Meta‐analysis reveals enhanced growth of marine harmful algae from temperate regions with warming and elevated CO2 levels

Elevated pCO₂ and warming may promote algal growth and toxin production, and thereby possibly support the proliferation and toxicity of harmful algal blooms (HABs). Here, we tested whether empirical data support this hypothesis using a meta‐analytic approach and investigated the responses of growth rate and toxin content or toxicity of numerous marine and estuarine HAB species to elevated pCO₂ and warming. Most of the available data on HAB responses towards the two tested climate change variables concern dinoflagellates, as many members of this phytoplankton group are known to cause HAB outbreaks. Toxin content and toxicity did not reveal a consistent response towards both tested climate change variables, while growth rate increased consistently with elevated pCO₂. Warming also led to higher growth rates, but only for species isolated at higher latitudes. The observed gradient in temperature growth responses shows the potential for enhanced development of HABs at higher latitudes. Increases in growth rates with more CO₂ may present an additional competitive advantage for HAB species, particularly as CO₂ was not shown to enhance growth rate of other non‐HAB phytoplankton species. However, this may also be related to the difference in representation of dinoflagellate and diatom species in the respective HAB and non‐HAB phytoplankton groups. Since the proliferation of HAB species may strongly depend on their growth rates, our results warn for a greater potential of dinoflagellate HAB development in future coastal waters, particularly in temperate regions.     

Combination treatment of elevated UVB radiation,CO2 and temperature has little effect on silver birch (Betula pendula) growth and phytochemistry

Elevations of carbon dioxide, temperature and ultraviolet‐B (UBV) radiation in the growth environment may have a high impact on the accumulation of carbon in plants, and the different factors may work in opposite directions or induce additive effects. To detect the changes in the growth and phytochemistry of silver birch (Betula pendula) seedlings, six genotypes were exposed to combinations of ambient or elevated levels of CO₂, temperature and UVB radiation in top‐closed chambers for 7 weeks. The genotypes were relatively similar in their responses, and no significant interactive effects of three‐level climate factors on the measured parameters were observed. Elevated UVB had no effect on growth, nor did it alter plant responses to CO₂ and/or temperature in combined treatments. Growth in all plant parts increased under elevated CO₂, and height and stem biomass increased under elevated temperature. Increased carbon distribution to biomass did not reduce its allocation to phytochemicals: condensed tannins, most flavonols and phenolic acids accumulated under elevated CO₂ and elevated UVB, but this effect disappeared under elevated temperature. Leaf nitrogen content decreased under elevated CO₂. We conclude that, as a result of high genetic variability in phytochemicals, B. pendula seedlings have potential to adapt to the tested environmental changes. The induction in protective flavonoids under UVB radiation together with the positive impact of elevated CO₂ and temperature mitigates possible UVB stress effects, and thus atmospheric CO₂ concentration and temperature are the climate change factors that will dictate the establishment and success of birch at higher altitudes in the future.     

Dryland biological soil crust cyanobacteria show unexpected decreases in abundance under long‐term elevated CO2

Biological soil crusts (biocrusts) cover soil surfaces in many drylands globally. The impacts of 10 years of elevated atmospheric CO₂ on the cyanobacteria in biocrusts of an arid shrubland were examined at a large manipulated experiment in Nevada, USA. Cyanobacteria‐specific quantitative PCR surveys of cyanobacteria small‐subunit (SSU) rRNA genes suggested a reduction in biocrust cyanobacterial biomass in the elevated CO₂ treatment relative to the ambient controls. Additionally, SSU rRNA gene libraries and shotgun metagenomes showed reduced representation of cyanobacteria in the total microbial community. Taxonomic composition of the cyanobacteria was similar under ambient and elevated CO₂ conditions, indicating the decline was manifest across multiple cyanobacterial lineages. Recruitment of cyanobacteria sequences from replicate shotgun metagenomes to cyanobacterial genomes representing major biocrust orders also suggested decreased abundance of cyanobacteria sequences across the majority of genomes tested. Functional assignment of cyanobacteria‐related shotgun metagenome sequences indicated that four subsystem categories, three related to oxidative stress, were differentially abundant in relation to the elevated CO₂ treatment. Taken together, these results suggest that elevated CO₂ affected a generalized decrease in cyanobacteria in the biocrusts and may have favoured cyanobacteria with altered gene inventories for coping with oxidative stress.     

Embryonic response to long‐term exposure of the marine crustacean Nephrops norvegicus to ocean acidification and elevated temperature

Due to anthropogenic CO₂ emissions, our oceans have gradually become warmer and more acidic. To better understand the consequences of this, there is a need for long‐term (months) and multistressor experiments. Earlier research demonstrates that the effects of global climate change are specific to species and life stages. We exposed berried Norway lobsters (Nephrops norvegicus), during 4 months to the combination of six ecologically relevant temperatures (5–18°C) and reduced pH (by 0.4 units). Embryonic responses were investigated by quantifying proxies for development rate and fitness including: % yolk consumption, mean heart rate, rate of oxygen consumption, and oxidative stress. We found no interactions between temperature and pH, and reduced pH only affected the level of oxidative stress significantly, with a higher level of oxidative stress in the controls. Increased temperature and % yolk consumed had positive effects on all parameters except on oxidative stress, which did not change in response to temperature. There was a difference in development rate between the ranges of 5–10°C (Q₁₀: 5.4) and 10–18°C (Q₁₀: 2.9), implicating a thermal break point at 10°C or below. No thermal limit to a further increased development rate was found. The insensitivity of N. norvegicus embryos to low pH might be explained by adaptation to a pH‐reduced external habitat and/or internal hypercapnia during incubation. Our results thus indicate that this species would benefit from global warming and be able to withstand the predicted decrease in ocean pH in the next century during their earliest life stages. However, future studies need to combine low pH and elevated temperature treatments with hypoxia as hypoxic events are frequently and increasingly occurring in the habitat of benthic species.     

Rising CO2 Levels Will Intensify Phytoplankton Blooms in Eutrophic and Hypertrophic Lakes

Harmful algal blooms threaten the water quality of many eutrophic and hypertrophic lakes and cause severe ecological and economic damage worldwide. Dense blooms often deplete the dissolved CO₂ concentration and raise pH. Yet, quantitative prediction of the feedbacks between phytoplankton growth, CO₂ drawdown and the inorganic carbon chemistry of aquatic ecosystems has received surprisingly little attention. Here, we develop a mathematical model to predict dynamic changes in dissolved inorganic carbon (DIC), pH and alkalinity during phytoplankton bloom development. We tested the model in chemostat experiments with the freshwater cyanobacterium Microcystis aeruginosa at different CO₂ levels. The experiments showed that dense blooms sequestered large amounts of atmospheric CO₂, not only by their own biomass production but also by inducing a high pH and alkalinity that enhanced the capacity for DIC storage in the system. We used the model to explore how phytoplankton blooms of eutrophic waters will respond to rising CO₂ levels. The model predicts that (1) dense phytoplankton blooms in low- and moderately alkaline waters can deplete the dissolved CO₂ concentration to limiting levels and raise the pH over a relatively wide range of atmospheric CO₂ conditions, (2) rising atmospheric CO₂ levels will enhance phytoplankton blooms in low- and moderately alkaline waters with high nutrient loads, and (3) above some threshold, rising atmospheric CO₂ will alleviate phytoplankton blooms from carbon limitation, resulting in less intense CO₂ depletion and a lesser increase in pH. Sensitivity analysis indicated that the model predictions were qualitatively robust. Quantitatively, the predictions were sensitive to variation in lake depth, DIC input and CO₂ gas transfer across the air-water interface, but relatively robust to variation in the carbon uptake mechanisms of phytoplankton. In total, these findings warn that rising CO₂ levels may result in a marked intensification of phytoplankton blooms in eutrophic and hypertrophic waters.     

The effect of pre-industrial and predicted atmospheric CO2 concentrations on the development of diazotrophic and non-diazotrophic cyanobacterium: Dolichospermum circinale and Microcystis aeruginosa

Photoautotrophs are capable of consuming high quantities of CO₂, yet scant research exists examining the influence of different CO₂ concentrations on the growth of freshwater diazotrophic or non-diazotrophic cyanobacteria. In this study, we cultured two cyanobacteria taxa (Dolichospermum circinale and Microcystis aeruginosa) within controlled atmospheric CO₂ chambers at pre-industrial, and post-industrial concentrations. Biovolume and chlorophyll a (Chl-a) differed as a consequence of the adjusted CO₂ gradients. Significantly higher biovolume measurements were observed in the elevated CO₂ treatment for the diazotrophic species in the initial experiment. However, a follow-up experiment, with a corrected culture replenishment regime showed Chl-a measurements were greater for the diazotrophic and non-diazotrophic species in the elevated CO₂ treatment. Increasing CO₂ presents a risk to already compromised eutrophic and hyper-eutrophic ecosystems, and we reason increasing CO₂ concentrations could affect photosynthetic performance and CO₂ assimilation of surface dwelling cyanobacteria. Further experimental work is required to establish ecological thresholds for freshwater ecosystems, as pH levels showed a measurable reduction within the elevated CO₂ treatments. As cyanobacteria species may respond quite differently to future CO₂ concentrations similar comparative studies should be carried out that focus on CO₂ dynamics and pH. The findings of the study indicate diazotrophic cyanobacteria growth in particular may benefit from elevated atmospheric CO₂ concentrations.     

Interactive effects of elevated temperature and CO2 levels on metabolism and oxidative stress in two common marine bivalves (Crassostrea virginica and Mercenaria mercenaria)

Marine bivalves such as the hard shell clams Mercenaria mercenaria and eastern oysters Crassostrea virginica are affected by multiple stressors, including fluctuations in temperature and CO₂ levels in estuaries, and these stresses are expected to be exacerbated by ongoing global climate change. Hypercapnia (elevated CO₂ levels) and temperature stress can affect survival, growth and development of marine bivalves, but the cellular mechanisms of these effects are not yet fully understood. In this study, we investigated whether oxidative stress is implicated in cellular responses to elevated temperature and CO₂ levels in marine bivalves. We measured the whole-organism standard metabolic rate (SMR), total antioxidant capacity (TAOC), and levels of oxidative stress biomarkers in the muscle tissues of clams and oysters exposed to different temperatures (22 and 27 °C) and CO₂ levels (the present day conditions of ~ 400 ppm CO₂ and 800 ppm CO₂ predicted by a consensus business-as-usual IPCC emission scenario for the year 2100). SMR was significantly higher and the antioxidant capacity was lower in oysters than in clams. Aerobic metabolism was largely temperature-independent in these two species in the studied temperature range (22–27 °C). However, the combined exposure to elevated temperature and hypercapnia led to elevated SMR in clams indicating elevated costs of basal maintenance. No persistent oxidative stress signal (measured by the levels of protein carbonyls, and protein conjugates with malondialdehyde and 4-hydroxynonenal) was observed during the long-term exposure to moderate warming (+ 5 °C) and hypercapnia (~ 800 ppm CO₂). This indicates that long-term exposure to moderately elevated CO₂ and temperature minimally affects the cellular redox status in these bivalve species and that the earlier observed negative physiological effects of elevated CO₂ and temperature must be explained by other cellular mechanisms.     

Interspecific protection against oxidative stress: green algae protect harmful cyanobacteria against hydrogen peroxide

Oceanographic studies have shown that heterotrophic bacteria can protect marine cyanobacteria against oxidative stress caused by hydrogen peroxide (H₂O₂). Could a similar interspecific protection play a role in freshwater ecosystems? In a series of laboratory experiments and two lake treatments, we demonstrate that freshwater cyanobacteria are sensitive to H₂O₂ but can be protected by less-sensitive species such as green algae. Our laboratory results show that green algae degrade H₂O₂ much faster than cyanobacteria. Consequently, the cyanobacterium Microcystis was able to survive at higher H₂O₂ concentrations in mixtures with the green alga Chlorella than in monoculture. Interestingly, even the lysate of destructed Chlorella was capable to protect Microcystis, indicating a two-component H₂O₂ degradation system in which Chlorella provided antioxidant enzymes and Microcystis the reductants. The level of interspecific protection provided to Microcystis depended on the density of Chlorella. These findings have implications for the mitigation of toxic cyanobacterial blooms, which threaten the water quality of many eutrophic lakes and reservoirs worldwide. In several lakes, H₂O₂ has been successfully applied to suppress cyanobacterial blooms. Our results demonstrate that high densities of green algae can interfere with these lake treatments, as they may rapidly degrade the added H₂O₂ and thereby protect the bloom-forming cyanobacteria.     

Physiological ecology of desert biocrust moss following 10 years exposure to elevated CO₂: evidence for enhanced photosynthetic thermotolerance

In arid regions, biomes particularly responsive to climate change, mosses play an important biogeochemical role as key components of biocrusts. Using the biocrust moss Syntrichia caninervis collected from the Nevada Desert Free Air CO₂ Enrichment Facility, we examined the physiological effects of 10 years of exposure to elevated CO₂, and the effect of high temperature events on the photosynthetic performance of moss grown in CO₂‐enriched air. Moss exposed to elevated CO₂ exhibited a 46% decrease in chlorophyll, a 20% increase in carbon and no difference in either nitrogen content or photosynthetic performance. However, when subjected to high temperatures (35–40°C), mosses from the elevated CO₂ environment showed higher photosynthetic performance and photosystem II (PSII) efficiency compared to those grown in ambient conditions, potentially reflective of a shift in nitrogen allocation to components that offer a higher resistance of PSII to heat stress. This result suggests that mosses may respond to climate change in markedly different ways than vascular plants, and observed CO₂‐induced photosynthetic thermotolerance in S. caninervis will likely have consequences for future desert biogeochemistry.     

The interacting effects of elevated atmospheric CO<Subscript>2</Subscript> concentration,drought and leaf-to-air vapour pressure deficit on ecosystem isoprene fluxes

Isoprene is the most abundant biogenic hydrocarbon released from vegetation and it plays a major role in tropospheric chemistry. Because of its link to climate change, there is interest in understanding the relationship between CO₂, water availability and isoprene emission. We explored the effect of atmospheric elevated CO₂ concentration and its interaction with vapour pressure deficit (VPD) and water stress, on gross isoprene production (GIP) and net ecosystem exchange of CO₂ (NEE) in two Populus deltoides plantations grown at ambient and elevated atmospheric CO₂ concentration in the Biosphere 2 Laboratory facility. Although GIP and NEE showed a similar response to light and temperature, their responses to CO₂ and VPD were opposite; NEE was stimulated by elevated CO₂ and depressed by high VPD, while GIP was inhibited by elevated CO₂ and stimulated by high VPD. The difference in response between isoprene production and photosynthesis was also evident during water stress. GIP was stimulated in the short term and declined only when the stress was severe, whereas NEE started to decrease from the beginning of the experiment. This contrasting response led the carbon lost as isoprene in both the ambient and the elevated CO₂ treatments to increase as water stress progressed. Our results suggest that water limitation can override the inhibitory effect of elevated CO₂ leading to increased global isoprene emissions in a climate change scenario with warmer and drier climate.     

Climate change (elevated CO₂, elevated temperature and moderate drought) triggers the antioxidant enzymes' response of grapevine cv. Tempranillo, avoiding oxidative damage

Photosynthetic carbon fixation (A_N) and photosynthetic electron transport rate (ETR) are affected by different environmental stress factors, such as those associated with climate change. Under stress conditions, it can be generated an electron excess that cannot be consumed, which can react with O₂, producing reactive oxygen species. This work was aimed to evaluate the influence of climate change (elevated CO₂, elevated temperature and moderate drought) on the antioxidant status of grapevine (Vitis vinifera) cv. Tempranillo leaves, from veraison to ripeness. The lowest ratios between electrons generated (ETR) and consumed (A_N + respiration + photorespiration) were observed in plants treated with elevated CO₂ and elevated temperature. In partially irrigated plants under current ambient conditions, electrons not consumed seemed to be diverted to alternative ways. Oxidative damage to chlorophylls and carotenoids was not observed. However, these plants had increases in thiobarbituric acid reacting substances, an indication of lipid peroxidation. These increases matched well with an early rise of H₂O₂ and antioxidant enzyme activities, superoxide dismutase (EC 1.15.1.1), ascorbate peroxidase (EC 1.11.1.11) and catalase (EC 1.11.1.6). Enzymatic activities were maintained high until ripeness. In conclusion, plants grown under current ambient conditions and moderate drought were less efficient to cope with oxidative damage than well‐irrigated plants, and more interestingly, plants grown under moderate drought but treated with elevated CO₂ and elevated temperature were not affected by oxidative damage, mainly because of higher rates of electrons consumed in photosynthetic carbon fixation.     

Effects of elevated CO2 and temperature on leaf characteristics,photosynthesis and carbon storage in aboveground biomass of a boreal bioenergy crop (Phalaris arundinacea L.) under varying water regimes

This work examined the effects of elevated CO₂ and temperature and water regimes, alone and in interaction, on the leaf characteristics [leaf area (LA), specific leaf weight (SLW), leaf nitrogen content (N_L) based on LA], photosynthesis (light‐saturated net carbon fixation rate, P_sat) and carbon storage in aboveground biomass of leaves (C_l) and stem (C_s) for a perennial reed canary grass (Phalaris arundinacea L., Finnish local cultivar). For this purpose, plants were grown under different water regimes (ranging from high to low soil moisture) in climate‐controlled growth chambers under the elevated CO₂ and/or temperature (following a factorial design) over a whole growing season (May–September in 2009). The results showed that the elevated temperature increased the leaf growth, photosynthesis and carbon storage of aboveground biomass the most in the early growing periods, compared with ambient temperature. However, the plant growth declined rapidly thereafter with a lower carbon storage at the end of growing season. This was related to the accelerated phenology regulation and consequent earlier growth senescence. Consequently, the elevation of CO₂ increased the P_sat, LA and SLW during the growing season, with a significant concurrent increase in the carbon storage in aboveground biomass. Low soil moisture decreased the P_sat, leaf stomatal conductance, LA and carbon storage in above ground biomass compared with high and normal soil moisture. This water stress effect was the largest under the elevated temperature. The elevated CO₂ partially mitigated the adverse effects of high temperature and low soil moisture. However, the combination of elevated temperature and CO₂ did not significantly increase the carbon storage in aboveground biomass of the plants.     

Analysis of environmental drivers influencing interspecific variations and associations among bloom-forming cyanobacteria in large,shallow eutrophic lakes

Non-diazotrophic Microcystis and filamentous N₂-fixing Aphanizomenon and Dolichospermum (formerly Anabaena) co-occur or successively dominate freshwaters globally. Previous studies indicate that dual nitrogen (N) and phosphorus (P) reduction is needed to control cyanobacterial blooms; however, N limitation may cause replacement of non-N₂-fixing by N₂-fixing taxa. To evaluate potentially counterproductive scenarios, the effects of temperature, nutrients, and zooplankton on the spatio-temporal variations of cyanobacteria were investigated in three large, shallow eutrophic lakes in China. The results illustrate that the community composition of cyanobacteria is primarily driven by physical factors and the zooplankton community, and their interactions. Niche differentiation between Microcystis and two N₂-fixing taxa in Lake Taihu and Lake Chaohu was observed, whereas small temperature fluctuations in Lake Dianchi supported co-dominance. Through structural equation modelling, predictor variables were aggregated into ‘composites’ representing their combined effects on species-specific biomass. The model results showed that Microcystis biomass was affected by water temperature and P concentrations across the studied lakes. The biomass of two filamentous taxa, by contrast, exhibited lake-specific responses. Understanding of driving forces of the succession and competition among bloom-forming cyanobacteria will help to guide lake restoration in the context of climate warming and N:P stoichiometry imbalances.     

Global changes may be promoting a rise in select cyanobacteria in nutrient‐poor northern lakes

The interacting effects of global changes—including increased temperature, altered precipitation, reduced acidification and increased dissolved organic matter loads to lakes—are anticipated to create favourable environmental conditions for cyanobacteria in northern lakes. However, responses of cyanobacteria to these global changes are complex, if not contradictory. We hypothesized that absolute and relative biovolumes of cyanobacteria (both total and specific genera) are increasing in Swedish nutrient‐poor lakes and that these increases are associated with global changes. We tested these hypotheses using data from 28 nutrient‐poor Swedish lakes over 16 years (1998–2013). Increases in cyanobacteria relative biovolume were identified in 21% of the study sites, primarily in the southeastern region of Sweden, and were composed mostly of increases from three specific genera: Merismopedia, Chroococcus and Dolichospermum. Taxon‐specific changes were related to different environmental stressors; that is, increased surface water temperature favoured higher Merismopedia relative biovolume in low pH lakes with high nitrogen to phosphorus ratios, whereas acidification recovery was statistically related to increased relative biovolumes of Chroococcus and Dolichospermum. In addition, enhanced dissolved organic matter loads were identified as potential determinants of Chroococcus suppression and Dolichospermum promotion. Our findings highlight that specific genera of cyanobacteria benefit from different environmental changes. Our ability to predict the risk of cyanobacteria prevalence requires consideration of the environmental condition of a lake and the sensitivities of the cyanobacteria genera within the lake. Regional patterns may emerge due to spatial autocorrelations within and among lake history, rates and direction of environmental change and the niche space occupied by specific cyanobacteria.     

Different tolerances and responses to low temperature and darkness between waterbloom forming cyanobacterium <Emphasis Type="Italic">Microcystis</Emphasis> and a green alga <Emphasis Type="Italic">Scenedesmus</Emphasis>

The dynamics of planktonic cyanobacteria in eutrophicated freshwaters play an important role in formation of annual summer blooms, yet overwintering mechanisms of these water bloom forming cyanobacteria remain unknown. The responses to darkness and low temperature of three strains (unicellular Microcystis aeruginosa FACHB-905, colonial M. aeruginosa FACHB-938, and a green alga Scenedesmus quadricauda FACHB-45) were investigated in the present study. After a 30-day incubation under darkness and low temperature, cell morphology, cell numbers, chlorophyll a, photosynthetic activity (ETR_max and I _k), and malodialdehyde (MDA) content exhibited significant changes in Scenedesmus. In contrast, Microcystis aeruginosa cells did not change markedly in morphology, chlorophyll a, photosynthetic activity, and MDA content. The stress caused by low temperature and darkness resulted in an increase of the antioxidative enzyme-catalase (CAT) in all three strains. When the three strains re-grew under routine cultivated condition subjected to darkness and low temperature, specific growth rate of Scenedesmus was lower than that of Microcystis. Flow cytometry (FCM) examination indicated that two distinct types of metabolic response to darkness and low temperature existed in the three strains. The results from the present study reveal that the cyanobacterium Microcystis, especially colonial Microcystis, has greater endurance and adaptation ability to the stress of darkness and low temperature than the green alga Scenedesmus. Handling editor: D. Hamilton     

Genetic analysis on Dolichospermum (Cyanobacteria; sensu Anabaena) populations based on the culture-independent clone libraries revealed the dominant genotypes existing in Lake Taihu,China

Lake Taihu has been severely eutrophied during the last few decades and dense cyanobacterial blooms have led to a decrease in phytoplankton diversity. The cyanobacterial blooms in Lake Taihu were mainly composed of unicellular colony-forming Microcystis and filamentous heterocystous Dolichospermum (formerly known as planktonic species of Anabaena). In contrast to that of Microcystis spp., the fundamental knowledge about diversity, abundance and dynamics of Dolichospermum populations in Lake Taihu is lacking. The present study was conducted to understand genotypic distribution, dynamics and succession of Dolichospermum populations in Lake Taihu. By sequencing 688 internal transcribed spacer (ITS) regions between the 16S and 23S rRNA genes of Dolichospermum, we were able to confirm that all the sequences were Dolichospermum rather than Aphanizomenon. 118 different genotypes were identified from the obtained sequences, and two genotypes (W-type and L-type) were found to dominate in the lake, representing 36.6% and 26.2% of the total sequences, respectively. These two dominant genotypes of Dolichospermum displayed the significant seasonal pattern. Stepwise regressions analysis revealed that water temperature was associated with the two dominant genotypes. The combined results implied the possible existence of ecotypes in bloom-forming cyanobacteria, probably triggered by water temperature in the lake.     

Stomatal responses of Douglas-fir seedlings to elevated carbon dioxide and temperature during the third and fourth years of exposure

Two major components of climate change, increasing atmospheric [CO₂] and increasing temperature, may substantially alter the effects of water availability to plants through effects on the rate of water loss from leaves. We examined the interactive effects of elevated [CO₂] and temperature on seasonal patterns of stomatal conductance (g_s), transpiration (E) and instantaneous transpiration efficiency (ITE) in Douglas‐fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Seedlings were grown in sunlit chambers at either ambient CO₂ (AC) or ambient + 180 µmol mol^?1 CO₂ (EC), and at ambient temperature (AT) or ambient + 3·5 °C (ET) in a full‐factorial design. Needle gas exchange at the target growth conditions was measured approximately monthly over 21 months. Across the study period and across temperature treatments, growth in elevated [CO₂] decreased E by an average of 12% and increased ITE by an average of 46%. The absolute reduction of E associated with elevated [CO₂] significantly increased with seasonal increases in the needle‐to‐air vapour pressure deficit (D). Across CO₂ treatments, growth in elevated temperature increased E an average of 37%, and did not affect ITE. Combined, growth in elevated [CO₂] and elevated temperature increased E an average of 19% compared with the ACAT treatment. The CO₂ supply and growth temperature did not significantly affect stomatal sensitivity to D or the relationship between g_s and net photosynthetic rates. This study suggests that elevated [CO₂] may not completely ameliorate the effect of elevated temperature on E, and that climate change may substantially alter needle‐level water loss and water use efficiency of Douglas‐fir seedlings.     

Elevated CO2 ameliorates birch response to high temperature and frost stress: implications for modeling climate-induced geographic range shifts

Despite predictions that both atmospheric CO₂ concentrations and air temperature will rise together, very limited data are currently available to assess the possible interactive effects of these two global change factors on temperate forest tree species. Using yellow birch (Betula alleghaniensis) as a model species, we studied how elevated CO₂ (800 vs. 400 μl l⁻¹) influences seedling growth and physiological responses to a 5°C increase in summer air temperatures (31/26 vs. 26/21°C day/night), and how both elevated CO₂ and air temperature during the growing season influence seedling ability to survive freezing stress during the winter dormant season. Our results show that while increased temperature decreases seedling growth, temperature-induced growth reductions are significantly lower at elevated CO₂ concentrations (43% vs. 73%). The amelioration of high-temperature stress was related to CO₂-induced reductions in both whole-shoot dark respiration and transpiration. Our results also show that increased summer air temperature, and to a lesser degree CO₂ concentration, make dormant winter buds less susceptible to freezing stress. We show the relevance of these results to models used to predict how climate change will influence future forest species distribution and productivity, without considering the direct or interactive effects of CO_2. Received: 5 June 1997 / Accepted: 16 December 1997