The importance of low atmospheric CO2 and fire in promoting the spread of grasslands and savannas

The distribution and abundance of trees can be strongly affected by disturbance such as fire. In mixed tree/grass ecosystems, recurrent grass‐fuelled fires can strongly suppress tree saplings and therefore control tree dominance. We propose that changes in atmospheric [CO₂] could influence tree cover in such metastable ecosystems by altering their postburn recovery rates relative to flammable herbaceous growth forms such as grasses. Slow sapling recovery rates at low [CO₂] would favour the spread of grasses and a reduction of tree cover. To test the possible importance of [CO₂]/fire interactions, we first used a Dynamic Global Vegetation Model (DGVM) to simulate biomass in grassy ecosystems in South Africa with and without fire. The results indicate that fire has a major effect under higher rainfall conditions suggesting an important role for fire/[CO₂] interactions. We then used a demographic model of the effects of fire on mesic savanna trees to test the importance of grass/tree differences in postburn recovery rates. We adjusted grass and tree growth in the model according to the DGVM output of net primary production at different [CO₂] relative to current conditions. The simulations predicted elimination of trees at [CO₂] typical of the last glacial period (180 ppm) because tree growth rate is too slow (15 years) to grow to a fire‐proof size of ca. 3 m. Simulated grass growth would produce an adequate fuel load for a burn in only 2 years. Simulations of preindustrial [CO₂] (270 ppm) predict occurrence of trees but at low densities. The greatest increase in trees occurs from preindustrial to current [CO₂] (360 ppm). The simulations are consistent with palaeo‐records which indicate that trees disappeared from sites that are currently savannas in South Africa in the last glacial. Savanna trees reappeared in the Holocene. There has also been a large increase in trees over the last 50–100 years. We suggest that slow tree recovery after fire, rather than differential photosynthetic efficiencies in C₃ and C₄ plants, might have been the significant factor in the Late Tertiary spread of flammable grasslands under low [CO₂] because open, high light environments would have been a prerequisite for the spread of C₄ grasses. Our simulations suggest further that low [CO₂] could have been a significant factor in the reduction of trees during glacial times, because of their slower regrowth after disturbance, with fire favouring the spread of grasses.     

Growth and nitrogen accretion of dinitrogen-fixing Alnus glutinosa (L.) Gaertn. under elevated carbon dioxide

Short-term studies of tree growth at elevated CO₂ suggest that forest productivity may increase as atmospheric CO₂ concentrations rise, although low soil N availability may limit the magnitude of this response. There have been few studies of growth and N₂ fixation by symbiotic N₂-fixing woody species under elevated CO₂ and the N inputs these plants could provide to forest ecosystems in the future. We investigated the effect of twice ambient CO₂ on growth, tissue N accretion, and N₂ fixation of nodulated Alnus glutinosa (L.) Gaertn. grown under low soil N conditions for 160 d. Root, nodule, stem, and leaf dry weight (DW) and N accretion increased significantly in response to elevated CO₂. Whole-plant biomass and N accretion increased 54% and 40%, respectively. Delta-¹⁵N analysis of leaf tissue indicated that plants from both treatments derived similar proportions of their total N from symbiotic fixation suggesting that elevated CO₂ grown plants fixed approximately 40% more N than did ambient CO₂ grown plants. Leaves from both CO₂ treatments showed similar relative declines in leaf N content prior to autumnal leaf abscission, but total N in leaf litter increased 24% in elevated compared to ambient CO₂ grown plants. These results suggest that with rising atmospheric CO₂ N₂-fixing woody species will accumulate greater amounts of biomass N through N₂ fixation and may enhance soil N levels by increased litter N inputs.     

Physiological and growth responses of two African species, Acacia karroo and Themeda triandra, to combined increases in CO2 and UV-B radiation

The interactive effects of increased carbon dioxide (CO₂) concentration and ultraviolet-B (UV-B, 280–320 nm) radiation on Acacia karroo Hayne, a C₃ tree, and Themeda triandra Forsk., a C₄ grass, were investigated. We tested the hypothesis that A. karroo would show greater CO₂-induced growth stimulation than T. triandra, which would partially explain current encroachment of A. karroo into C₄ grasslands, but that increased UV-B could mitigate this advantage. Seedlings were grown in open-top chambers in a greenhouse in ambient (360 μmol mol^-1) and elevated (650 μmol mol^-1) CO₂, combined with ambient (1.56 to 8.66 kJ m^-2 day^-1) or increased (2.22 to 11.93 kJ m^-2 day^-1) biologically effective (weighted) UV-B irradiances. After 30 weeks, elevated CO₂ had no effect on biomass of A. karroo, despite increased net CO₂ assimilation rates. Interaction between UV-B and CO₂ on stomatal conductance was found, with conductances decreasing only where elevated CO₂ and UV-B were supplied separately. Increases in water use efficiencies, foliar starch concentrations, root nodule numbers and total nodule mass were measured in elevated CO₂. Elevated UV-B caused only an increase in foliar carbon concentrations. In T. triandra, net CO₂ assimilation rates were unaffected in elevated CO₂, but stomatal conductances and foliar nitrogen concentrations decreased, and water use efficiencies increased. Biomass of all vegetative fractions, particularly leaf sheaths, was increased in elevated CO₂. and was accompanied by increased leaf blade lengths and individual leaf and leaf sheath masses. However, tiller numbers were reduced in elevated CO₂. Significantly moderating effects of elevated UV-B were apparent only in individual masses of leaf blades and sheaths, and in total sheath and shoot biomass. The direct CO₂-induced growth responses of the species therefore do not support the hypothesis of CO₂-driven woody encroachment of C₄ grasslands. Rather, differential changes in resource use efficiency between grass and woody species, or morphological responses of grass species, could alter the competitive balance. Increased UV-B radiation is unlikely to substantially alter the CO₂ response of these species.     

高浓度CO2对树木生理生态的影响研究进展

大气CO2浓度升高已成为世界范围内的重要环境问题。CO2浓度升高势必会对植物的生理生态变化产生重要影响。综述了国内外有关高浓度CO2对树木生理生态影响研究的最新进展,具体包括高浓度CO2对树木生长发育、光合和呼吸作用、抗氧化系统、树木代谢物质、挥发性有机化合物以及树木凋落物等方面的影响。高浓度CO2一般会促进树木地上植株的生长和发育,但也因树种差异而有所不同。最新研究表明,高浓度CO2促进了树木细根周转,树木根系生长在大气CO2浓度升高条件下表现为促进作用,这种作用加快了全球森林生态系统的C循环。高浓度CO2虽然在一定程度上促进树木光合速率的增加,但长期熏蒸也往往会发生光合驯化,这种现象产生的生理学机制目前仍无定论。高浓度CO2对树木呼吸作用尤其是根系呼吸的影响将是未来研究的重点和难点。高浓度CO2一般会提高树木抗氧化酶活性与抗氧化剂含量,但不同树种响应高浓度CO2的过程和机理也有所差异。研究表明,高浓度CO2一般对树木凋落物的分解产生不利影响,但也因树种而异。需要强调的是,目前关于树木地下部分、树木对高浓度CO2的适应机理和重要过程(碳氮水耦合及基因调控等)以及多个树种包括不同类型树种及不同品种之间比较研究较少;关于某一重要生理生态机制(如根系生理代谢)尤其是多个生态因子复合条件下缺乏长期深入的研究。在此基础上给出了大气CO2浓度升高下树木生理生态学研究的未来发展方向,包括高CO2浓度条件下树木根系生理代谢及机制、树木碳氮水耦合的生理过程及机制、不同生态因子复合作用对树木生理影响机制以及树木分子作用机理等方面的研究。这些研究不仅将丰富森林树木应对未来气候变化的有关科学理论,也为全球气候变化背景下实现森林树种生态功能的优化选择及森林生态系统的可持续发展与经营提供重要的生理生态学理论依据和参考。     

Growth responses of African savanna trees implicate atmospheric [CO2] as a driver of past and current changes in savanna tree cover

Atmospheric CO₂ has more than doubled since the last glacial maximum (LGM) and could double again within this century, largely due to anthropogenic activity. It has been suggested that low [CO₂] contributed to reduced tree cover in savanna and grassland biomes at LGM, and that increasing [CO₂] over the last century promoted increases in woody plants in these ecosystems over the past few decades. Despite the implications of this idea for understanding global carbon cycle dynamics and key global role of the savanna biome, there are still very few experimental studies quantifying the effects of CO₂ on tree growth and demography in savannas and grasslands. In this paper we present photosynthetic, growth and carbon allocation responses of African savanna trees (Acacia karroo and Acacia nilotica) and a C₄ grass, Themeda triandra, exposed to a gradient of CO₂ concentrations from 180 (typical of LGM) to 1000 µmol mol^?1 in open‐top chambers in a glasshouse as a first empirical test of this idea. Photosynthesis, total stem length, total stem diameter, shoot dry weight and root dry weight of the acacias increased significantly across the CO₂ gradient, saturating at higher CO₂ concentrations. After clipping to simulate fire, plants showed an even greater response in total stem length, total stem diameter and shoot dry weight, signalling the importance of re‐sprouting following disturbances such as fire or herbivory in savanna systems. Root starch (per unit root mass and total root starch per plant) increased steeply along the CO₂ gradient, explaining the re‐sprouting response. In contrast to the strong response of tree seedlings to the CO₂ gradient, grass productivity showed little variation, even at low CO₂ concentrations. These results suggest that CO₂ has significant direct effects on tree recruitment in grassy ecosystems, influencing the ability of trees to recover from fire damage and herbivory. Fire and herbivore regimes that were effective in controlling tree increases in grassy ecosystems could thus be much less effective in a CO₂‐rich world, but field‐based tests are needed to confirm this suggestion.     

Elevated CO2 and tree fecundity: the role of tree size, interannual variability, and population heterogeneity

Long‐term population effects of changes in atmospheric CO₂ will be largely determined by reproductive effort. Our research objectives were to quantify variability in seed production and rate of maturation among individual Pinus taeda L. (Pinaceae) trees growing in elevated CO₂ (ambient plus 200 μL L^?1) since 1996. Estimating tree fecundity in nature is frustrated by the difficulty of counting seeds from individual trees and the need for long‐term data. We have used a hierarchical Bayes approach to model individual tree fecundity, accounting for the complexity of experimentation in a natural setting over multiple years. The study presented here demonstrates large variability in natural fecundity rates and contributes to our understanding of how both interannual variation and population heterogeneity influence elevated CO₂ effects. We found that trees growing under elevated CO₂ matured earlier and produced more seeds and cones per unit basal area than ambient grown trees. By 2004, trees grown in high CO₂ had produced an average 300 more seeds per tree than ambient grown trees. Although there was a trend toward decreasing mean CO₂ effect (difference in fecundity between elevated and ambient treatments) over time, the hierarchical analysis indicates that this decrease comes from the emergence of a few highly fecund ambient grown trees by 2002, rather than acclimation or downregulation among the fumigated trees. The most important effect of increased CO₂ in forest ecosystems may be the increase in fecundity reported here. Although biomass responses can sometimes be large, the increase in fecundity can have long‐term impacts on forest dynamics that transcend the current generation.     

Predicting the Effects of Woody Encroachment on Mammal Communities,Grazing Biomass and Fire Frequency in African Savannas

With grasslands and savannas covering 20% of the world’s land surface, accounting for 30–35% of worldwide Net Primary Productivity and supporting hundreds of millions of people, predicting changes in tree/grass systems is priority. Inappropriate land management and rising atmospheric CO₂ levels result in increased woody cover in savannas. Although woody encroachment occurs world-wide, Africa’s tourism and livestock grazing industries may be particularly vulnerable. Forecasts of responses of African wildlife and available grazing biomass to increases in woody cover are thus urgently needed. These predictions are hard to make due to non-linear responses and poorly understood feedback mechanisms between woody cover and other ecological responders, problems further amplified by the lack of long-term and large-scale datasets. We propose that a space-for-time analysis along an existing woody cover gradient overcomes some of these forecasting problems. Here we show, using an existing woody cover gradient (0–65%) across the Kruger National Park, South Africa, that increased woody cover is associated with (i) changed herbivore assemblage composition, (ii) reduced grass biomass, and (iii) reduced fire frequency. Furthermore, although increased woody cover is associated with reduced livestock production, we found indigenous herbivore biomass (excluding elephants) remains unchanged between 20–65% woody cover. This is due to a significant reorganization in the herbivore assemblage composition, mostly as a result of meso-grazers being substituted by browsers at increasing woody cover. Our results suggest that woody encroachment will have cascading consequences for Africa’s grazing systems, fire regimes and iconic wildlife. These effects will pose challenges and require adaptation of livelihoods and industries dependent on conditions currently prevailing.     

Cheatgrass is favored by warming but not CO2 enrichment in a semi‐arid grassland

Elevated CO₂ and warming may alter terrestrial ecosystems by promoting invasive plants with strong community and ecosystem impacts. Invasive plant responses to elevated CO₂ and warming are difficult to predict, however, because of the many mechanisms involved, including modification of phenology, physiology, and cycling of nitrogen and water. Understanding the relative and interactive importance of these processes requires multifactor experiments under realistic field conditions. Here, we test how free‐air CO₂ enrichment (to 600 ppmv) and infrared warming (+1.5 °C day/3 °C night) influence a functionally and phenologically distinct invasive plant in semi‐arid mixed‐grass prairie. Bromus tectorum (cheatgrass), a fast‐growing Eurasian winter annual grass, increases fire frequency and reduces biological diversity across millions of hectares in western North America. Across 2 years, we found that warming more than tripled B. tectorum biomass and seed production, due to a combination of increased recruitment and increased growth. These results were observed with and without competition from native species, under wet and dry conditions (corresponding with tenfold differences in B. tectorum biomass), and despite the fact that warming reduced soil water. In contrast, elevated CO₂ had little effect on B. tectorum invasion or soil water, while reducing soil and plant nitrogen (N). We conclude that (1) warming may expand B. tectorum's phenological niche, allowing it to more successfully colonize the extensive, invasion‐resistant northern mixed‐grass prairie, and (2) in ecosystems where elevated CO₂ decreases N availability, CO₂ may have limited effects on B. tectorum and other nitrophilic invasive species.     

The resprouting response of co‐occurring temperate woody plant and grass species to elevated [CO2]: An insight into woody plant encroachment of grasslands

The phenomenon of woody plant thickening in grasslands has been observed globally and is likely to have widespread ecological consequences. It has been proposed that woody plant thickening is driven in part by rising atmospheric [CO₂] enhancing the resprouting ability of woody plants relative to grasses so they respond more strongly after disturbances such as herbivory and fire. The aim of this study was to examine the CO₂ effect on the resprouting ability of 16 co‐occurring temperate woody plant and grass species (eight species from each growth form). Plants were grown in a controlled glasshouse experiment under ambient (400 ppm) and elevated [CO₂] (600 ppm) for 14 weeks after which their resprouting ability was measured. Root non‐structural carbohydrate (NSC_mass) and nitrogen (N_mass) storage was used as proxies to measure the resprouting ability of woody plants while for the grasses it was measured directly. We found that both the woody plants (22% on average; P = 0.003) and grasses (20% on average; P = 0.003) produced more biomass under elevated [CO₂]. Despite the woody plants not allocating additional carbon to belowground storage under elevated [CO₂], they had significantly greater root NSC_mass (23% on average; P = 0.007) due to increased root biomass production (8% on average; P = 0.007). In contrast, root N_mass of the woody plants did not differ between CO₂ treatments (P = 0.373). Surprisingly, the resprouting ability of the grasses did not significantly differ between the CO₂ treatments (P = 0.067). These results provide evidence that the differing resprouting response of woody plants and grasses under elevated [CO₂] may be contributing to woody plant encroachment of grasslands worldwide.     

Responses of wild C4 and C3 grass (Poaceae) species to elevated atmospheric CO2 concentration: a meta-analytic test of current theories and perceptions

C4 plants contribute ≈ 20% of global gross primary productivity, and uncertainties regarding their responses to rising atmospheric CO₂ concentrations may limit predictions of future global change impacts on C4-dominated ecosystems. These uncertainties have not yet been considered rigorously due to expectations of C4 low responsiveness based on photosynthetic theory and early experiments. We carried out a literature review (1980–97) and meta-analysis in order to identify emerging patterns of C4 grass responses to elevated CO₂, as compared with those of C3 grasses. The focus was on nondomesticated Poaceae alone, to the exclusion of C4 dicotyledonous and C4 crop species. This provides a clear test, controlled for genotypic variability at family level, of differences between the CO₂-responsiveness of these functional types. Eleven responses were considered, ranging from physiological behaviour at the leaf level to carbon allocation patterns at the whole plant level. Results were also assessed in the context of environmental stress conditions (light, temperature, water and nutrient stress), and experimental growing conditions (pot size, experimental duration and fumigation method). Both C4 and C3 species increased total biomass significantly in elevated CO₂, by 33% and 44%, respectively. Differing tendencies between types in shoot structural response were revealed: C3 species showed a greater increase in tillering, whereas C4 species showed a greater increase in leaf area in elevated CO₂. At the leaf level, significant stomatal closure and increased leaf water use efficiency were confirmed in both types, and higher carbon assimilation rates were found in both C3 and C4 species (33% and 25%, respectively). Environmental stress did not alter the C4 CO₂-response, except for the loss of a significant positive CO₂-response for above-ground biomass and leaf area under water stress. In C3 species, stimulation of carbon assimilation rate was reduced by stress (overall), and nutrient stress tended to reduce the mean biomass response to elevated CO₂. Leaf carbohydrate status increased and leaf nitrogen concentration decreased significantly in elevated CO₂ only in C3 species. We conclude that the relative responses of the C4 and C3 photosynthetic types to elevated CO₂ concur only to some extent with expectations based on photosynthetic theory. The significant positive responses of C4 grass species at both the leaf and the whole plant level demand a re-evaluation of the assumption of low responsiveness in C4 plants at both levels, and not only with regard to water relations. The combined shoot structural and water use efficiency responses of these functional types will have consequential implications for the water balance of important catchments and range-lands throughout the world, especially in semiarid subtropical and temperate regions. It may be premature to predict that C4 grass species will lose their competitive advantage over C3 grass species in elevated CO₂.     

The effects of elevated CO2 on symbiotic N2 fixation: a link between the carbon and nitrogen cycles in grassland ecosystems

The response of plants to elevated CO₂ is dependent on the availability of nutrients, especially nitrogen. It is generally accepted that an increase in the atmospheric CO₂ concentration increases the C:N ratio of plant residues and exudates. This promotes temporary N-immobilization which might, in turn, reduce the availability of soil nitrogen. In addition, both a CO₂ stimulated increase in plant growth (thus requiring more nitrogen) and an increased N demand for the decomposition of soil residues with a large C:N will result under elevated CO₂ in a larger N-sink of the whole grassland ecosystem. One way to maintain the balance between the C and N cycles in elevated CO₂ would be to increase N-import to the grassland ecosystem through symbiotic N₂ fixation. Whether this might happen in the context of temperate ecosystems is discussed, by assessing the following hypothesis: i) symbiotic N₂ fixation in legumes will be enhanced under elevated CO₂, ii) this enhancement of N₂ fixation will result in a larger N-input to the grassland ecosystem, and iii) a larger N-input will allow the sequestration of additional carbon, either above or below-ground, into the ecosystem. Data from long-term experiments with model grassland ecosystems, consisting of monocultures or mixtures of perennial ryegrass and white clover, grown under elevated CO₂ under free-air or field-like conditions, supports the first two hypothesis, since: i) both the percentage and the amount of fixed N increases in white clover grown under elevated CO₂, ii) the contribution of fixed N to the nitrogen nutrition of the mixed grass also increases in elevated CO₂. Concerning the third hypothesis, an increased nitrogen input to the grassland ecosystem from N₂ fixation usually promotes shoot growth (above-ground C storage) in elevated CO₂. However, the consequences of this larger N input under elevated CO₂ on the below-ground carbon fluxes are not fully understood. On one hand, the positive effect of elevated CO₂ on the quantity of plant residues might be overwhelming and lead to an increased long-term below-ground C storage; on the other hand, the enhancement of the decomposition process by the N-rich legume material might favour carbon turn-over and, hence, limit the storage of below-ground carbon.     

First-year growth response of trees in an intact forest exposed to elevated CO2

Although elevated atmospheric CO₂ has been shown to increase growth of tree seedlings and saplings, the response of intact forest ecosystems and established trees is unclear. We report results from the first large-scale experimental system designed to study the effects of elevated CO₂ on an intact forest with the full complement of species interactions and environmental stresses. During the first year of exposure to ^ 1.5 Ë ambient CO₂, canopy loblolly pine (Pinus taeda, L.) trees increased basal area growth rate by 24% but understorey trees of loblolly pine, sweetgum (Liquidambar styraciflua L.), and red maple (Acer rubrum L.) did not respond. Winged elm (Ulmus alata Michx.) had a marginally significant increase in growth rate (P = 0.069). These data suggest that this ecosystem has the capacity to respond immediately to a step increase in atmospheric CO₂; however, as exposure time increases, nutrient limitations may reduce this initial growth stimulation.     

Aphid response to elevated ozone and CO2

Numerous reports indicate that pollution stress caused by sulphur dioxide (SO₂), oxies of nitrogen or fluorides promote aphid growth on herbaceous and woody plants. At SO₂ exposures, the response curve of aphids is bell-shaped having the peak at 100 ppb. This curvilinear response is related to physiological stress responses of host plants exposed to pollutants. On the other hand, observations of aphid performance on ozone-exposed (O₃) or elevated carbon dioxide-exposed (CO₂) plants have given very variable results. Depending on the duration and concentration of O₃ or elevated CO₂ exposure or the age of the exposed plants, aphid growth on the same plants either decreased or increased in comparison to growth on control plants grown in filtered air. The results of these studies suggest that there is no general air pollution-induced plant stress that triggers aphid outbreaks on plants. Plants grown in elevated CO₂ usually have higher C/N ratios than plants grown in current ambient CO₂ atmosphere. A reduced proportion of nitrogen in the plant foliage decreases growth of chewing herbivorous insects, but the few studies of elevated CO₂ effects on sucking insects such as aphids have not yielded similar consistent effects. The present paper reviews recent studies of elevated CO₂ effects on aphids and discusses the effects of combined elevated O₃ and CO₂ exposures on aphid performance on woody plants using pine and birch aphids as examples.     

Contemporary evolution of an invasive grass in response to elevated atmospheric CO2 at a Mojave Desert FACE site

Elevated atmospheric CO₂ has been shown to rapidly alter plant physiology and ecosystem productivity, but contemporary evolutionary responses to increased CO₂ have yet to be demonstrated in the field. At a Mojave Desert FACE (free‐air CO₂ enrichment) facility, we tested whether an annual grass weed (Bromus madritensis ssp. rubens) has evolved in response to elevated atmospheric CO₂. Within 7 years, field populations exposed to elevated CO₂ evolved lower rates of leaf stomatal conductance; a physiological adaptation known to conserve water in other desert or water‐limited ecosystems. Evolution of lower conductance was accompanied by reduced plasticity in upregulating conductance when CO₂ was more limiting; this reduction in conductance plasticity suggests that genetic assimilation may be ongoing. Reproductive fitness costs associated with this reduction in phenotypic plasticity were demonstrated under ambient levels of CO₂. Our findings suggest that contemporary evolution may facilitate this invasive species' spread in this desert ecosystem.     

Nitrification,denitrification, and related functional genes under elevated CO2: A meta-analysis in terrestrial ecosystems

Global change may have profound effects on soil nitrogen (N) cycling that can induce positive feedback to climate change through increased nitrous oxide (N₂O) emissions mediated by nitrification and denitrification. We conducted a meta-analysis of the effects of elevated CO₂ on nitrification and denitrification based on 879 observations from 58 publications and 46 independent elevated CO₂ experiments in terrestrial ecosystems. We investigated the effects of elevated CO₂ alone or combined with elevated temperature, increased precipitation, drought, and N addition. We assessed the response to elevated CO₂ of gross and potential nitrification, potential denitrification, and abundances of related functional genes (archaeal amoA, bacterial amoA, nirK, nirS, and nosZ). Elevated CO₂ increased potential nitrification (+28%) and the abundance of bacterial amoA functional gene (+62%) in cropland ecosystems. Elevated CO₂ increased potential denitrification when combined with N addition and higher precipitation (+116%). Elevated CO₂ also increased the abundance of nirK (+25%) and nirS (+27%) functional genes in terrestrial ecosystems and of nosZ (+32%) functional gene in cropland ecosystems. The increase in the abundance of nosZ under elevated CO₂ was larger at elevated temperature and high N (+62%). Four out of 14 two-way interactions tested between elevated CO₂ and elevated temperature, elevated CO₂ and increased precipitation, and elevated CO₂ and N addition were marginally significant and mostly synergistic. The effects of elevated CO₂ on potential nitrification and abundances of bacterial amoA and nirS functional genes increased with mean annual temperature and mean annual precipitation. Our meta-analysis thus suggests that warming and increased precipitation in large areas of the world could reinforce positive responses of nitrification and denitrification to elevated CO₂ and urges the need for more investigations in the tropical zone and on interactive effects among multiple global change factors, as we may largely underestimate the effects of global change on soil N₂O emissions.     

Do fires in savannas consume woody biomass? A comment on approaches to modeling savanna dynamics

Savanna ecosystems have long been fertile ground for mathematical modeling of vegetation structure and the role of resources and disturbance in tree-grass coexistence. In recent years, several authors have presented models that explore how savanna fires suppress the woody community, alter ecosystem dynamics, and promote grass persistence. We argue, however, that the assumption that fires influence savanna dynamics by consuming woody biomass may be wrong because, in reality, fires kill seedlings and saplings that constitute little biomass relative to adult trees. We present a simple alternative that separates the woody community into a subadult (fire-sensitive) class and an adult (fire-resistant) class and explore how this ecologically more realistic, but still simplified, model may provide better simulations of demographic processes and response to fires in savannas.     

Elevated atmospheric CO2 alters stomatal responses to variable sunlight in a C4 grass

Native tallgrass prairie in NE Kansas was exposed to elevated (twice ambient) or ambient atmospheric CO₂ levels in open-top chambers. Within chambers or in adjacent unchambered plots, the dominant C₄ grass, Andropogon gerardii, was subjected to fluctuations in sunlight similar to that produced by clouds or within canopy shading (full sun > 1500 μmol m⁻² s⁻¹ versus 350 μmol m⁻² s⁻¹ shade) and responses in gas exchange were measured. These field experiments demonstrated that stomatal conductance in A. gerardii achieved new steady state levels more rapidly after abrupt changes in sunlight at elevated CO₂ when compared to plants at ambient CO₂. This was due primarily to the 50% reduction in stomatal conductance at elevated CO₂, but was also a result of more rapid stomatal responses. Time constants describing stomatal responses were significantly reduced (29–33%) at elevated CO₂. As a result, water loss was decreased by as much as 57% (6.5% due to more rapid stomatal responses). Concurrent increases in leaf xylem pressure potential during periods of sunlight variability provided additional evidence that more rapid stomatal responses at elevated CO₂ enhanced plant water status. CO₂-induced alterations in the kinetics of stomatal responses to variable sunlight will likely enhance direct effects of elevated CO₂ on plant water relations in all ecosystems.     

Interaction between C4 barnyard grass and C3 upland rice under elevated CO2: Impact of mycorrhizae

Atmospheric CO₂ enrichment may impact arbuscular mycorrhizae (AM) development and function, which could have subsequent effects on host plant species interactions by differentially affecting plant nutrient acquisition. However, direct evidence illustrating this scenario is limited. We examined how elevated CO₂ affects plant growth and whether mycorrhizae mediate interactions between C₄ barnyard grass (Echinochloa crusgalli (L.) Beauv.) and C₃ upland rice (Oryza sativa L.) in a low nutrient soil. The monocultures and combinations with or without mycorrhizal inoculation were grown at ambient (400 ± 20 μmol mol^?1) and elevated CO₂ (700 ± 20 μmol mol^?1) levels. The ¹⁵N isotope tracer was introduced to quantify the mycorrhizally mediated N acquisition of plants. Elevated CO₂ stimulated the growth of C₃ upland rice but not that of C₄ barnyard grass under monoculture. Elevated CO₂ also increased mycorrhizal colonization of C₄ barnyard grass but did not affect mycorrhizal colonization of C₃ upland rice. Mycorrhizal inoculation increased the shoot biomass ratio of C₄ barnyard grass to C₃ upland rice under both CO₂ concentrations but had a greater impact under the elevated than ambient CO₂ level. Mycorrhizae decreased relative interaction index (RII) of C₃ plants under both ambient and elevated CO₂, but mycorrhizae increased RII of C₄ plants only under elevated CO₂. Elevated CO₂ and mycorrhizal inoculation enhanced ¹⁵N and total N and P uptake of C₄ barnyard grass in mixture but had no effects on N and P acquisition of C₃ upland rice, thus altering the distribution of N and P between the species in mixture. These results implied that CO₂ stimulation of mycorrhizae and their nutrient acquisition may impact competitive interaction of C₄ barnyard grass and C₃ upland rice under future CO₂ scenarios.     

Increased leaf reflectance in tropical trees under elevated CO2

Globally increasing atmospheric CO₂ concentrations are known to affect many aspects of plant physiology and development; however, little attention has been given to leaf and canopy optical properties. Three tropical trees in the Leguminosae, an important canopy tree family in many tropical forests, responded similarly to an experimental doubling of CO₂ partial pressure with a 9–23% increase in spectral leaf reflectance to light in the visible (400–700 nm) waveband. Decreased leaf chlorophyll content under elevated CO₂ may explain part of the observed increase in reflectance. However, analyses that statistically corrected for chlorophyll content effects on reflectance still indicated a significant CO₂ effect. This results, in conjunction with the spectral pattern of the response, suggests that the primary mechanism is increased optical masking of chlorophyll under elevated CO₂. The magnitude of the increase in leaf reflectance is sufficient to suggest that increased canopy reflectance of tropical forests (and possibly other terrestrial ecosystems) may be an important negative feedback in the response of global net radiative climate forcing to increasing atmospheric CO₂.     

Effects of elevated CO2 and temperature-grown red and sugar maple on gypsy moth performance

Few studies have investigated how tree species grown under elevated CO₂ and elevated temperature alter the performance of leaf‐feeding insects. The indirect effects of an elevated CO₂ concentration and temperature on leaf phytochemistry, along with potential direct effects on insect growth and consumption, may independently or interactively affect insects. To investigate this, we bagged larvae of the gypsy moth on leaves of red and sugar maple growing in open‐top chambers in four CO₂/temperature treatment combinations: (i) ambient temperature, ambient CO₂; (ii) ambient temperature, elevated CO₂ (+ 300 μL L^?1 CO₂); (iii) elevated temperature (+ 3.5°C), ambient CO₂; and (iv) elevated temperature, elevated CO₂. For both tree species, leaves grown at elevated CO₂ concentration were significantly reduced in leaf nitrogen concentration and increased in C: N ratio, while neither temperature nor its interaction with CO₂ concentration had any effect. Depending on the tree species, leaf water content declined (red maple) and carbon‐based phenolics increased (sugar maple) on plants grown in an enriched CO₂ atmosphere. The only observed effect of elevated temperature on leaf phytochemistry was a reduction in leaf water content of sugar maple leaves. Gypsy moth larval responses were dependent on tree species. Larvae feeding on elevated CO₂‐grown red maple leaves had reduced growth, while temperature had no effect on the growth or consumption of larvae. No significant effects of either temperature or CO₂ concentration were observed for larvae feeding on sugar maple leaves. Our data demonstrate strong effects of CO₂ enrichment on leaf phytochemical constituents important to folivorous insects, while an elevated temperature largely has little effect. We conclude that alterations in leaf chemistry due to an elevated CO₂ atmosphere are more important in this plant–folivorous insect system than either the direct short‐term effects of temperature on insect performance or its indirect effects on leaf chemistry.