The response of plants to elevated CO2

Tree saplings, two groups of three species from each of two deciduous tree communities, were grown in competition at three CO₂ concentrations and two light levels. After one growing season, biomass was measured to assess the effect of CO₂ on community structure, and nitrogen and phosphorus concentrations were measured for leaves, stems, and roots of all trees. Gas-exchange measurements were made on the same species grown under the same CO₂ concentrations.Photosynthetic capacity (rate of photosynthesis at saturating CO₂ and light) tended to decline as CO₂ concentration increased, but differences were not statistically significant. Stomatal conductance declined significantly as CO₂ increased. Nitrogen and phosphorus concentrations generally declined as CO₂ increased, but there were some unexpected patterns in roots and stems. CO₂ concentration did not significantly affect the overall growth of either community after one season, but the relative biomass of each species changed in a complex way, depending on CO₂ light level, and community.     

The growth response of plants to elevated CO2 under non-optimal environmental conditions

Under benign environmental conditions, plant growth is generally stimulated by elevated atmospheric CO₂ concentrations. When environmental conditions become sub- or supra-optimal for growth, changes in the biomass enhancement ratio (BER; total plant biomass at elevated CO₂ divided by plant biomass at the current CO₂ level) may occur. We analysed literature sources that studied CO₂2environment interactions on the growth of herbaceous species and tree seedlings during the vegetative phase. For each experiment we calculated the difference in BER for plants that were grown under 'optimal' and 'non-optimal' conditions. Assuming that interactions would be most apparent if the environmental stress strongly diminished growth, we scaled the difference in the BER values by the growth reduction due to the stress factor. In our compilation we found a large variability in CO₂2environment interactions between experiments. To test the impact of experimental design, we simulated a range of analyses with a plant-to-plant variation in size common in experimental plant populations, in combination with a number of replicates generally used in CO₂2environment studies. A similar variation in results was found as in the compilation of real experiments, showing the strong impact of stochasticity. We therefore caution against strong inferences derived from single experiments and suggest rather a reliance on average interactions across a range of experiments. Averaged over the literature data available, low soil nutrient supply or sub-optimal temperatures were found to reduce the proportional growth stimulation of elevated CO₂. In contrast, BER increased when plants were grown at low water supply, albeit relatively modestly. Reduced irradiance or high salinity caused BER to increase in some cases and decrease in others, resulting in an average interaction with elevated CO₂ that was not significant. Under high ozone concentrations, the relative growth enhancement by elevated CO₂ was strongly increased, to the extent that high CO₂ even compensated in an absolute way for the harmful effect of ozone on growth. No systematic difference in response was found between herbaceous and woody species for any of the environmental variables considered.     

Interspecific variation in the growth response of plants to an elevated ambient CO2 concentration

The effect of a doubling in the atmospheric CO₂ concentration on the growth of vegetative whole plants was investigated. In a compilation of literature sources, the growth stimulation of 156 plant species was found to be on average 37%. This enhancement is small compared to what could be expected on the basis of CO₂-response curves of photosynthesis. The causes for this stimulation being so modest were investigated, partly on the basis of an experiment with 10 wild plant species. Both the source-sink relationship and size constraints on growth can cause the growth-stimulating effect to be transient.Data on the 156 plant species were used to explore interspecific variation in the response of plants to high CO₂. The growth stimulation was larger for C₃ species than for C₄ plants. However the difference in growth stimulation is not as large as expected as C₄ plants also significantly increased in weight (41% for C₃ vs. 22% for C₄). The few investigated CAM species were stimulated less in growth (15%) than the average C₄ species. Within the group of C₃ species, herbaceous crop plants responded more strongly than herbaceous wild species (58%vs. 35%) and potentially fast-growing wild species increased more in weight than slow-growing species (54%vs. 23%). C₃ species capable of symbiosis with N₂-fixing organisms had higher growth stimulations compared to other C₃ species. A common denominator in these 3 groups of more responsive C₃ plants might be their large sink strength. Finally, there was some tendency for herbaceous dicots to show a larger response than monocots. Thus, on the basis of this literature compilation, it is concluded that also within the group of C₃ species differences exist in the growth response to high CO₂.Abbreviations LAR leaf area ratio - LWR leaf weight ratio - NAR net assimilation rate - PS_a rate of photosynthesis per unit leaf area - RGR relative growth rate - RWR root weight ratio - SLA specific leaf area     

Soil microbial response in tallgrass prairie to elevated CO2

Terrestrial responses to increasing atmospheric CO₂ are important to the global carbon budget. Increased plant production under elevated CO₂ is expected to increase soil C which may induce N limitations. The objectives of this study were to determine the effects of increased CO₂ on 1) the amount of carbon and nitrogen stored in soil organic matter and microbial biomass and 2) soil microbial activity. A tallgrass prairie ecosystem was exposed to ambient and twice-ambient CO₂ concentrations in open-top chambers in the field from 1989 to 1992 and compared to unchambered ambient CO₂ during the entire growing season. During 1990 and 1991, N fertilizer was included as a treatment. The soil microbial response to CO₂ was measured during 1991 and 1992. Soil organic C and N were not significantly affected by enriched atmospheric CO₂. The response of microbial biomass to CO₂ enrichment was dependent upon soil water conditions. In 1991, a dry year, CO₂ enrichment significantly increased microbial biomass C and N. In 1992, a wet year, microbial biomass C and N were unaffected by the CO₂ treatments. Added N increased microbial C and N under CO₂ enrichment. Microbial activity was consistently greater under CO₂ enrichment because of better soil water conditions. Added N stimulated microbial activity under CO₂ enrichment. Increased microbial N with CO₂ enrichment may indicate plant production could be limited by N availability. The soil system also could compensate for the limited N by increasing the labile pool to support increased plant production with elevated atmospheric CO₂. Longer-term studies are needed to determine how tallgrass prairie will respond to increased C input.     

红松幼苗对CO2浓度升高的生理生态反应

研究了用开顶箱控制CO     

Proteomic response of rice seedling leaves to elevated CO2 levels

Previous investigations of plant responses to higher CO 2 levels were mostly based on physiological measurements and biochemical assays. In this study, a proteomic approach was employed to investigate plant response to higher CO 2 levels using rice as a model. Ten-day-old seedlings were progressively exposed to 760 ppm, 1140 ppm, and 1520 ppm CO 2 concentrations for 24 h each. The net photosynthesis rate ( P n), stomatal conductance ( G s), transpiration rate ( E), and intercellular to ambient CO 2 concentration ratio ( C i/ C a) were measured. P n, G s, and E showed a maximum increase at 1140 ppm CO 2, but further exposure to 1520 ppm for 24 h resulted in down regulation of these. Proteins extracted from leaves were subjected to 2-DE analysis, and 57 spots showing differential expression patterns, as detected by profile analysis, were identified by MALDI-TOF/TOF-MS. Most of the proteins belonged to photosynthesis, carbon metabolism, and energy pathways. Several molecular chaperones and ascorbate peroxidase were also found to respond to higher CO 2 levels. Concomitant with the down regulation of P n and G s, the levels of enzymes of the regeneration phase of the Calvin cycle were decreased. Correlations between the protein profiles and the photosynthetic measurements at the three CO 2 levels were explored.     

Photosynthetic and growth response to fumigation with SO2 at elevated CO2 for C3 and C4 plants

Summary Six early successional plant species with differing photosynthetic pathways (3 C₃ species and 3 C₄ species) were grown at either 300, 600, or 1,200 ppm CO₂ and at either 0.0 or 0.25 ppm SO₂. Total plant growth increased with CO₂ concentration for the C₃ species and varied only slightly with CO₂ for the C₄ species. Fumigation with SO₂ caused reduced growth of the C₃ species at 300 ppm CO₂ but not at the higher concentrations of CO₂. Fumigation with SO₂ reduced growth of the C₄ species at high CO₂ and increased growth at 300 ppm CO₂. Leaf area increased with increasing CO₂ for all plant species. Fumigation with SO₂ reduced leaf area of C₃ plants more at low CO₂ than at high CO₂ while leaf area of C₄ plants was reduced more at high CO₂ than at low CO₂. These results support the notion that C₃ species are more sensitive to SO₂ fumigation than are C₄ species at concentrations of CO₂ equal to that found in normal ambient air. However, the difference in sensitivity to SO₂ between C₃ and C₄ species was found to be reversed at higher concentrations of CO₂. A possible explanation for this reversal based upon differences in stomatal response to elevated CO₂ between C₃ and C₄ species is discussed.     

Rhizosphere microflora of winter wheat plants cultivated under elevated CO2

We studied an effect of elevated atmospheric CO₂ on rhizosphere microorganisms in a hydroponics system where young wheat plants provided the only source of C for microorganisms. Plants were cultivated in mineral solution in sterile silica sand and exposed to control (ambient) and elevated (double) CO₂ concentrations for periods of 13, 20, 25 and 34 days.Microbial biomass C (C content in fraction of size 0.3–2.7 µm) was not affected by the elevated CO₂ concentration during the first 25 days of plant growth and was increased after 34 days of plant growth. A content of poly--hydroxybutyrate (PHB) reserve compounds (measured as derivatized product of 3-hydroxy-butyric acid and N-tert-butyldimethylsilyl-N-methyltrifluoracetamide using GC–MS) was lowered significantly (p<0.001) in the elevated CO₂ after 25 and 34 days. It was accompanied with a shift of bacterial distribution towards the nutritional groups utilising more complex organic material (number of CFUs on media with different sources of C and N). A coincidence of several events connected with plant and microbial carbon economy (decrease of an assimilation rate and relative growth rate of plants, small increase of microbial biomass, PHB decrease and suppression within the bacterial nutritional group requiring the most readily available source of C and energy) was observed in the system under elevated CO₂ on the 25th day.A modification of the GC–MS method for the detection of low levels of PHB compounds in natural samples was developed. We excluded the lipids fractionation step and we used EI MS/MS detection of the main fragment ions of the derivatized compound. This guarantees that the ion profiles have high signal-to-noise ratio at correct retention time. The detection limit is then about 30 pg g^-1 of sand or soil.The rhizosphere microflora responded very sensitively to the short-term changes in C partitioning in plants caused by the elevated CO₂.     

The fraction of expanding to expanded leaves determines the biomass response of Populus to elevated CO2

We examined whether the effects of elevated CO₂ on growth of 1-year old Populus deltoides saplings was a function of the assimilation responses of particular leaf developmental stages. Saplings were grown for 100 days at ambient (approximately 350 ppm) and elevated (ambient + 200 ppm) CO₂ in forced-air greenhouses. Biomass, biomass distribution, growth rates, and leaf initiation and expansion rates were unaffected by elevated CO₂. Leaf nitrogen (N), the leaf C:N ratio, and leaf lignin concentrations were also unaffected. Carbon gain was significantly greater in expanding leaves of saplings grown at elevated compared to ambient CO₂. The Rubisco content in expanding leaves was not affected by CO₂ concentration. Carbon gain and Rubisco content were significantly lower in fully expanded leaves of saplings grown at elevated compared to ambient CO₂, indicating CO₂-induced down-regulation in fully expanded leaves. Elevated CO₂ likely had no overall effect on biomass accumulation due to the more rapid decline in carbon gain as leaves matured in saplings grown at elevated compared to ambient CO₂. This decline in carbon gain has been documented in other species and shown to be related to a balance between sink/source balance and acclimation. Our data suggest that variation in growth responses to elevated CO₂ can result from differences in leaf assimilation responses in expanding versus expanded leaves as they develop under elevated CO₂. Received: 28 September 1998 / Accepted: 23 June 1999     

Alfalfa response to elevated atmospheric CO2 varies with the symbiotic rhizobial strain

Increased atmospheric CO₂ was shown to affect a variety of physiological processes in plants, including photosynthesis and growth with repercussions on crop yield and nutritive value. Perennial alfalfa (Medicago sativa L.) is a sustainable crop with a deep root system, living in symbiosis with rhizobium for nitrogen (N) fixation. The objective of the project was to determine the combined effects of elevated CO₂ and rhizobial strains on photosynthesis, growth, N fixation, and nutritive value of alfalfa, and on soil microflora. Alfalfa inoculated with two different strains of rhizobia (Sinorhizobium meliloti strains A2 and NRG34) was grown 2 months at day/night temperatures of 22/17°C under either 400 (near ambient) or 800 (elevated) μmol mol⁻¹ of CO₂. The photosynthetic response of alfalfa to elevated CO₂ differed according to the rhizobial strain. At the end of the experiment, elevated CO₂ stimulated photosynthetic rates by 50% in plants associated with A2 but there was no significant increase in plants nodulated with NRG34. Nitrogenase activity (+38%) and shoot growth (+60%) were stimulated under 800 μmol mol⁻¹ of CO₂ for alfalfa inoculated with both strains. Root dry weight was significantly higher at 800 μmol mol⁻¹ of CO₂ only with strain A2. Fibre concentration decreased in response to elevated atmospheric CO₂ in alfalfa inoculated with strain A2 resulting in plant material with greater nutritive value when inoculated with A2 compared to NRG34. In the soil, elevated CO₂ increased the proportion of fungi in the microbial community while decreasing Gram⁻ bacteria. For alfalfa inoculated with rhizobial strain A2, photosynthetic rates, nitrogenase activity, and growth were all stimulated by increased atmospheric CO₂ compared to less consistently positive responses to elevated CO₂ when inoculated with NRG34. Our results show that it is possible to identify rhizobial strains to improve plant performance under predicted future CO₂ concentrations with no negative effect on nutritive value. The Canadian Government’s right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

我国主要类型昆虫对CO_2升高响应的研究进展

大气CO2浓度增加已经受到国内外的极大关注。自2002年以来,在自行设计、组装的一系列密闭式动态CO2气室和开顶式CO2浓度控制箱基础上,研究了我国主要类型昆虫对CO2浓度升高响应的特征。结果显示,大气CO2浓度升高降低了棉铃虫Helicoverpa armigera的适合度和对棉花的危害作用,增加了棉花对棉铃虫为害的补偿作用,使以咀嚼式口器昆虫为代表的棉铃虫种群发生与危害下降;但大气CO2浓度升高改变了植物组织营养物质的组成与含量,提高了蚜虫Aphis gosypii对氨基酸营养的利用与补偿效率,降低了3种麦蚜的种间竞争,导致蚜虫种群发生与危害严重;而对烟粉虱Bemisia tabaci的种群特征影响较少。大气CO2浓度升高对天敌昆虫的影响存在种的特异性,表现出种群上升、下降和变化不大等特征。未来大气CO2浓度升高下,由于作物生长发育加快,生物量增加,蚜虫种群增多,导致吡虫啉农药防治效果下降,由此未来大气CO2浓度升高下农民将被迫使用更多的化学农药防治蚜虫类害虫,进而加重环境污染。     

Progressive N limitation of plant response to elevated CO2: a microbiological perspective

A major uncertainty in predicting long-term ecosystem C balance is whether stimulation of net primary production will be sustained in future atmospheric CO₂ scenarios. Immobilization of nutrients (N in particular) in plant biomass and soil organic matter (SOM) provides negative feedbacks to plant growth and may lead to progressive N limitation (PNL) of plant response to CO₂ enrichment. Soil microbes mediate N availability to plants by controlling litter decomposition and N transformations as well as dominating biological N fixation. CO₂-induced changes in C inputs, plant nutrient demand and water use efficiency often have interactive and contrasting effects on microbes and microbially mediated N processes. One critical question is whether CO₂-induced N accumulation in plant biomass and SOM will result in N limitation of microbes and subsequently cause them to obtain N from alternative sources or to alter the ecosystem N balance. We reviewed the experimental results that examined elevated CO₂ effects on microbial parameters, focusing on those published since 2000. These results in general show that increased C inputs dominate the CO₂ impact on microbes, microbial activities and their subsequent controls over ecosystem N dynamics, potentially enhancing microbial N acquisition and ecosystem N retention. We reason that microbial mediation of N availability for plants under future CO₂ scenarios will strongly depend on the initial ecosystem N status, and the nature and magnitude of external N inputs. Consequently, microbial processes that exert critical controls over long-term N availability for plants would be ecosystem-specific. The challenge remains to quantify CO₂-induced changes in these processes, and to extrapolate the results from short-term studies with step-up CO₂ increases to native ecosystems that are already experiencing gradual changes in the CO₂ concentration.     

Arbuscular mycorrhizae respond to plants exposed to elevated atmospheric CO2 as a function of soil depth

The importance of arbuscular mycorrhizae (AM) in plant and ecosystem responses to global changes, e.g. elevated atmospheric CO₂, is widely acknowledged. Frequently, increases in AM root colonization occur in response to increased CO₂, but also the lack of significant changes has been reported. The goal of this study was to test whether arbuscular mycorrhizae (root colonization and composition of root colonization) respond to plants grown in elevated CO₂ as a function of soil depth. We grew Bromus hordeaceus L. and Lotus wrangelianus Fischer & C. Meyer monocultures in large pots with a synthetic serpentine soil profile for 4 yr in an experiment, in which CO₂ concentration was crossed factorially with NPK fertilization. When analyzing root infection separately for topsoil (0–15 cm) and subsoil (15–45 cm), we found large (e.g., about 5-fold) increases of AM fungal root colonization in the subsoil in response to CO₂, but no significant changes in the corresponding topsoil of Bromus. Only the coarse endophyte AM fungi, not the fine endophyte AM fungi, were responsible for the observed increase in the bottom soil layer, indicating a depth-dependent shift in the AM community colonizing the roots, even at this coarse morphological level. Other response variables also had significant soil layer ^* CO₂ interaction terms. The subsoil response would have been hidden in an unstratified assessment of the total root system, since most of the root length was concentrated in the top soil layer. The increased presence of mycorrhizae in roots deeper in the soil should be considered in sampling protocols, as it may be indicative of changed patterns of nutrient acquisition and carbon sequestration.     

The mangrove ant, Camponotus anderseni, switches to anaerobic respiration in response to elevated CO2 levels

The small tree-living mangrove ant Camponotus anderseni is remarkably adapted for surviving tidal inundation. By blocking the nest entrance with a soldier's head, water intrusion into the nest cavity can be effectively prevented, but lack of gas-exchange caused extremely high concentrations of CO(2)(>30%) and very low O(2) concentrations (<1%). The O(2) uptake in experiments with CO(2) absorption showed a linear decrease until about 4%, whereas the O(2) uptake in chambers without absorbent showed a decrease with a different pattern, consisting of three parts. The first component of this decrease is a linear decrease to about 18%, which is the normal O(2) concentration in open natural nests. The second phase is an exponential decrease continuing to about 4% O(2), showing that the CO(2) concentrations have influence on the O(2) uptake. The final component is also exponential, but with a much smaller slope. The respiratory quotient (RQ) was 0.92 until CO(2) concentration increased to about 15-17%, and after that it showed a strong increase, which is due to the initiation of anaerobic respiration. Anaerobic respiration has not been demonstrated for social insects before, but it is not surprising that it is found in this ant species, which lives in the extreme conditions of a hollow twig in an inundated mangrove.     

Nitrogen deposition limits photosynthetic response to elevated CO2 differentially in a dioecious species

Sexual dimorphisms of dioecious plants are important in controlling and maintaining sex ratios under changing climate environments. Yet, little is known about sex-specific responses to elevated CO₂ with soil nitrogen (N) deposition. To investigate sex-related physiological and biochemical responses to elevated CO₂ with N deposition, Populus cathayana Rehd. was employed as a model species. The cuttings were subjected to two CO₂ regimes (350 and 700???mol?mol^?1) with two N levels (0 and 5?g?N?m^?2?year^?1). Our results showed that elevated CO₂ and N deposition separately increased the total number of leaves, leaf area (LA), leaf mass, net photosynthetic rate (P _n), light saturated photosynthetic rate (P _max), chlorophyll a (Chl a), and chlorophyll a to chlorophyll b ratio (Chl a/b) in both males and females of P. cathayana. However, the effects on LA, leaf mass, P _n, P _max, Chl a and Chl a/b were weakened under the combined treatment of elevated CO₂ and N deposition. Males had higher leaf mass, P _n, P _max, apparent quantum yield (??), carboxylation efficiency (CE), Chl a, Chl a/b, leaf N, and root carbon to N ratio (C/N) than did females under elevated CO₂ with N deposition. In contrast to males, females had significantly higher levels of soluble sugars in leaves and greater starch accumulation in roots and stems under the same condition. The results of the present work imply that P. cathayana females are more responsive and suffer from greater negative effects on growth and photosynthetic capacity than do males when grown under elevated CO₂ with soil N deposition.     

Genetic variation in plant below-ground response to elevated CO2 and two herbivore species

Aims

It is unclear how changing atmospheric conditions, including rising carbon dioxide concentration, influence interactions between above and below-ground systems and if intraspecific variation exists in this response.

Methods

We assessed interactive effects of atmospheric CO₂ concentration, above-ground herbivory, and plant genotype on root traits and mycorrhizal associations. Plants from five families of Asclepias syriaca, a perennial forb, were grown under ambient and elevated atmospheric CO₂ concentrations. Foliar herbivory by either lepidopteran caterpillars or phloem-feeding aphids was imposed. Mycorrhizal colonization, below-ground biomass, root biomass, and secondary defensive chemistry in roots were quantified.

Results

We observed substantial genetic variation among A. syriaca families in their mycorrhizal colonization levels in response to elevated CO₂ and herbivory treatments. Elevated CO₂ treatment increased root biomass in all genetic families, whereas foliar herbivory tended to decrease root biomass. Root cardenolide concentration and composition varied greatly among plant families, and elevated CO₂ treatment increased root cardenolides in two of the five plant families. Moreover, herbivores differentially affected the composition of cardenolides expressed below ground.

Conclusions

Increased atmospheric CO₂ has the potential to influence interactions among plants, herbivores and mycorrhizal fungi and intraspecific variation suggests that such interactions can evolve.     

Transgenerational effects of global environmental change: long-term CO2 and nitrogen treatments influence offspring growth response to elevated CO2

Global environmental changes can have immediate impacts on plant growth, physiology, and phenology. Long-term effects that are only observable after one or more generations are also likely to occur. These transgenerational effects can result either from maternal environmental effects or from evolutionary responses to novel selection pressures and are important because they may alter the ultimate ecological impact of the environmental change. Here, we show that transgenerational effects of atmospheric carbon dioxide (CO₂) and soil nitrogen (N) treatments influence the magnitude of plant growth responses to elevated CO₂ (eCO₂). We collected seeds from Lupinus perennis, Poa pratensis, and Schizachyrium scoparium populations that had experienced five growing seasons of ambient CO₂ (aCO₂) or eCO₂ treatments and ambient or increased N deposition and planted these seeds into aCO₂ or eCO₂ environments. We found that the offspring eCO₂ treatments stimulated immediate increases in L. perennis and P. pratensis growth and that the maternal CO₂ environment influenced the magnitude of this growth response for L. perennis: biomass responses of offspring from the eCO₂ maternal treatments were only 54% that of the offspring from the aCO₂ maternal treatments. Similar trends were observed for P. pratensis and S. scoparium. We detected some evidence that long-term N treatments also altered growth responses to eCO₂; offspring reared from seed from maternal N-addition treatments tended to show greater positive growth responses to eCO₂ than offspring from ambient N maternal treatments. However, the effects of long-term N treatments on offspring survival showed the opposite pattern. Combined, our results suggest that transgenerational effects of eCO₂ and N-addition may influence the growth stimulation effects of eCO₂, potentially altering the long-term impacts of eCO₂ on plant populations.