decreasing soil fertility, i.e. in many natural systems.Secondly, a decrease in rainfall in some parts of the world may cause a shortage in water supply which favours the carbon partitioning to roots. Water stress is likely to reduce rates of root respiration per unit root mass, but enhance the fraction of total assimilates required for root respiration, due to greater allocation of biomass to roots.Increased temperatures are unlikely to affect the specific rate of root respiration in all species. Broadly generalized, the effect of temperature on biomass allocation is that the relative investment of biomass in roots is lowest at a certain optimum temperature and increases at both higher and lower temperatures. The root respiration of some species acclimates to growth temperature, so that the effect of global temperature rise is entirely accounted for by the effect of temperature on biomass allocation. The specific rate of root respiration of other species will increase with global warming. In response to global warming the carbon requirement of roots is likely to decrease in temperate regions, when temperatures are suboptimal for the roots' capacity to acquire water. Here global warming will induce a smaller biomass allocation to the roots. Conversely, the carbon requirements are more likely to increase in mediterranean environments, where temperatures are often supraoptimal and a rise in temperature will induce greater allocation of biomass to the roots.     

Combined effects of elevated CO2 and soil drought on carbon and nitrogen allocation of the desert shrub Caragana intermedia

Impacts of either elevated CO₂ or drought stress on plant growth have been studied extensively, but interactive effects of these on plant carbon and nitrogen allocation is inadequately understood yet. In this study the response of the dominant desert shrub, Caragana intermedia Kuanget H.c.Fu, to the interaction of elevated CO₂ (700 ± 20 μmol mol⁻¹) and soil drought were determined in two large environmental growth chambers (18 m²). Elevated CO₂ increased the allocation of biomass and carbon into roots and the ratio of carbon to nitrogen (C:N) as well as the leaf soluble sugar content, but decreased the allocation of biomass and carbon into leaves, leaf nitrogen and leaf soluble protein concentrations. Elevated CO₂ significantly decreased the partitioning of nitrogen into leaves, but increased that into roots, especially under soil drought. Elevated CO₂ significantly decreased the carbon isotope discrimination (Δ) in leaves, but increased them in roots, and the ratio of Δ values between root and leaf, indicating an increased allocation into below-ground parts. It is concluded that stimulation of plant growth by CO₂ enrichment may be negated under soil drought, and under the future environment, elevated CO₂ may partially offset the negative effects of enhanced drought by regulating the partitioning of carbon and nitrogen.     

Carbon source–sink limitations differ between two species with contrasting growth strategies

Understanding how carbon source and sink strengths limit plant growth is a critical knowledge gap that hinders efforts to maximize crop yield. We investigated how differences in growth rate arise from source–sink limitations, using a model system comparing a fast‐growing domesticated annual barley (Hordeum vulgare cv. NFC Tipple) with a slow‐growing wild perennial relative (Hordeum bulbosum). Source strength was manipulated by growing plants at sub‐ambient and elevated CO₂ concentrations ([CO₂]). Limitations on vegetative growth imposed by source and sink were diagnosed by measuring relative growth rate, developmental plasticity, photosynthesis and major carbon and nitrogen metabolite pools. Growth was sink limited in the annual but source limited in the perennial. RGR and carbon acquisition were higher in the annual, but photosynthesis responded weakly to elevated [CO₂] indicating that source strength was near maximal at current [CO₂]. In contrast, photosynthetic rate and sink development responded strongly to elevated [CO₂] in the perennial, indicating significant source limitation. Sink limitation was avoided in the perennial by high sink plasticity: a marked increase in tillering and root:shoot ratio at elevated [CO₂], and lower non‐structural carbohydrate accumulation. Alleviating sink limitation during vegetative development could be important for maximizing growth of elite cereals under future elevated [CO₂].     

The Effect of Nitrogen Supply to Urtica dioica L. Plants on the Distribution of Assimilate Between Shoot and Roots

In this study the influence of nitrogen nutrition on the patterns of carbon distribution was investigated with Urtica dioica. The nettles were grown in sand culture at 3 levels of NO^?₃, namely 3 (low), 15 (medium) and 22 (high) mM. These levels encompassed a range within which nitrogen did not affect total biomass production. The ratio of root: shoot biomass of the low nitrogen plants was, however, significantly higher than that of the nettles grown at medium and high N supply. Carbon allocation from one leaf of each pair of leaves was examined after a ¹⁴CO₂-pulse and a subsequent ¹⁴C distribution period of one night. Only the youngest two leaf pairs did not export assimilates. Carbon (¹⁴C) export to the shoot apex and to the roots, as measured at the individual nodes responded to the nitrogen status: At medium and high nitrogen supply the 3rd, 4th and 5th leaf pairs exported to the shoot apex, while lower leaves exported to the root. At low nitrogen supply only the 3rd leaf exported towards the shoot apex. The results illustrate the plastic response of carbon distribution patterns to the nitrogen supply, even when net photosynthesis, carbon export from the source leaves and biomass production were not affected by the nitrogen supply to the plant.     

Modeled responses of terrestrial ecosystems to elevated atmospheric CO2: a comparison of simulations by the biogeochemistry models of the Vegetation/Ecosystem Modeling and Analysis Project (VEMAP)

Although there is a great deal of information concerning responses to increases in atmospheric CO₂ at the tissue and plant levels, there are substantially fewer studies that have investigated ecosystem-level responses in the context of integrated carbon, water, and nutrient cycles. Because our understanding of ecosystem responses to elevated CO₂ is incomplete, modeling is a tool that can be used to investigate the role of plant and soil interactions in the response of terrestrial ecosystems to elevated CO₂. In this study, we analyze the responses of net primary production (NPP) to doubled CO₂ from 355 to 710 ppmv among three biogeochemistry models in the Vegetation/Ecosystem Modeling and Analysis Project (VEMAP): BIOME-BGC (BioGeochemical Cycles), Century, and the Terrestrial Ecosystem Model (TEM). For the conterminous United States, doubled atmospheric CO₂ causes NPP to increase by 5% in Century, 8% in TEM, and 11% in BIOME-BGC. Multiple regression analyses between the NPP response to doubled CO₂ and the mean annual temperature and annual precipitation of biomes or grid cells indicate that there are negative relationships between precipitation and the response of NPP to doubled CO₂ for all three models. In contrast, there are different relationships between temperature and the response of NPP to doubled CO₂ for the three models: there is a negative relationship in the responses of BIOME-BGC, no relationship in the responses of Century, and a positive relationship in the responses of TEM. In BIOME-BGC, the NPP response to doubled CO₂ is controlled by the change in transpiration associated with reduced leaf conductance to water vapor. This change affects soil water, then leaf area development and, finally, NPP. In Century, the response of NPP to doubled CO₂ is controlled by changes in decomposition rates associated with increased soil moisture that results from reduced evapotranspiration. This change affects nitrogen availability for plants, which influences NPP. In TEM, the NPP response to doubled CO₂ is controlled by increased carboxylation which is modified by canopy conductance and the degree to which nitrogen constraints cause down-regulation of photosynthesis. The implementation of these different mechanisms has consequences for the spatial pattern of NPP responses, and represents, in part, conceptual uncertainty about controls over NPP responses. Progress in reducing these uncertainties requires research focused at the ecosystem level to understand how interactions between the carbon, nitrogen, and water cycles influence the response of NPP to elevated atmospheric CO₂. Received: 13 December 1996 / Accepted: 20 November 1997     

大气CO2浓度升高对土壤碳库稳定性的影响

工业革命以来,大气CO₂浓度持续上升,升高的CO₂浓度会改变植物光合产物积累、土壤碳库的碳输入和碳输出过程,进而通过影响有机碳组成和周转特征来调控土壤碳库动态变化。土壤碳库是陆地生态系统碳库的重要组成部分,其碳储量的微小变化都会对大气CO₂浓度和气候变化产生巨大影响。但目前关于CO₂浓度升高对土壤碳库动态和稳定性的影响还不清楚,很大程度上限制了预测陆地生态系统碳循环对气候变化的反馈。系统综述国内外大气CO₂浓度升高对植被生产力、植被碳输入和土壤碳库影响的研究进展,旨在揭示土壤碳库物理、化学组成以及周转特征对CO₂浓度升高的响应过程和机理,探讨CO₂升高情境下土壤微生物特征对土壤碳库稳定性的影响和驱动机制,为深入理解全球变化下的土壤碳循环特征提供理论支撑。