Oxidative Stress in Submerged Cultures of Fungi

ABSTRACT:?

It has been known for many years that oxygen (O₂) may have toxic effects on aerobically growing microorganisms, mainly due to the threat arising from reactive oxygen species (ROS). In submerged culture industrial fermentation processes, maintenance of adequate levels of O₂ (usually measured as dissolved oxygen tension (DOT)) can often be critical to the success of the manufacturing process. In viscous cultures of filamentous cultures, actively respiring, supplying adequate levels of O₂ to the cultures by conventional air sparging is difficult and various strategies have been adopted to improve or enhance O₂ transfer. However, adoption of those strategies to maintain adequate levels of DOT, that is, to avoid O₂ limitation, may expose the fungi to potential oxidative damage caused by enhanced flux through the respiratory system. In the past, there have been numerous studies investigating the effects of DOT on fungal bioprocesses. Generally, in these studies moderately enhanced levels of O₂ supply resulted in improvement in growth, product formation and acceptable morphological changes, while the negative impact of higher levels of DOT on morphology and product synthesis were generally assumed to be a consequence of “oxidative stress.” However, very little research has actually been focused on investigation of this implicit link, and the mechanisms by which such effects might be mediated within industrial fungal processes. To elucidate this neglected topic, this review first surveys the basic knowledge of the chemistry of ROS, defensive systems in fungi and the effects of DOT on fungal growth, metabolism and morphology. The physiological responses of fungal cells to oxidative stress imposed by artificial and endogenous stressors are then critically reviewed. It is clear that fungi have a range of methods available to minimize the negative impacts of elevated ROS, but also that development of the various defensive systems or responses, can itself have profound consequences upon many process-related parameters. It is also clear that many of the practically convenient and widely used experimental methods of simulating oxidative stress, for example, addition of exogenous menadione or hydrogen peroxide, have effects on fungal cultures quite distinct from the effects of elevated levels of O₂, and care must thus be exercised in the interpretation of results from such studies. The review critically evaluates our current understanding of the responses of fungal cultures to elevated O₂ levels, and highlights key areas requiring further research to remedy gaps in knowledge.     

Species richness and nitrogen supply regulate the productivity and respiration of ectomycorrhizal fungi in pure culture

The effects of biodiversity of aboveground organisms have been widely investigated in a range of ecosystems, yet whether similar responses are also seen in belowground microbial communities, such as ectomycorrhizal (EM) fungi, are little understood. We investigated, in vitro, the effects of a gradient of 1–8 species of EM fungi interacting with substratum carbon:nitrogen (C:N) ratio on biomass production and CO₂ efflux. The model experimental systems enabled us to recover and measure biomass of individuals within communities and calculate net selection and complementarity effects. Both biomass and CO₂ efflux increased with species richness particularly under high N concentrations. Moreover, net biodiversity effects were largely positive, driven by both selection and complementarity effects. Our results reveal, in pure culture, the implications of EM species richness on community productivity and C cycling, particularly under high N conditions, and constitute the basis for future experiments under natural conditions.     

Effects of elevated [CO2] at the community level mediated by root symbionts

This review examines the effects of elevated [CO₂] on plant symbioses with mycorrhizal fungi and root nodule bacteria, with emphasis on community and ecosystem processes. The effects of elevated [CO₂] on the relationships between single plant species and root symbionts are considered first. There is some evidence that plant infection by and/or biomass of root symbionts are stimulated by elevated [CO₂], but growth enhancement of the host seemingly depends on its degree of dependence on symbiosis and on soil nutrient availability. Second, the effects of elevated [CO₂] on the relationships between plant multispecies assemblages and soil, and likely impacts on above-ground and belowground diversity, are analysed. Experimental and modelling work have suggested the existence of complex feedbacks in the responses of plants and the rhizosphere to CO₂ enrichment. By modifying C inputs from plants to soil, elevated [CO₂] may affect the biomass, the infectivity, and the species/isolate composition of root symbionts. This has the potential to alter community structure and ecosystem functioning. Finally, the incorporation of type and degree of symbiotic dependence into the definition of plant functional types, and into experimental work within the context of global change research, are discussed. More experimental work on the effects of elevated [CO₂] at the community/ecosystem level, explicitly considering the role of root symbioses, is urgently needed.     

Mycorrhizal dynamics under elevated CO2 and nitrogen fertilization in a warm temperate forest

We examined the response of mycorrhizal fungi to free-air CO₂ enrichment (FACE) and nitrogen (N) fertilization in a warm temperate forest to better understand potential influences over plant nutrient uptake and soil carbon (C) storage. In particular, we hypothesized that mycorrhizal fungi and glomalin would become more prevalent under elevated CO₂ but decrease under N fertilization. In addition, we predicted that N fertilization would mitigate any positive effects of elevated CO₂ on mycorrhizal abundance. Overall, we observed a 14% increase in ectomycorrhizal (ECM) root colonization under CO₂ enrichment, which implies that elevated CO₂ results in greater C investments in these fungi. Arbuscular mycorrhizal (AM) hyphal length and glomalin stocks did not respond substantially to CO₂ enrichment, and effects of CO₂ on AM root colonization varied by date. Nitrogen effects on AM fungi were not consistent with our hypothesis, as we found an increase in AM colonization under N fertilization. Lastly, neither glomalin concentrations nor ECM colonization responded significantly to N fertilization or to an N-by-CO₂ interaction. A longer duration of N fertilization may be required to detect effects on these parameters.     

Mycorrhizas and global environmental change: research at different scales

Global environmental change (GEC), in particular rising atmospheric CO₂ concentration and temperature, will affect most ecosystems. The varied responses of plants to these aspects of GEC are well documented. As with other key below-ground components of terrestrial ecosystems, the response of the ubiquitous mycorrhizal fungal root symbionts has received limited attention. Most of the research on the effects of GEC on mycorrhizal fungi has been pot-based with a few field (especially monoculture) studies. A major question that arises in all these studies is whether the GEC effects on the mycorrhizal fungi are independent of the effects on their plant hosts. We evaluate the current knowledge on the effects of elevated CO₂ and increased temperature on mycorrhizal fungi and focus on the few available field examples. The value of using long-term and large-scale field experiments is emphasised. We conclude that the laboratory evidence to date shows that the effect of elevated CO₂ on mycorrhizal fungi is dependent on plant growth and that temperature effects seen in the past might have reflected a similar dependence. Therefore, how temperature directly affects mycorrhizal fungi remains unknown. In natural ecosystems, we predict that GEC effects on mycorrhizal fungal communities will be strongly mediated by the effects on plant communities to the extent that community level interactions will prove to be the key mechanism for determining GEC-induced changes in mycorrhizal fungal communities.     

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.     

Detecting changes in soil carbon in CO2 enrichment experiments

After four growing seasons, elevated CO₂ did not significantly alter surface soil C pools in two intact annual grasslands. However, soil C pools in these systems are large compared to the likely changes caused by elevated CO₂. We calculated statistical power to detect changes in soil C, using an approach applicable to all elevated CO₂ experiments. The distinctive isotopic signature of the fossil-fuel-derived CO₂ added to the elevated CO₂ treatment provides a C tracer to determine the rate of incorporation of newly-fixed C into soil. This rate constrains the size of the possible effect of eievated CO₂ on soil C. Even after four years of treatment, statistical power to detect plausible changes in soil C under elevated CO₂ is quite low. Analysis of other elevated CO₂ experiments in the literature indicates that either CO₂ does not affect soil C content, or that reported CO₂ effects on soil C are too large to be a simple consequence of increased plant carbon inputs, suggesting that other mechanisms are involved, or that the differences are due to chance. Determining the effects of elevated CO₂ on total soil C and long-term C storage requires more powerful experimental techniques or experiments of longer duration.     

Source-sink relations affect growth but not the allocation pattern of birch (Betula pendula Roth) seedlings under elevated [CO2]

ABSTRACT

Birch (Betula pendula Roth.) seedlings were kept for two growing seasons under ambient (～350 µmol mol^-1) and elevated (～700 µmol mol^-1) [CO₂]. The present study was designed to examine the effects of [CO₂] and pot size on growth and carbon allocation under conditions of non-limiting water and nutrient supply, in order to separate the effects of source-sink interaction from the effects of nutrient deficiency. The manipulation of the source-sink relations had a strong influence on the growth response to elevated [CO₂]. When the rooting volume was inadequate, it resulted in a source-sink imbalance which constrained growth under elevated [CO₂]. When root exploration was unconstrained, total dry mass was significantly increased (by about 24%) under elevated [CO₂]. However, the allometric relationships in allocation pattern and in morphogenetic development were not affected by either [CO₂] or pot treatments when the saplings were of the same size. Thus, by constraining dry mass production, small sinks affected the magnitude of the growth responses to elevated [CO₂], but did not affect the plant allocation pattern and allometric relationships when nutrient supply was non-limiting. However, by slowing down growth, sink restrictions counteract the speed-up of ontogeny which is the main effect of elevated [CO₂] on tree growth.     

Responses of soil biota to elevated atmospheric carbon dioxide

Increasing concentrations of atmospheric CO₂ could have dramatic effects upon terrestrial ecosystems including changes in ecosystem structure, nutrient cycling rates, net primary production, C source-sink relationships and successional patterns. All of these potential changes will be constrained to some degree by below ground processes and mediated by responses of soil biota to indirect effects of CO₂ enrichment. A review of our current state of knowledge regarding responses of soil biota is presented, covering responses of mycorrhizae, N-fixing bacteria and actinomycetes, soil microbiota, plant pathogens, and soil fauna. Emphasis will be placed on consequences to biota of increasing C input through the rhizosphere and resulting feedbacks to above ground systems. Rising CO₂ may also result in altered nutrient concentrations of plant litter, potentially changing decomposition rates through indirect effects upon decomposer communities. Thus, this review will also cover current information on decomposition of litter produced at elevated CO₂. Summary Predictably, the responses of soil biota to CO₂ enrichment and the degree of experimental emphasis on them increase with proximity to, and intimacy with, roots. Symbiotic associations are all stimulated to some degree. Total plant mycorrhization increases with elevated CO₂. VAM fungi increase proportionately with fine root length/mass increase. ECM fungi, however, exhibit greater colonization per unit root length/mass at elevated CO₂ than at current atmospheric levels. Total N-fixation per plant increases in all species examined, although the mechanisms of increase, as well as the eventual benefit to the host relative to N uptake may vary. Microbial responses are unclear. The assumption that changes in root exudation will drive increased mineralization and facilitate nutrient uptake should be examined experimentally, in light of recent models. Microbial results to date suggest that metabolic activity (measured as changes in process rates) is stimulated by root C input, rather than population size (measured by cell or colony counts). Insufficient evidence exists to predict responses of either soil-borne plant pathogens or soil fauna (i.e., food web responses). These are areas requiring attention, the first for its potential to limit ecosystem production through disease and the second because of its importance to nutrient cycling processes. Preliminary data on foliar litter decomposition suggests that neither nutrient ratios nor decomposition rates will be affected by rising CO₂. This is another important area that may be better understood as the number of longer term studies with more realistic CO₂ exposures increase. Evidence continues to mount that C fixation increases with CO₂ enrichment and that the bulk of this C enters the belowground component of ecosystems. The global fate and effects of this additional C may affect all hierarchical levels, from organisms to ecosystems, and will be largely determined by responses of soil biota.     

Crop responses to CO2 enrichment

Carbon dioxide is rising in the global atmosphere, and this increase can be expected to continue into the foreseeable future. This compound is an essential input to plant life. Crop function is affected across all scales from biochemical to agro-ecosystem. An array of methods (leaf cuvettes, field chambers, free-air release systems) are available for experimental studies of CO₂ effects. Carbon dioxide enrichment of the air in which crops grow usually stimulates their growth and yield. Plant structure and physiology are markedly altered. Interactions between CO₂ and environmental factors that influence plants are known to occur. Implications for crop growth and yield are enormous. Strategies designed to assure future global food security must include a consideration of crop responses to elevated atmospheric CO₂. Future research should include these targets: search for new insights, development of new techniques, construction of better simulation models, investigation of belowground processes, study of interactions, and the elimination of major discrepancies in the scientific knowledge base.     

Dark septate root endophytic fungi increase growth of Scots pine seedlings under elevated CO2 through enhanced nitrogen use efficiency

Although increasing concentrations of atmospheric CO₂ are predicted to have substantial impacts on plant growth and functioning of ecosystems, there is insufficient understanding of the responses of belowground processes to such increases. We investigated the effects of different dark septate root endophytic (DSE) fungi on growth and nutrient acquisition by Pinus sylvestris seedlings under conditions of N limitation and at ambient and elevated CO₂ (350 or 700 μ1 CO₂ l^?1). Each seedling was inoculated with one of the following species: Phialocephala fortinii (two strains), Cadophora finlandica, Chloridium paucisporum, Scytalidium vaccinii, Meliniomyces variabilis and M. vraolstadiae. The trial lasted 125 days. During the final 27 days, the seedlings were labeled with ¹⁴CO₂ and ¹⁵NH ₄ ⁺ . We measured extraradical hyphal length, internal colonization, plant biomass, ¹⁴C allocation, and plant N and ¹⁵N content. Under elevated CO₂, the biomass of seedlings inoculated with DSE fungi was on average 17% higher than in control seedlings. Simultaneously, below-ground respiration doubled or trebled, and as a consequence carbon use efficiency by the DSE fungi significantly decreased. Shoot N concentration decreased on average by 57% under elevated CO₂ and was lowest in seedlings inoculated with S. vaccinii. Carbon gain by the seedlings despite reduced shoot N concentration indicates that DSE fungi increase plant nutrient use efficiency and are therefore more beneficial to the plant under elevated CO₂.     

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。     

Plant species-specific changes in root-inhabiting fungi in a California annual grassland: responses to elevated CO2 and nutrients

Five co-occurring plant species from an annual mediterranean grassland were grown in monoculture for 4 months in pots inside open-top chambers at the Jasper Ridge Biological Preserve (San Mateo County, California). The plants were exposed to elevated atmospheric CO₂ and soil nutrient enrichment in a complete factorial experiment. The response of root-inhabiting non-mycorrhizal and arbuscular mycorrhizal fungi to the altered resource base depended strongly on the plant species. Elevated CO₂ and fertilization altered the ratio of non-mycorrhizal to mycorrhizal fungal colonization for some plant species, but not for others. Percent root infection by non-mycorrhizal fungi increased by over 500% for Linanthus parviflorus in elevated CO₂, but decreased by over 80% for Bromus hordeaceus. By contrast, the mean percent infection by mycorrhizal fungi increased in response to elevated CO₂ for all species, but significantly only for Avena barbata and B. hordeaceus. Percent infection by mycorrhizal fungi increased, decreased, or remained unchanged for different plant hosts in response to fertilization. There was evidence of a strong interaction between the two treatments for some plant species and non-mycorrhizal and mycorrhizal fungi. This study demonstrated plant species- and soil fertility-dependent shifts in below-ground plant resource allocation to different morpho-groups of fungal symbionts. This may have consequences for plant community responses to elevated CO₂ in this California grassland ecosystem. Received: 2 June 1997 / Accepted: 22 August 1997     

Potential effects of elevated carbon dioxide on leaf-feeding forest insects

The elevated concentration of atmospheric CO₂ may result in a decline of leaf nutritional quality (especially N) and an increase in some kinds of defensive secondary components (such as phenolics). The changes in the phytochemistry of trees, combined with the effect of elevated CO₂ per se, have a potential negative influence on insect herbivores. Here, we review the effect of elevated CO₂ on the performance of leaffeeding forest insects at individual-level and community-level. The elevated CO₂ per se have little influence on the metabolism of insects. Over half of the tree-insect experimental systems show that the performance of individual insect become poorer under high-CO₂ grown trees; but the others show that the insects have just little or no response to the treatments. The direction and magnitude of the changes in the performance of insects could be mediated by various factors. The effects of treatment are strongly species-dependent. The magnitude of changes in the phytochemistry, the sensitivity and adaptive capacity of insects to the poorer leaf quality, the differences in plant growth conditions and experimental methods, and the mediated effects of other environmental factors (such as soil nutrient availability, light, temperature, O₃) were all closely related to the final performance of insects. However, the larvae’s consumption usually increased under enriched CO₂ treatment, which was widely thought to be a compensatory response to poorer plant quality. The experiments on forest community-level found identically a reduction in herbivory, which was contrary to the results from small-scale experiments. The changes in insect population and the actual response of consumption by leaf-feeding forest insects under CO₂ enrichment remain unclear, and more field-based experiments need to be conducted. __________ Translated from Chinese Journal of Applied Ecology, 2006, 17(4): 720–726 [译自: 应用生态学报]     

P uptake by arbuscular mycorrhizal hyphae: effect of soil temperature and atmospheric CO2 enrichment

Mycorrhizas are ubiquitous symbioses that may have an important role in the movement of C from air to soil. Studies on the effects of climate change factors on mycorrhizas have been concentrated on the effects of atmospheric [CO₂] whereas temperature effects have been neglected. Based on previous results showing no effect of varying atmospheric [CO₂] on the development and P uptake of the arbuscular mycorrhizal fungi (AMF) colonizing plants growing in controlled conditions, we hypothesized that soil temperature would have a higher impact on AMF development and nutrient uptake than the effects of [CO₂] on the host plant. Pea plants were grown in association with either a single isolate of Glomus caledonium or AMF from field soil in factorial combination with the corresponding current (10 °C) or elevated (15 °C) soil temperatures at current (350 p.p.m) or elevated (700 p.p.m) atmospheric [CO₂]. ³³P uptake by extraradical AMF hyphae was measured independently from root P uptake in a root exclusion compartment. Intraradical colonization developed well at both soil temperatures and almost duplicated from 10 to 15 °C. Extraradical mycelium developed only at 15 °C in the root exclusion compartment and hyphal P uptake could therefore be studied at 15 °C only. Hyphal P uptake differed markedly between inoculum types, but was not altered by growing the host plants at two atmospheric [CO₂] levels. No significant [CO₂] × soil temperature interactions were observed. The results suggested that, in the system tested, AMF development and function is likely more influenced by the temperature component of climate change than by its [CO₂] component. We suggest that much more attention should be paid to temperature effects in future studies.     

Issues and perspectives for investigating root responses to elevated atmospheric carbon dioxide

A thorough assessment of how plants and ecosystems will respond to increasing concentrations of atmospheric CO₂ requires that the responses of root systems and associated belowground processes be understood. Static measures of root-to-shoot ratio have not been satisfactory for describing the integrated responses of plants to CO₂-enriched atmospheres, but research with a process orientation has suggested that elevated CO₂ can stimulate root growth or root activity and provide a positive feedback on plant growth. There are, however, critical questions concerning the relevance of root data from short-term studies with potted plants when scaling to questions about plants in the field. Data on root responses to CO₂ enrichment in the field are fragmentary, but they allow us to more clearly define research questions for further investigation. Three perspectives for analyzing the significance of root responses as a component of the overall response of the terrestrial biosphere to increasing atmospheric CO₂ are suggested: (1) roots as a platform for nutrient acquisition and a mediator of whole-plant response to CO₂; (2) carbon storage in roots as a component of whole-plant carbon storage; and (3) effects of CO₂ enrichment on root turnover and the implications for carbon storage as soil organic matter. The relative importance of these different perspectives will vary depending on the ecosystem of interest and the larger-scale issues being considered.     

Effect of carbon dioxide concentration on microbial respiration in soil

In order to assess the validity of conventional methods for measuring CO₂ flux from soil, the relationship between soil microbial respiration and ambient CO₂ concentration was studied using an open-flow infra-red gas analyser (IRGA) method. Andosol from an upland field in central Japan was used as a soil sample. Soil microbial respiration activity was depressed with the increase of CO₂ concentration in ventilated air from 0 to 1000 ppmv. At 1000 ppmv, the respiration rate was less than half of that at 0 ppmv. Thus, it is likely that soil respiration rate is overestimated by the alkali absorption method, because CO₂ concentration in the absorption chamber is much lower than the normal level. Metabolic responses to CO₂ concentration were different among groups of soil microorganisms. The bacteria actinomycetes group cultivated on agar medium showed a more sensitive response to the CO₂ concentration than the filamentous fungi group.     

No Detectable Maternal Effects of Elevated CO2 on Arabidopsis thaliana Over 15 Generations

Maternal environment has been demonstrated to produce considerable impact on offspring growth. However, few studies have been carried out to investigate multi-generational maternal effects of elevated CO₂ on plant growth and development. Here we present the first report on the responses of plant reproductive, photosynthetic, and cellular characteristics to elevated CO₂ over 15 generations using Arabidopsis thaliana as a model system. We found that within an individual generation, elevated CO₂ significantly advanced plant flowering, increased photosynthetic rate, increased the size and number of starch grains per chloroplast, reduced stomatal density, stomatal conductance, and transpiration rate, and resulted in a higher reproductive mass. Elevated CO₂ did not significantly influence silique length and number of seeds per silique. Across 15 generations grown at elevated CO₂ concentrations, however, there were no significant differences in these traits. In addition, a reciprocal sowing experiment demonstrated that elevated CO₂ did not produce detectable maternal effects on the offspring after fifteen generations. Taken together, these results suggested that the maternal effects of elevated CO₂ failed to extend to the offspring due to the potential lack of genetic variation for CO₂ responsiveness, and future plants may not evolve specific adaptations to elevated CO₂ concentrations.     

Changes in the microbial community structure of bacteria,archaea and fungi in response to elevated CO2 and warming in an Australian native grassland soil

The microbial community structure of bacteria, archaea and fungi is described in an Australian native grassland soil after more than 5 years exposure to different atmospheric CO₂ concentrations ([CO₂]) (ambient, + 550 ppm) and temperatures (ambient, + 2°C) under different plant functional types (C ₃ and C ₄ grasses) and at two soil depths (0–5 cm and 5–10 cm). Archaeal community diversity was influenced by elevated [CO₂], while under warming archaeal 16S rRNA gene copy numbers increased for C ₄ plant Themeda triandra and decreased for the C ₃ plant community (P < 0.05). Fungal community diversity resulted in three groups based upon elevated [CO₂], elevated [CO₂] plus warming and ambient [CO₂]. Overall bacterial community diversity was influenced primarily by depth. Specific bacterial taxa changed in richness and relative abundance in response to climate change factors when assessed by a high‐resolution 16S rRNA microarray (PhyloChip). Operational taxonomic unit signal intensities increased under elevated [CO₂] for both Firmicutes and Bacteroidetes, and increased under warming for Actinobacteria and Alphaproteobacteria. For the interaction of elevated [CO₂] and warming there were 103 significant operational taxonomic units (P < 0.01) representing 15 phyla and 30 classes. The majority of these operational taxonomic units increased in abundance for elevated [CO₂] plus warming plots, while abundance declined in warmed or elevated [CO₂] plots. Bacterial abundance (16S rRNA gene copy number) was significantly different for the interaction of elevated [CO₂] and depth (P < 0.05) with decreased abundance under elevated [CO₂] at 5–10 cm, and for Firmicutes under elevated [CO₂] (P < 0.05). Bacteria, archaea and fungi in soil responded differently to elevated [CO₂], warming and their interaction. Taxa identified as significantly climate‐responsive could show differing trends in the direction of response (‘+’ or ‘?’) under elevated CO₂ or warming, which could then not be used to predict their interactive effects supporting the need to investigate interactive effects for climate change. The approach of focusing on specific taxonomic groups provides greater potential for understanding complex microbial community changes in ecosystems under climate change.