Impacts of 3 years of elevated atmospheric CO2 on rhizosphere carbon flow and microbial community dynamics

Carbon (C) uptake by terrestrial ecosystems represents an important option for partially mitigating anthropogenic CO₂ emissions. Short‐term atmospheric elevated CO₂ exposure has been shown to create major shifts in C flow routes and diversity of the active soil‐borne microbial community. Long‐term increases in CO₂ have been hypothesized to have subtle effects due to the potential adaptation of soil microorganism to the increased flow of organic C. Here, we studied the effects of prolonged elevated atmospheric CO₂ exposure on microbial C flow and microbial communities in the rhizosphere. Carex arenaria (a nonmycorrhizal plant species) and Festuca rubra (a mycorrhizal plant species) were grown at defined atmospheric conditions differing in CO₂ concentration (350 and 700 ppm) for 3 years. During this period, C flow was assessed repeatedly (after 6 months, 1, 2, and 3 years) by ¹³C pulse‐chase experiments, and label was tracked through the rhizosphere bacterial, general fungal, and arbuscular mycorrhizal fungal (AMF) communities. Fatty acid biomarker analyses and RNA‐stable isotope probing (RNA‐SIP), in combination with real‐time PCR and PCR‐DGGE, were used to examine microbial community dynamics and abundance. Throughout the experiment the influence of elevated CO₂ was highly plant dependent, with the mycorrhizal plant exerting a greater influence on both bacterial and fungal communities. Biomarker data confirmed that rhizodeposited C was first processed by AMF and subsequently transferred to bacterial and fungal communities in the rhizosphere soil. Over the course of 3 years, elevated CO₂ caused a continuous increase in the ¹³C enrichment retained in AMF and an increasing delay in the transfer of C to the bacterial community. These results show that, not only do elevated atmospheric CO₂ conditions induce changes in rhizosphere C flow and dynamics but also continue to develop over multiple seasons, thereby affecting terrestrial ecosystems C utilization processes.     

Interacting effects of atmospheric CO2 enrichment and solar radiation on growth of the aquatic fern Azolla filiculoides

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Elevated atmospheric CO2 stimulates soil fungal diversity through increased fine root production in a semiarid shrubland ecosystem

Soil fungal communities are likely to be central in mediating microbial feedbacks to climate change through their effects on soil carbon (C) storage, nutrient cycling, and plant health. Plants often produce increased fine root biomass in response to elevated atmospheric carbon dioxide (CO₂), but the responses of soil microbial communities are variable and uncertain, particularly in terms of species diversity. In this study, we describe the responses of the soil fungal community to free air CO₂ enrichment (FACE) in a semiarid chaparral shrubland in Southern California (dominated by Adenomstoma fasciculatum) using large subunit rRNA gene sequencing. Community composition varied greatly over the landscape and responses to FACE were subtle, involving a few specific groups. Increased frequency of Sordariomycetes and Leotiomycetes, the latter including the Helotiales, a group that includes many dark septate endophytes known to associate positively with roots, was observed in the FACE plots. Fungal diversity, both in terms of richness and evenness, increased consistently in the FACE treatment, and was relatively high compared to other studies that used similar methods. Increases in diversity were observed across multiple phylogenetic levels, from genus to class, and were distributed broadly across fungal lineages. Diversity was also higher in samples collected close to (5 cm) plants compared to samples in canopy gaps (30 cm away from plants). Fungal biomass correlated well with soil organic matter (SOM) content, but patterns of diversity were correlated with fine root production rather than SOM. We conclude that the fungal community in this ecosystem is tightly linked to plant fine root production, and that future changes in the fungal community in response to elevated CO₂ and other climatic changes will be primarily driven by changes in plant belowground allocation. Potential feedbacks mediated by soil fungi, such as soil C sequestration, nutrient cycling, and pathogenesis, are discussed.     

Carbon for nutrient exchange between arbuscular mycorrhizal fungi and wheat varies according to cultivar and changes in atmospheric carbon dioxide concentration

Arbuscular mycorrhizal fungi (AMF) form symbioses with most crops, potentially improving their nutrient assimilation and growth. The effects of cultivar and atmospheric CO₂ concentration ([CO₂]) on wheat–AMF carbon‐for‐nutrient exchange remain critical knowledge gaps in the exploitation of AMF for future sustainable agricultural practices within the context of global climate change. We used stable and radioisotope tracers (¹⁵N, ³³P, ¹⁴C) to quantify AMF‐mediated nutrient uptake and fungal acquisition of plant carbon in three wheat (Triticum aestivum L.) cultivars. We grew plants under current ambient (440 ppm) and projected future atmospheric CO₂ concentrations (800 ppm). We found significant ¹⁵N transfer from fungus to plant in all cultivars, and cultivar‐specific differences in total N content. There was a trend for reduced N uptake under elevated atmospheric [CO₂]. Similarly, ³³P uptake via AMF was affected by cultivar and atmospheric [CO₂]. Total P uptake varied significantly among wheat cultivars and was greater at the future than current atmospheric [CO₂]. We found limited evidence of cultivar or atmospheric [CO₂] effects on plant‐fixed carbon transfer to the mycorrhizal fungi. Our results suggest that AMF will continue to provide a route for nutrient uptake by wheat in the future, despite predicted rises in atmospheric [CO₂]. Consideration should therefore be paid to cultivar‐specific AMF receptivity and function in the development of climate smart germplasm for the future.     

Combined effects of elevated CO2 concentration and Wolbachia on Hylyphantes graminicola (Araneae: Linyphiidae)

The increasing concentration of carbon dioxide in atmosphere is not only a major cause of global warming, but it also adversely affects the ecological diversity of invertebrates. This study was conducted to evaluate the effect of elevated CO₂ concentration (ambient, 400 ppm and high, 800 ppm) and Wolbachia (Wolbachia‐infected, W⁺ and Wolbachia‐uninfected, W^?) on Hylyphantes graminicola. The total survival rate, developmental duration, carapace width and length, body weight, sex ratio, net reproductive rate, nutrition content, and enzyme activity in H. graminicola were examined under four treatments: W^? 400 ppm, W^? 800 ppm, W⁺ 400 ppm, and W⁺ 800 ppm. Results showed that Wolbachia‐infected spiders had significantly decreased the total developmental duration. Different instars showed variations up to some extent, but no obvious effect was found under elevated CO₂ concentration. Total survival rate, sex ratio, and net reproductive rate were not affected by elevated CO₂ concentration or Wolbachia infection. The carapace width of Wolbachia‐uninfected spiders decreased significantly under elevated CO₂ concentration, while the width, length and weight were not significantly affected in Wolbachia‐infected spiders reared at ambient CO₂ concentration. The levels of protein, specific activities of peroxidase, and amylase were significantly increased under elevated CO₂ concentration or Wolbachia‐infected spiders, while the total amino content was only increased in Wolbachia‐infected spiders. Thus, our current finding suggested that elevated CO₂ concentration and Wolbachia enhance nutrient contents and enzyme activity of H. graminicola and decrease development duration hence explore the interactive effects of factors which were responsible for reproduction regulation, but it also gives a theoretical direction for spider's protection in such a dynamic environment. Increased activities of enzymes and nutrients caused by Wolbachia infection aids for better survival of H. graminicola under stress.     

Fungal root colonization responses in natural grasslands after long-term exposure to elevated atmospheric CO2

Arbuscular mycorrhizae, ubiquitous mutualistic symbioses between plant roots and fungi in the order Glomales, are believed to be important controllers of plant responses to global change, in particular to elevated atmospheric CO₂. In order to test if any effects on the symbiosis can persist after long-term treatment, we examined root colonization by arbuscular mycorrhizal (AM) and other fungi of several plant species from two grassland communities after continuous exposure to elevated atmospheric CO₂ for six growing seasons in the field. For plant species from both a sandstone and a serpentine annual grassland there was evidence for changes in fungal root colonization, with changes occurring as a function of plant host species. We documented decreases in percentage nonmycorrhizal fungal root colonization in elevated CO₂ for several plant species. Total AM root colonization (%) only increased significantly for one out of the five plant species in each grassland. However, when dividing AM fungal hyphae into two groups of hyphae (fine endophyte and coarse endophyte), we could document significant responses of AM fungi that were hidden when only total percentage colonization was measured. We also documented changes in elevated CO₂ in the percentage of root colonized by both AM hyphal types simultaneously. Our results demonstrate that changes in fungal root colonization can occur after long-term CO₂ enrichment, and that the level of resolution of the study of AM fungal responses may have to be increased to uncover significant changes to the CO₂ treatment. This study is also one of the first to document compositional changes in the AM fungi colonizing roots of plants grown in elevated CO₂. Although it is difficult to relate the structural data directly to functional changes, possible implications of the observed changes for plant communities are discussed.     

Seed germination and seedling establishment of an invasive tropical tree species under different climate change scenarios

Increasing air temperature and atmospheric CO₂ levels may affect the distribution of invasive species. Whereas there is wide knowledge on the effect of global change on temperate species, responses of tropical invasive species to these two global change drivers are largely unknown. We conducted a greenhouse experiment on Terminalia catappa L. (Combretaceae), an invasive tree species on Brazilian coastal areas, to evaluate the effects of increased air temperature and CO₂ concentration on seed germination and seedling growth on the island of Santa Catarina (Florianópolis, Brazil). Seeds of the invasive tree were subjected to two temperature levels (ambient and +1.6 °C) and two CO₂ levels (ambient and ~650 ppmv) with a factorial design. Increased temperature enhanced germination rate and shortened germination time of T. catappa seeds. It also increased plant height, number of leaves and above‐ground biomass. By contrast, increased atmospheric CO₂ concentration had no significant effects, and the interaction between temperature and CO₂ concentration did not affect any of the measured traits. Terminalia catappa adapts to a relatively broad range of environmental conditions, being able to tolerate cooler temperatures in its invasive range. As T. catappa is native to tropical areas, global warming might favour its establishment along the coast of subtropical South America, while increased CO₂ levels seem not to have significant effects on seed germination or seedling growth.     

Short‐term carbon cycling responses of a mature eucalypt woodland to gradual stepwise enrichment of atmospheric CO2 concentration

Projections of future climate are highly sensitive to uncertainties regarding carbon (C) uptake and storage by terrestrial ecosystems. The Eucalyptus Free‐Air CO₂ Enrichment (EucFACE) experiment was established to study the effects of elevated atmospheric CO₂ concentrations (eCO₂) on a native mature eucalypt woodland with low fertility soils in southeast Australia. In contrast to other FACE experiments, the concentration of CO₂ at EucFACE was increased gradually in steps above ambient (+0, 30, 60, 90, 120, and 150 ppm CO₂ above ambient of ~400 ppm), with each step lasting approximately 5 weeks. This provided a unique opportunity to study the short‐term (weeks to months) response of C cycle flux components to eCO₂ across a range of CO₂ concentrations in an intact ecosystem. Soil CO₂ efflux (i.e., soil respiration or R_soil) increased in response to initial enrichment (e.g., +30 and +60 ppm CO₂) but did not continue to increase as the CO₂ enrichment was stepped up to higher concentrations. Light‐saturated photosynthesis of canopy leaves (A_sat) also showed similar stimulation by elevated CO₂ at +60 ppm as at +150 ppm CO₂. The lack of significant effects of eCO₂ on soil moisture, microbial biomass, or activity suggests that the increase in R_soil likely reflected increased root and rhizosphere respiration rather than increased microbial decomposition of soil organic matter. This rapid increase in R_soil suggests that under eCO_2, additional photosynthate was produced, transported belowground, and respired. The consequences of this increased belowground activity and whether it is sustained through time in mature ecosystems under eCO₂ are a priority for future research.     

Photosynthetic activity and proteomic analysis highlights the utilization of atmospheric CO2 by Ulva prolifera (Chlorophyta) for rapid growth

Free‐floating Ulva prolifera is one of the causative species of green tides. When green tides occur, massive mats of floating U. prolifera thalli accumulate rapidly in surface waters with daily growth rates as high as 56%. The upper thalli of the mats experience environmental changes such as the change in carbon source, high salinity, and desiccation. In this study, the photosynthetic performances of PSI and PSII in U. prolifera thalli exposed to different atmospheric carbon dioxide (CO₂) levels were measured. Changes in photosynthesis within salinity treatments and dehydration under different CO₂ concentrations were also analyzed. The results showed that PSII activity was enhanced as CO₂ increased, suggesting that CO₂ assimilation was enhanced and U. prolifera thalli can utilize CO₂ in the atmosphere directly, even when under moderate stress. In addition, changes in the proteome of U. prolifera in response to salt stress were investigated. Stress‐tolerance proteins appeared to have an important role in the response to salinity stress, whereas the abundance of proteins related to metabolism showed no significant change under low salinity treatments. These findings may be one of the main reasons for the extremely high growth rate of free‐floating U. prolifera when green tides occur.     

Effect of Elevated Atmospheric CO2 and Temperature Increases on the Severity of Basil Downy Mildew Caused by Peronospora belbahrii Under Phytotron Conditions

Three experimental trials have been carried out on the basil (Ocimum basilicum)–downy mildew (Peronospora belbahrii) pathosystem, under phytotron conditions, to evaluate the effect of simulated elevated atmospheric CO₂ concentrations and temperatures as well as that of their interaction. Six CO₂ and temperature combinations were tested to establish their effect on disease development. The photosynthetic efficiency (PI) and chlorophyll content index (CCI) of the basil plants were monitored throughout the trials. Average disease incidence was 43.8% under standard conditions (18–22°C and 400–450 ppm of CO₂)_, while average disease severity was 22.1%. In the same temperature regime, a doubled level of CO₂ caused a significant increase in both disease incidence and severity. When temperatures ranged between 18 and 26°C, CO₂ at 800–850 ppm increased disease incidence. At the highest temperatures tested, that is at 26–30°C, which are not favourable for downy mildew development, the increase in CO₂ had no significant effect on disease incidence. A decreasing trend of PI was observed for the PI values of the inoculated plants. This trend was particularly pronounced for high CO₂ levels at the end of the experiment. In the same way as for disease development, lower values were recorded for the inoculated plants at the end of the experiment at 18–22°C for both CO₂ concentrations and at 22–26°C for 850 ppm of CO₂. The non‐inoculated plants showed higher photosynthetic efficiency than the inoculated plants. Similar trends were also observed for the CCI, thus confirming that downy mildew incidence and severity, which in particular caused foliar damage at high CO₂ concentrations, led to a decrease in the physiological performances.     

Pea aphid promotes amino acid metabolism both in Medicago truncatula and bacteriocytes to favor aphid population growth under elevated CO2

Rising atmospheric CO₂ levels can dilute the nitrogen (N) resource in plant tissue, which is disadvantageous to many herbivorous insects. Aphids appear to be an exception that warrants further study. The effects of elevated CO₂ (750 ppm vs. 390 ppm) were evaluated on N assimilation and transamination by two Medicago truncatula genotypes, a N‐fixing‐deficient mutant (dnf1) and its wild‐type control (Jemalong), with and without pea aphid (Acyrthosiphon pisum) infestation. Elevated CO₂ increased population abundance and feeding efficiency of aphids fed on Jemalong, but reduced those on dnf1. Without aphid infestation, elevated CO₂ increased photosynthetic rate, chlorophyll content, nodule number, biomass, and pod number for Jemalong, but only increased pod number and chlorophyll content for dnf1. Furthermore, aphid infested Jemalong plants had enhanced activities of N assimilation‐related enzymes (glutamine synthetase, Glutamate synthase) and transamination‐related enzymes (glutamate oxalate transaminase, glutamine phenylpyruvate transaminase), which presumably increased amino acid concentration in leaves and phloem sap under elevated CO₂. In contrast, aphid infested dnf1 plants had decreased activities of N assimilation‐related enzymes and transmination‐related enzymes and amino acid concentrations under elevated CO₂. Furthermore, elevated CO₂ up‐regulated expression of genes relevant to amino acid metabolism in bacteriocytes of aphids associated with Jemalong, but down‐regulated those associated with dnf1. Our results suggest that pea aphids actively elicit host responses that promote amino acid metabolism in both the host plant and in its bacteriocytes to favor the population growth of the aphid under elevated CO₂.     

Mycelial production, spread and root colonisation by the ectomycorrhizal fungi Hebeloma crustuliniforme and Paxillus involutus under elevated atmospheric CO2

Effects of elevated atmospheric carbon dioxide (CO₂) levels on the production and spread of ectomycorrhizal fungal mycelium from colonised Scots pine roots were investigated. Pinus sylvestris (L.) Karst. seedlings inoculated with either Hebeloma crustuliniforme (Bull:Fr.) Quél. or Paxillus involutus (Fr.) Fr. were grown at either ambient (350 ppm) or elevated (700 ppm) levels of CO₂. Mycelial production was measured after 6 weeks in pots, and mycelial spread from inoculated seedlings was studied after 4 months growth in perlite in shallow boxes containing uncolonised bait seedlings. Plant and fungal biomass were analysed, as well as carbon and nitrogen content of seedling shoots. Mycelial biomass production by H. crustuliniforme was significantly greater under elevated CO₂ (up to a 3-fold increase was observed). Significantly lower concentrations and total amounts of N were found in plants exposed to elevated CO₂.     

The influence of increased temperature and carbon dioxide levels on the benthic/sea ice diatom <Emphasis Type="Italic">Navicula</Emphasis><Emphasis Type="Italic">directa</Emphasis>

Polar oceans are very susceptible to increased levels of atmospheric CO₂ and may act as the world’s largest sink for anthropogenic CO₂. Simultaneously, as atmospheric CO₂ increases, sea surface temperature rises due to global warming. These two factors are important in regulating microalgal ecophysiology, and it has been suggested that future global changes may significantly alter phytoplankton species composition. This study aims to investigate potential consequences of global change in terms of increased temperature and CO₂ enrichment on the benthic/sea ice diatom Navicula directa. In a laboratory experiment, the physiological response to elevated temperature and partial pressure of CO₂ (pCO₂) was investigated in terms of growth, photosynthetic activity and photosynthetic pigment composition. The experiment was performed under manipulated levels of pCO₂ (380 and 960 ppm) and temperature (0.5 and 4.5°C) to simulate a change from present levels to predicted levels during a worst-case scenario by the year 2100. After 7 days of treatment, no synergetic effects between temperature and pCO₂ were detected. However, elevated temperature promoted effective quantum yield of photosynthesis (∆F/F￠_m F^\prime_{\rm m} ) and increased growth rates by approximately 43%. Increased temperature also resulted in an altered pigment composition. In addition, enrichment of CO₂ appeared to reduce specific growth rates of N. directa. Even though growth rates were only reduced by approximately 5%, we hereby report that increased pCO₂ levels might also have potential negative effects on certain diatom strains.     

Elevated Atmospheric CO2 Alters Soil Microbial Communities Associated with Trembling Aspen (Populus tremuloides) Roots

Global atmospheric CO₂ levels are expected to double within the next 50 years. To assess the effects of increased atmospheric CO₂ on soil ecosystems, cloned trembling aspen (Populus tremuloides) seedlings were grown individually in 1 m³ open bottom root boxes under either elevated (720 ppm, ELEV) or ambient CO₂ (360 ppm, AMB). After 5 years, soil cores (40 cm depth) were collected from the root boxes and divided into 0–20 cm and 20–40 cm fractions. ELEV treatment resulted in significant decreases in both soil nitrate and total soil nitrogen in both the 0–20 cm and 20–40 cm soil fractions, with a 47% decrease in soil nitrate and a 50% decrease in total soil nitrogen occurring in the 0–20 cm fraction. ELEV treatment did not result in a significant change in the amount of soil microbial biomass. However, analysis of indicator phospholipid fatty acids (PLFA) indicated that ELEV treatment did result in significant increases in PLFA indicators for fungi and Gram-negative bacteria in the 0–20 cm fraction. Terminal restriction fragment length polymorphism (T-RFLP) analysis was used to analyze the composition of the soil bacterial communities (using primers targeting the 16SrRNA gene) and the soil fungal communities (using primers targeting the intergenic transcribed spacer region). T-RFLP analysis revealed shifts in both bacterial and fungal community structure, as well as increases in both bacterial and fungal species richness with ELEV treatment. These results indicated that increased atmospheric CO₂ had significant effects on both soil nutrient availability and the community composition of soil microbes associated with aspen roots.     

The effects of elevated CO2 and eutrophication on surface elevation gain in a European salt marsh

Salt marshes can play a vital role in mitigating the effects of global environmental change by dissipating incident storm wave energy and, through accretion, tracking increasing water depths consequent upon sea level rise. Atmospheric CO₂ concentrations and nutrient availability are two key variables that can affect the biological processes that contribute to marsh surface elevation gain. We measured the effects of CO₂ concentrations and nutrient availability on surface elevation change in intact mixed‐species blocks of UK salt marsh using six open‐top chambers receiving CO₂‐enriched (800 ppm) or ambient (400 ppm) air. We found more rapid surface elevation gain in elevated CO₂ conditions: an average increase of 3.4 mm over the growing season relative to ambient CO₂. Boosted regression analysis to determine the relative influence of different parameters on elevation change identified that a 10% reduction in microbial activity in elevated CO₂‐grown blocks had a positive influence on elevation. The biomass of Puccinellia maritima also had a positive influence on elevation, while other salt marsh species (e.g. Suaeda maritima) had no influence or a negative impact on elevation. Reduced rates of water use by the vegetation in the high CO₂ treatment could be contributing to elevation gain, either directly through reduced soil shrinkage or indirectly by decreasing microbial respiration rates due to lower redox levels in the soil. Eutrophication did not influence elevation change in either CO₂ treatment despite doubling aboveground biomass. The role of belowground processes (transpiration, root growth and decomposition) in the vertical adjustment of European salt marshes, which are primarily minerogenic in composition, could increase as atmospheric CO₂ concentrations rise and should be considered in future wetland models for the region. Elevated CO₂ conditions could enhance resilience in vulnerable systems such as those with low mineral sediment supply or where migration upwards within the tidal frame is constrained.     

Maximum impacts of future reforestation or deforestation on atmospheric CO2

There is scope for land‐use changes to increase or decrease CO₂ concentrations in the atmosphere over the next century. Here we make simple but robust calculations of the maximum impact of such changes. Historical land‐use changes (mostly deforestation) and fossil fuel emissions have caused an increase in atmospheric concentration of CO₂ of 90 ppm between the pre‐industrial era and year 2000. The projected range of CO₂ concentrations in 2100, under a range of emissions scenarios developed for the IPCC, is 170–600 ppm above 2000 levels. This range is mostly due to different assumptions regarding fossil fuel emissions. If all of the carbon so far released by land‐use changes could be restored to the terrestrial biosphere, atmospheric CO₂ concentration at the end of the century would be about 40–70 ppm less than it would be if no such intervention had occurred. Conversely, complete global deforestation over the same time frame would increase atmospheric concentrations by about 130–290 ppm. These are extreme assumptions; the maximum feasible reforestation and afforestation activities over the next 50 years would result in a reduction in CO₂ concentration of about 15–30 ppm by the end of the century. Thus the time course of fossil fuel emissions will be the major factor in determining atmospheric CO₂ concentrations for the foreseeable future.     

Elevated co2 has Differential Effects on Five Species of Microalgae from a Subtropical Freshwater Lake: Possible Implications for Phytoplankton Species Composition

Rising atmospheric CO₂ concentrations are predicted to have a significant impact on global phytoplankton populations. Of particular interest in freshwater systems are those species that produce toxins or impact water quality, though evidence for how these species, and many others, will respond is limited. This study investigated the effects of elevated CO₂ (1,000 ppm) relative to current atmospheric CO₂ partial pressures (400 ppm), on growth, cell size, carbon acquisition, and photophysiology of five freshwater phytoplankton species including a toxic cyanophyte, Raphidiopsis raciborskii, from Lake Wivenhoe, Australia. Effects of elevated CO₂ on growth rate varied between species; notably growth rate was considerably higher for Staurastrum sp. and significantly lower for Stichococcus sp. with a trend to lower growth rate for R. raciborskii. Surface area to volume ratio was significantly lower with elevated CO₂, for all species except Cyclotella sp. Timing of maximum cell concentrations of those genera studied in monoculture occurred in the lake in order of CO₂ affinity when free CO₂ concentrations dropped below air equilibrium. The results presented here suggest that as atmospheric levels of CO₂ rise, R. raciborskii may become less of a problem to water quality, while some species of chlorophytes may become more dominant. This has implications for stakeholders of many freshwater systems.