A meta-analysis of elevated [CO2] effects on soybean (Glycine max) physiology, growth and yield

The effects of elevated [CO₂] on 25 variables describing soybean physiology, growth and yield are reviewed using meta‐analytic techniques. This is the first meta‐analysis to our knowledge performed on a single crop species and summarizes the effects of 111 studies. These primary studies include numerous soybean growth forms, various stress and experimental treatments, and a range of elevated [CO₂] levels (from 450 to 1250 p.p.m.), with a mean of 689 p.p.m. across all studies. Stimulation of soybean leaf CO₂ assimilation rate with growth at elevated [CO₂] was 39%, despite a 40% decrease in stomatal conductance and a 11% decrease in Rubisco activity. Increased leaf CO₂ uptake combined with an 18% stimulation in leaf area to provide a 59% increase in canopy photosynthetic rate. The increase in total dry weight was lower at 37%, and seed yield still lower at 24%. This shows that even in an agronomic species selected for maximum investment in seed, several plant level feedbacks prevent additional investment in reproduction, such that yield fails to reflect fully the increase in whole plant carbon uptake. Large soil containers (> 9 L) have been considered adequate for assessing plant responses to elevated [CO₂]. However, in open‐top chamber experiments, soybeans grown in large pots showed a significant threefold smaller stimulation in yield than soybeans grown in the ground. This suggests that conclusions about plant yield based on pot studies, even when using very large containers, are a poor reflection of performance in the absence of any physical restriction on root growth. This review supports a number of current paradigms of plant responses to elevated [CO₂]. Namely, stimulation of photosynthesis is greater in plants that fix N and have additional carbohydrate sinks in nodules. This supports the notion that photosynthetic capacity decreases when plants are N‐limited, but not when plants have adequate N and sink strength. The root : shoot ratio did not change with growth at elevated [CO₂], sustaining the charge that biomass allocation is unaffected by growth at elevated [CO₂] when plant size and ontogeny are considered.     

Global response of terrestrial ecosystem structure and function to CO2 and climate change: results from six dynamic global vegetation models

The possible responses of ecosystem processes to rising atmospheric CO₂ concentration and climate change are illustrated using six dynamic global vegetation models that explicitly represent the interactions of ecosystem carbon and water exchanges with vegetation dynamics. The models are driven by the IPCC IS92a scenario of rising CO₂ ( Wigley et al. 1991 ), and by climate changes resulting from effective CO₂ concentrations corresponding to IS92a, simulated by the coupled ocean atmosphere model HadCM2‐SUL. Simulations with changing CO₂ alone show a widely distributed terrestrial carbon sink of 1.4–3.8 Pg C y^?1 during the 1990s, rising to 3.7–8.6 Pg C y^?1 a century later. Simulations including climate change show a reduced sink both today (0.6–3.0 Pg C y^?1) and a century later (0.3–6.6 Pg C y^?1) as a result of the impacts of climate change on NEP of tropical and southern hemisphere ecosystems. In all models, the rate of increase of NEP begins to level off around 2030 as a consequence of the ‘diminishing return’ of physiological CO₂ effects at high CO₂ concentrations. Four out of the six models show a further, climate‐induced decline in NEP resulting from increased heterotrophic respiration and declining tropical NPP after 2050. Changes in vegetation structure influence the magnitude and spatial pattern of the carbon sink and, in combination with changing climate, also freshwater availability (runoff). It is shown that these changes, once set in motion, would continue to evolve for at least a century even if atmospheric CO₂ concentration and climate could be instantaneously stabilized. The results should be considered illustrative in the sense that the choice of CO₂ concentration scenario was arbitrary and only one climate model scenario was used. However, the results serve to indicate a range of possible biospheric responses to CO₂ and climate change. They reveal major uncertainties about the response of NEP to climate change resulting, primarily, from differences in the way that modelled global NPP responds to a changing climate. The simulations illustrate, however, that the magnitude of possible biospheric influences on the carbon balance requires that this factor is taken into account for future scenarios of atmospheric CO₂ and climate change.     

博斯腾湖人工和天然芦苇湿地土壤CO2、CH4和N2O排放通量

为研究干旱区淡水湖泊人工、天然芦苇湿地土壤温室气体源汇强度及其影响因素,采用静态箱-气相色谱法,于2015年1月—12月对博斯腾湖人工和天然芦苇湿地土壤CO_2、CH_4和N_2O通量进行全年观测。结果表明,人工芦苇湿地土壤CO_2、CH_4和N_2O排放通量变化范围分别为:10.1—588.4mg m~(-2)h~(-1)、2.9—82.4μg m~(-2)h~(-1)和1.32—29.7μg m~(-2)h~(-1),天然芦苇湿地土壤CO_2、CH_4和N_2O排放通量变化范围分别为10.3—469.6mg m~(-2)h~(-1)、3.1—64.8μg m~(-2)h~(-1)和1.9—14.3μg m~(-2)h~(-1)。人工和天然芦苇湿地夏季土壤CO_2排放通量均明显高于其他季节,而土壤CH_4和N_2O排放通量较大值多集中在春末夏初。全年观测期间,人工芦苇湿地土壤CO_2、CH_4和N_2O排放通量高于天然芦苇湿地(P0.05);温度是影响人工、天然芦苇湿地土壤CO_2和N_2O排放通量的关键因素,近地面温度和5cm土壤温度与CO_2和N_2O排放通量呈现极显著的正相关关系(P0.01)。土壤CH_4排放通量是温度和水分二者共同影响的,由近地表温度、5cm土壤温度和土壤含水量共同拟合的方程可以分别解释人工、天然芦苇湿地土壤CH_4排放通量的71%、74.5%;土壤有机碳、pH、盐分、NH_4~+-N、NO_3~--N也是人工、天然芦苇湿地土壤CO_2、CH_4和N_2O排放通量的影响因素;人工和天然芦苇湿地土壤均是CO_2、CH_4和N_2O的"源"。基于100年尺度,由3种温室气体计算全球增温潜势得出,人工芦苇湿地全球增温潜势大于天然芦苇湿地(15150.18kg/hm~212484.21kg/hm~2)。     

Elevated atmospheric CO2: effects on phytochemistry, insect performance and insect-parasitoid interactions

This study was conducted to examine the effects of CO₂-mediated changes in tree chemistry on the performance of the gypsy moth ((Lymantria dispar L.) and the parasitold Cotesia melanoscela (Ratz.). We used carbon-nutrient balance theory to develop hypotheses regarding changes in tree chemistry and the performance of both insects under elevated CO₂. As predicted, levels of foliar nitrogen declined and concentrations of carbon-based compounds (e.g. starch and phenolics) increased under elevated CO₂. Gypsy moth performance (e.g. growth, development) was altered by CO₂-mediated changes in foliar chemistry, but the magnitude was small and varied across tree species. Larvae feeding on high CO₂ aspen exhibited the largest reduction in performance, relative to larvae feeding on birch, oak, or maple. Parasitism by C. melanoscela significantly prolonged gypsy moth development and reduced growth rates. Overall, the effect of parasitism on gypsy moth performance did not differ between CO₂ treatments. Altered gypsy moth performance on high CO₂ foliage in turn affected parasitoid performance, but the response was variable: parasitoid mortality increased and adult female size declined slightly under high CO₂, while development time and adult male size were unaffected. Our results suggest that CO₂-induced changes in plant chemistry were buffered to the extent that effects on third trophic level interactions were weak to non-existent for the system examined in this study.     

Warming and elevated CO2 affect the relationship between seed mass, germinability and seedling growth in Austrodanthonia caespitosa, a dominant Australian grass

While the influence of elevated CO₂ on the production, mass and quality of plant seeds has been well studied, the effect of warming on these characters is largely unknown; and there is practically no information on possible interactions between warming and elevated CO₂, despite the importance of these characters in population maintenance and recovery. Here, we present the impacts of elevated CO₂ and warming, both in isolation and combination, on seed production, mass, quality, germination success and subsequent seedling growth of Austrodanthonia caespitosa , a dominant temperate C₃ grass from Australia, using seeds collected from the TasFACE experiment. Mean seed production and mass were not significantly affected by either elevated CO₂ or warming, but elevated CO₂ more than doubled the proportion of very light, inviable seeds ( P < 0.05) and halved mean seed N concentration ( P < 0.04) and N content ( P < 0.03). The dependence of seed germination success on seed mass was affected by an elevated CO₂× warming interaction ( P < 0.004), such that maternal exposure to elevated CO₂ or warming reduced germination if applied in isolation, but not when applied in combination. Maternal effects were retained when seedlings were grown in a common environment for 6 weeks, with seedlings descended from warmed plants 20% smaller ( P < 0.008) with a higher root : shoot ratio ( P < 0.001) than those from unwarmed plants. Given that both elevated CO₂ and warming reduced seed mass, quality, germinability or seedling growth, it is likely that global change will reduce population growth or distribution of this dominant species.     

Effects of species composition, land surface cover, CO2 concentration and climate on isoprene emissions from European forests.

Emissions of isoprene from terrestrial vegetation are known to affect atmospheric chemical properties, like its oxidation capacity or the concentration of tropospheric ozone. The latter is of concern, since besides being a potent greenhouse gas, O(3) is toxic for humans, animals, and plants even at relatively low concentrations. Isoprene-emitting forests in the vicinity of NO(x) pollution sources (like cities) can contribute considerably to O(3) formation, and to the peak concentrations observed during hot summer weather. The biogenic contribution to O(3) concentrations is generally thought to increase in a future, warmer climate--pushing values beyond health thresholds possibly even more frequently and over larger areas--given that emissions of isoprene are highly temperature-dependent but also because of the CO(2) fertilisation of forest productivity and leaf growth. Most projections of future emissions, however, do not include the possible CO(2)-inhibition of leaf isoprene metabolism. We explore the regional distribution of emissions from European woody vegetation, using a mechanistic isoprene-dynamic vegetation model framework. We investigate the interactive effects of climate and CO(2) concentration on forest productivity, species composition, and isoprene emissions for the periods 1981-2000 and 2081-2100. Our projection of future emissions includes a direct CO(2)-isoprene inhibition. Across the model domain, we show that this direct effect has the potential to offset the stimulation of emissions that could be expected from warmer temperatures and from the increased productivity and leaf area of emitting vegetation. Changes in forest species composition that may result from climate change can play a substantial additional role in a region's future emissions. Changes in forest area or area planted in woody biofuels in general are not noticeable in the overall European forest isoprene budget, but--as was the case for changes in species composition--may substantially affect future projections in some regions of the continent.     

Genetic variation in germination, growth, and survivorship of red maple in response to subambient through elevated atmospheric CO2

Genetic variation in plant response to atmospheric carbon dioxide (CO₂) may have influenced paleo‐vegetation dynamics and could determine how future elevated CO₂ drives plant evolution and ecosystem productivity. We established how levels of relatedness – the maternal family, population, and provenance – affect variation in the CO₂ response of a species. This 2‐year growth chamber experiment focused on the germination, growth, biomass allocation, and survivorship responses of Acer rubrum to four concentrations of CO₂: 180, 270, 360, and 600 μL L^?1– representing Pleistocene through potential future conditions. We found that all levels of relatedness interacted with CO₂ to contribute to variation in response. Germination responses to CO₂ varied among families and populations, growth responses depended on families and regions of origin, and survivorship responses to CO₂ were particularly affected by regional identities. Differences among geographic regions accounted for 23% of the variation in biomass response to CO₂. If seeds produced under subambient CO₂ conditions behave similarly, our results suggest that A. rubrum may have experienced strong selection on seedling survivorship at Pleistocene CO₂ levels. Further, this species may evolve in response to globally rising CO₂ so as to increase productivity above that experimentally observed today. Species responses to future atmospheric CO₂ and the accompanying biotic effects on the global carbon cycle will vary among families, populations, and provenances.     

Simple carbon assimilation response functions from atmospheric CO2, and daily temperature and shortwave radiation

A global ‘CO₂ fertilizer effect’ multiplier is often used in crop or ecosystem models because of its simplicity. However, this approach does not take into account the interaction between CO₂, temperature and light on assimilation. This omission can lead to significant under- or overestimation of the magnitude of beneficial effects from elevated CO₂, depending on environmental conditions. We use a mechanistic model of the biochemistry of photosynthesis to represent the response of net assimilation to different levels of CO₂, temperature and radiation, on the daily time scale. Instantaneous assimilation rates for an idealized canopy model are integrated through diurnal cycles of environmental variables derived from historical climate data at three locations in North America. The calculated CO₂ fertilizer effect is greatest at high light and warm temperatures. The results are summarized by assimilation response surfaces specified by the CO₂ concentration, the canopy leaf area index, and by daily values of temperature and radiation available from climatic records. These summary functions are suitable for incorporation into crop or ecosystem models for predicting carbon assimilation or biomass production on a daily time step. An example application of the function reveals that for a relatively cool, high latitude location, the beneficial effects from a CO₂ doubling would be negligible during the early spring, even assuming a + 4°C global warming scenario. In contrast, the beneficial effects from increasing CO₂ at a relatively warm, lower latitude location are greatest in the spring, but decline in late summer because of excessively warm temperatures with a + 4°C global warming.     

Atmospheric CO2, plant nitrogen status and the susceptibility of plants to an acute increase in temperature

Abstract. Elevated levels of CO₂ in the atmosphere are expected to affect plant performance and may alter global temperature patterns. Changes in mean air temperatures that might be induced by rising levels of CO₂ and other greenhouse gases could also be accompanied by increased variability in daily temperatures such that acute increases in air temperature may be more likely than at present. Consequently, we investigated whether plants grown in a CO₂ enriched atmosphere would be differently affected by a heat shock than plants grown at ambient CO₂ levels. Plants of a C₃ annual (Abutilon theophrasti), a C₃ annual crop (Sinapis alba) and a C₄ annual (Amaranthus retroflexus) were grown from seed in growth chambers under either 400 or 700cm³ m^?3 CO₂, and were fertilized with either a high or low nutrient regime. Young seedlings of S. alba, as well as plants of all species in either the vegetative or reproductive phase of growth were exposed to a 4-h heat shock in which the temperature was raised an additional 14–23°C (depending on plant age). Total biomass and reproductive biomass were examined to determine the effect of CO₂, nutrient and heat shock treatments on plant performance. Heat shock, CO₂, and nutrient treatments, all had some significant effects on plant performance, but plants from both CO₂ treatments responded similarly to heat shocks. We also found, as expected, that plants grown under high CO₂ had dramatically decreased tissue N concentrations relative to plants grown under ambient conditions. We predicted that high-CO₂-grown plants would be more susceptible to a heat shock than ambient-CO₂-grown plants, because the reduced N concentrations of high-CO₂ grown plants could result in the reduced synthesis of heat shock proteins and reduced thermotolerance. Although we did not examine heat shock proteins, our results showed little relationship between plant nitrogen status and the ability of a plant to tolerate an acute increase in temperature.     

Interactive effects of low atmospheric CO2 and elevated temperature on growth, photosynthesis and respiration in Phaseolus vulgaris

For most of the past 250 000 years, atmospheric CO₂ has been 30–50% lower than the current level of 360 μmol CO₂ mol^–1 air. Although the effects of CO₂ on plant performance are well recognized, the effects of low CO₂ in combination with abiotic stress remain poorly understood. In this study, a growth chamber experiment using a two-by-two factorial design of CO₂ (380 μmol mol^–1, 200 μmol mol^–1) and temperature (25/20 °C day/night, 36/29 °C) was conducted to evaluate the interactive effects of CO₂ and temperature variation on growth, tissue chemistry and leaf gas exchange of Phaseolus vulgaris. Relative to plants grown at 380 μmol mol^–1 and 25/20 °C, whole plant biomass was 36% less at 380 μmol mol^–1× 36/29 °C, and 37% less at 200 μmol mol^–1× 25/20 °C. Most significantly, growth at 200 μmol mol^–1× 36/29 °C resulted in 77% less biomass relative to plants grown at 380 μmol mol^–1× 25/20 °C. The net CO₂ assimilation rate of leaves grown in 200 μmol mol^–1× 25/20 °C was 40% lower than in leaves from 380 μmol mol^–1× 25/20 °C, but similar to leaves in 200 μmol mol^–1× 36/29 °C. The leaves produced in low CO₂ and high temperature respired at a rate that was double that of leaves from the 380μmol mol^–1× 25/20 °C treatment. Despite this, there was little evidence that leaves at low CO₂ and high temperature were carbohydrate deficient, because soluble sugars, starch and total non-structural carbohydrates of leaves from the 200μmol mol^–1× 36/29 °C treatment were not significantly different in leaves from the 380μmol mol^–1× 25/20 °C treatment. Similarly, there was no significant difference in percentage root carbon, leaf chlorophyll and leaf/root nitrogen between the low CO₂× high temperature treatment and ambient CO₂ controls. Decreased plant growth was correlated with neither leaf gas exchange nor tissue chemistry. Rather, leaf and root growth were the most affected responses, declining in equivalent proportions as total biomass production. Because of this close association, the mechanisms controlling leaf and root growth appear to have the greatest control over the response to heat stress and CO₂ reduction in P. vulgaris.     

CO2 balance of boreal, temperate, and tropical forests derived from a global database

Terrestrial ecosystems sequester 2.1 Pg of atmospheric carbon annually. A large amount of the terrestrial sink is realized by forests. However, considerable uncertainties remain regarding the fate of this carbon over both short and long timescales. Relevant data to address these uncertainties are being collected at many sites around the world, but syntheses of these data are still sparse. To facilitate future synthesis activities, we have assembled a comprehensive global database for forest ecosystems, which includes carbon budget variables (fluxes and stocks), ecosystem traits (e.g. leaf area index, age), as well as ancillary site information such as management regime, climate, and soil characteristics. This publicly available database can be used to quantify global, regional or biome‐specific carbon budgets; to re‐examine established relationships; to test emerging hypotheses about ecosystem functioning [e.g. a constant net ecosystem production (NEP) to gross primary production (GPP) ratio]; and as benchmarks for model evaluations. In this paper, we present the first analysis of this database. We discuss the climatic influences on GPP, net primary production (NPP) and NEP and present the CO₂ balances for boreal, temperate, and tropical forest biomes based on micrometeorological, ecophysiological, and biometric flux and inventory estimates. Globally, GPP of forests benefited from higher temperatures and precipitation whereas NPP saturated above either a threshold of 1500 mm precipitation or a mean annual temperature of 10 °C. The global pattern in NEP was insensitive to climate and is hypothesized to be mainly determined by nonclimatic conditions such as successional stage, management, site history, and site disturbance. In all biomes, closing the CO₂ balance required the introduction of substantial biome‐specific closure terms. Nonclosure was taken as an indication that respiratory processes, advection, and non‐CO₂ carbon fluxes are not presently being adequately accounted for.     

Spatial variation in regional CO2 exchange for the Kuparuk River Basin, Alaska over the summer growing season

The spatial and temporal patterns in CO₂ flux for the Kuparuk River Basin, a 9200‐km² watershed located in NE Alaska were estimated using the Regional Arctic CO₂ Exchange Simulator (RACES) for the 1994–1995 growing seasons. RACES uses non‐linear models and a Geographical Information System database (GIS) consisting of the normalized difference vegetation index (NDVI) and dynamic temperature and radiation maps. The spatial and temporal patterns in the NDVI during both growing seasons suggest that ecosystem development occurred 2–4 weeks earlier and was relatively more rapid in the southern portion of the Kuparuk River Basin. Rates of gross primary production (GPP) and whole‐ecosystem respiration (R) were 2–4 fold higher in the southern basin than along the arctic coastal plain depending on time of year. The higher rate of GPP estimated for the southern basin was primarily due to higher NDVI values, while the higher R estimated for the southern basin was due in part to higher temperature and the NDVI. While GPP and R showed strong latitudinal trends, spatial and temporal trends in net ecosystem CO₂ exchange (NEE) were much more variable. Thus, while spatial trends in carbon gain (GPP) and loss (R) were highly correlated, small spatial and temporal differences in these large fluxes (GPP and/or R) lead to corresponding large spatial variations in the NEE.     

Photosynthesis, light and nitrogen relationships in a young deciduous forest canopy under open-air CO2 enrichment

Leaf photosynthesis (Ps), nitrogen (N) and light environment were measured on Populus tremuloides trees in a developing canopy under free‐air CO₂ enrichment in Wisconsin, USA. After 2 years of growth, the trees averaged 1·5 and 1·6 m tall under ambient and elevated CO₂, respectively, at the beginning of the study period in 1999. They grew to 2·6 and 2·9 m, respectively, by the end of the 1999 growing season. Daily integrated photon flux from cloud‐free days (PPFD_day,sat) around the lowermost branches was 16·8 ± 0·8 and 8·7 ± 0·2% of values at the top for the ambient and elevated CO₂ canopies, respectively. Elevated CO₂ significantly decreased leaf N on a mass, but not on an area, basis. N per unit leaf area was related linearly to PPFD_day,sat throughout the canopies, and elevated CO₂ did not affect that relationship. Leaf Ps light‐response curves responded differently to elevated CO₂, depending upon canopy position. Elevated CO₂ increased Ps_sat only in the upper (unshaded) canopy, whereas characteristics that would favour photosynthesis in shade were unaffected by elevated CO₂. Consequently, estimated daily integrated Ps on cloud‐free days (Ps_day,sat) was stimulated by elevated CO₂ only in the upper canopy. Ps_day,sat of the lowermost branches was actually lower with elevated CO₂ because of the darker light environment. The lack of CO₂ stimulation at the mid‐ and lower canopy was probably related to significant down‐regulation of photosynthetic capacity; there was no down‐regulation of Ps in the upper canopy. The relationship between Ps_day,sat and leaf N indicated that N was not optimally allocated within the canopy in a manner that would maximize whole‐canopy Ps or photosynthetic N use efficiency. Elevated CO₂ had no effect on the optimization of canopy N allocation.     

A long-term association between global temperature and biodiversity, origination and extinction in the fossil record

The past relationship between global temperature and levels of biological diversity is of increasing concern due to anthropogenic climate warming. However, no consistent link between these variables has yet been demonstrated. We analysed the fossil record for the last 520 Myr against estimates of low latitude sea surface temperature for the same period. We found that global biodiversity (the richness of families and genera) is related to temperature and has been relatively low during warm 'greenhouse' phases, while during the same phases extinction and origination rates of taxonomic lineages have been relatively high. These findings are consistent for terrestrial and marine environments and are robust to a number of alternative assumptions and potential biases. Our results provide the first clear evidence that global climate may explain substantial variation in the fossil record in a simple and consistent manner. Our findings may have implications for extinction and biodiversity change under future climate warming.