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.     

Increased root growth in elevated CO2: a biophysical analysis of root cell elongation

A biophysical analysis of root expansion was conducted in fourchalk downland herbs (Sanguisorba minor Scop., Lotus corniculatusL., Anthyllis vul-neraria L. and Plantago media L.) exposedto either ambient or elevated CO₂in controlled environment cabinets.Measurements of fine (F) and extra-fine (EF) root extensionrate (RER), water relations, and cell wall tensiometric extensibilityrevealed differences in the diurnal pattern of root growth betweenspecies. After 35 d of exposure to elevated CO₂, RER of bothF and EF roots increased significantly in darkness and on illuminationfor S. minor, whilst for A. vulneraria (EF roots only) and L.corniculatus a significant increase occurred at night whereasfor P. media a significant increase occurred during the day.Cells measured in the zone of elongation were longer in allspecies exposed to elevated CO₂. Water potential (     

The relationship between transpiration and nutrient uptake in wheat changes under elevated atmospheric CO2

The impact of elevated [CO₂] (e[CO₂]) on crops often includes a decrease in their nutrient concentrations where reduced transpiration‐driven mass flow of nutrients has been suggested to play a role. We used two independent approaches, a free‐air CO₂ enrichment (FACE) experiment in the South Eastern wheat belt of Australia and a simulation study employing the agricultural production systems simulator (APSIM), to show that transpiration (mm) and nutrient uptake (g m^?2) of nitrogen (N), potassium (K), sulfur (S), calcium (Ca), magnesium (Mg) and manganese (Mn) in wheat are correlated under e[CO₂], but that nutrient uptake per unit water transpired is higher under e[CO₂] than under ambient [CO₂] (a[CO₂]). This result suggests that transpiration‐driven mass flow of nutrients contributes to decreases in nutrient concentrations under e[CO₂], but cannot solely explain the overall decline.     

Response of Norway spruce root system to elevated atmospheric CO2 concentration

Root structure parameters, root biomass and allometric relationships between above- and belowground biomass were investigated in young Norway spruce (Picea abies [L.] Karst.) trees cultivated inside the glass domes with ambient (AC, 375 μmol(CO₂) mol^?1) and elevated (EC, A + 375 μmol(CO₂) mol^?1) atmospheric CO₂ concentrations ([CO₂]). After 8 years of fumigation, a mean EC tree in comparison with AC one exhibited about 37 % higher belowground biomass. The growth of primary root structure was unaffected by elevated [CO₂]; however, the biomass of secondary roots growing on the primary root structure and the biomass of secondary roots growing in the zone between the soil surface and the first primary root ramification were significantly higher in EC comparing with AC treatment about 58 and 70 %, respectively. The finest root’s (diameter up to 1 mm) biomass as well as length and surface area of both primary and secondary root structures showed the highest difference between the treatments; advancing EC to AC by 43 % on average. Therefore, Norway spruce trees cultivated under well-watered and rather nitrogen-poor soil conditions responded to the air elevated [CO₂] environment by the enhancement of the secondary root structure increment, by enlargement of root length and root absorbing area, and also by alternation of root to aboveground organ biomass proportion. Higher root to leaf and root to stem basal area ratios could be beneficial for Norway spruce trees to survive periods with limited soil water availability.     

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.     

大气CO2浓度升高对植物根系的影响

植物长期生长在CO2浓度不断升高的环境中,其结构和功能都将受到影响,这种影响不仅表现在植物的地上部分,同时也表现在植物的地下部分(根系),尤其是细根的长度、直径、产量、周转以及根与枝的分配模式等方面。植物根系结构和功能的改变影响植物地上部分和生态系统物质循环中的碳动态及土壤中碳库的变化。目前有关大气CO2浓度升高对根系动态影响的研究报道主要包括大气CO2浓度升高对根系结构(直径、分枝、长度、数量等)和根系生理(周转率、产量、碳分配模式等)的影响2个方面。目前,该领域研究还存在一些不足,例如在CO2浓度升高条件下,对植物根系内部的调控机制,以及由其引起的物质循环和能量流动的动态变化的了解较少;至今没有令人信服的证据说明大气CO2浓度升高使根系周转升高还是降低。今后应加强研究在CO2浓度升高条件下根系的周转变化和光合产物分配模式变化,CO2浓度升高和外界环境因素的共同作用对根系的影响,以及采用不同研究方法和研究对象在不同立地条件下开展升高CO2浓度对根系影响的对比研究等。     

氮素对高大气CO2浓度下小麦叶片光合作用的影响

通过测定小麦拔节期叶片的光合气体交换参数和光强-光合速率（P_n）响应曲线,研究了氮素对长期高大气CO₂浓度（760 μmol·mol^-1）下小麦叶片光合作用的影响.结果表明：在长期高大气CO₂浓度下,增施氮肥能提高小麦叶片P_n、蒸腾速率（T_r）和瞬时水分利用效率（WUE_i）;与正常大气CO₂浓度相比,高大气CO₂浓度下小麦叶片的P_n和WUE_i增加,气孔导度（G_s）和胞间CO₂浓度(C_i）降低.随光合有效辐射的增强,高大气CO₂浓度下小麦叶片的P_n和WUE_i均高于正常大气CO₂浓度处理,G_s则较低,而C_i和T_r无显著变化.高氮水平下小麦叶片G_s与P_n、T_r、WUE_i呈线性正相关,G_s与C_i在正常大气CO₂浓度下呈线性负相关,但高大气CO₂浓度下二者无相关性;低氮水平下小麦叶片的G_s与P_n、WUE_i无相关性,而与C_i和T_r呈线性正相关,表明高大气CO₂浓度下低氮水平的小麦叶片P_n由非气孔因素限制.     

Responses of root hair development to elevated CO2

This review highlights a potential signaling pathway of CO₂-dependent stimulation in root hair development. Elevated CO₂ firstly increases the carbohydrates production, which triggers the auxin or ethylene responsive signal transduction pathways and subsequently stimulates the generation of intracellular nitric oxide (NO). The NO acts on target Ca²⁺ and ion channels and induces activation of MAPK. Meanwhile, reactive oxygen species (ROS) activates cytoplasmic Ca²⁺ channels at the plasma membrane in the apex of the root tip. This complex pathway involves transduction cascades of multiple signals that lead to the fine tuning of epidermal cell initiation and elongation. The results suggest that elevated CO₂ plays an important role in cell differentiation processes at the root epidermis.Key words: elevated CO₂, root hairs, carbohydrate, auxin, ethylene, NO, ROS, Ca²⁺, genetic elementsIncreasing concentration of atmospheric CO₂ in the 21st century will impact many aspects of the human and natural world. Elevated CO₂ has some beneficial physiological effects on plants but nutrient limitation has generally been found to suppress these beneficial effects.¹ Therefore, under conditions of suboptimal supply of nutrients and elevated CO₂, the plants need to develop adaptive mechanisms to enhance nutrient acquisition, among which the plasticity of root development is of crucial importance.Root hairs make a significant contribution to increasing root surface area and facilitating physical anchorage to a substrate and providing a large interface for nutrient uptake.² Root-hair cells are highly polarized cellular structures resulting from tip growth of specific epidermal cells, which are controlled by multiple cellular factors and genetic processes.^3,4 Previous studies have shown that root hair development can influenced by various environmental factors, such as nutritional status,⁵ mycorrhizal infection and water stress,⁶ salinity⁷ and light intensity.⁸ Our current research has demonstrated a profound effect of elevated CO₂ on development of root hairs in Arabidopsis, which works through the well-characterized auxin signal transduction pathway.⁹ Since root hairs are an efficient strategy to alleviate the limitation of nutrients, one promising area of future research will be to discover the pathway that control root hair differentiation in crops under elevated CO₂. In this paper, we discussed a layer pathway in the interaction between CO₂ and some classical signals on regulating gene regulatory network to control development of root hairs.     

Benefits of increasing transpiration efficiency in wheat under elevated CO2 for rainfed regions

Higher transpiration efficiency (TE) has been proposed as a mechanism to increase crop yields in dry environments where water availability usually limits yield. The application of a coupled radiation and TE simulation model shows wheat yield advantage of a high‐TE cultivar (cv. Drysdale) over its almost identical low‐TE parent line (Hartog), from about ?7 to 558 kg/ha (mean 187 kg/ha) over the rainfed cropping region in Australia (221–1,351 mm annual rainfall), under the present‐day climate. The smallest absolute yield response occurred in the more extreme drier and wetter areas of the wheat belt. However, under elevated CO₂ conditions, the response of Drysdale was much greater overall, ranging from 51 to 886 kg/ha (mean 284 kg/ha) with the greatest response in the higher rainfall areas. Changes in simulated TE under elevated CO₂ conditions are seen across Australia with notable increased areas of higher TE under a drier climate in Western Australia, Queensland and parts of New South Wales and Victoria. This improved efficiency is subtly deceptive, with highest yields not necessarily directly correlated with highest TE. Nevertheless, the advantage of Drysdale over Hartog is clear with the benefit of the trait advantage attributed to TE ranging from 102% to 118% (mean 109%). The potential annual cost‐benefits of this increased genetic TE trait across the wheat growing areas of Australia (5 year average of area planted to wheat) totaled AUD 631 MIL (5‐year average wheat price of AUD/260 t) with an average of 187 kg/ha under the present climate. The benefit to an individual farmer will depend on location but elevated CO₂ raises this nation‐wide benefit to AUD 796 MIL in a 2°C warmer climate, slightly lower (AUD 715 MIL) if rainfall is also reduced by 20%.     

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.     

Photosynthetic acclimation to elevated CO2 in wheat cultivars

Wheat (T. aestivum) cvs. Kalyansona and Kundan grown under atmospheric (CA) and elevated CO₂ concentrations (650±50 cm³ m^-3 - CE) in open top chambers were examined for net photosynthetic rate (PN), stomatal limitation (l _s) of P _N, ribulose-1,5-bisphosphate carboxylase (RuBPC) activity, and saccharide content of the leaves. The P _N values of both CA- and CE-grown plants compared at the same CO₂ concentration showed a down regulation under CE at the post-anthesis stage. The negative acclimation of P _N appeared to be due to both stomatal and mesophyll components, and the RuBPC activity got also adjusted. There was a decrease in activation state of RuBPC under CE. In connection with this, an increased accumulation of saccharides in wheat leaf under CE was observed. Kalyansona, owing to its larger sink potential in terms of the number of grains, showed a greater enhancement under CE in both post-ear emergence dry matter production and grain yield. Under CE, this cultivar also showed a lower down regulation of P _N than Kundan. This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthetic acclimation to elevated CO2 in wheat cultivars

Wheat (T. aestivum) cvs. Kalyansona and Kundan grown under atmospheric (CA) and elevated CO₂ concentrations (650±50 cm³ m^-3 - CE) in open top chambers were examined for net photosynthetic rate (PN), stomatal limitation (l _s) of P _N, ribulose-1,5-bisphosphate carboxylase (RuBPC) activity, and saccharide content of the leaves. The P _N values of both CA- and CE-grown plants compared at the same CO₂ concentration showed a down regulation under CE at the post-anthesis stage. The negative acclimation of P _N appeared to be due to both stomatal and mesophyll components, and the RuBPC activity got also adjusted. There was a decrease in activation state of RuBPC under CE. In connection with this, an increased accumulation of saccharides in wheat leaf under CE was observed. Kalyansona, owing to its larger sink potential in terms of the number of grains, showed a greater enhancement under CE in both post-ear emergence dry matter production and grain yield. Under CE, this cultivar also showed a lower down regulation of P _N than Kundan.     

大气CO2浓度升高对干旱条件下冬小麦叶片光合适应的影响

大气CO2浓度升高是全球气候变化的主要特征,但大气CO2浓度长期升高条件下冬小麦叶片发生光合适应的机制尚不十分清楚.本研究以盆栽冬小麦'郑麦9023,为试验材料,在人工气候控制室内设置2个CO2浓度(400和600 μmol·mol-1)、2个水分条件(田间持水量的80％±5％和55％±5％),测定拔节期和抽穗期的光合...     

CO2浓度升高和施氮条件下小麦根际呼吸对土壤呼吸的贡献

依托FACE技术平台,采用稳定13C同位素技术,通过将小麦(C3作物)种植于长期单作玉米(C4作物)的土壤上,研究了大气CO2浓度升高和不同氮肥水平对土壤排放CO2的δ13C值及根际呼吸的影响.结果表明:种植小麦后土壤排放CO2的δ13C值随作物生长逐渐降低,CO2浓度升高200 μmol·mol-1显著降低了孕穗、抽穗期(施氮量为250 kg·hm-2,HN)与拔节、孕穗期(施氮量为150 kg·hm-2,LN)土壤排放CO2的δ13C值,显著提高了孕穗、抽穗期的根际呼吸比例.拔节至成熟期,根际呼吸占土壤呼吸的比例在高CO2浓度下为24％～48％ (HN)和21％ ～48％ (LN),在正常CO2浓度下为20％ ～36％ (HN)和19％～32％(LN).不同CO2浓度下土壤排放CO2的δ13C值和根际呼吸对氮肥增加的响应不同,CO2浓度与氮肥用量在拔节期对根际呼吸的交互效应显著.     

Rhizodeposition under ambient and elevated CO2 levels

As global CO₂ levels rise, can soils store more carbon and so buffer atmospheric CO₂ levels? Answering this question requires a knowledge of the rates of C inputs to soil and of CO₂ outputs via decomposition. Below-ground inputs from roots are a major component of the C flow into soils but are still poorly understood. In this article, new techniques for measuring rhizodeposition are reviewed and discussed and the need for cross-comparisons between methods is identified. One component of rhizodeposition, root exudation, is examined in more detail and evidence is presented which suggests that current estimates of exudate flow into soils are incorrect. A mechanistic mathematical model is used to explore how exudate flows might change under elevated CO₂.     

Forest carbon balance under elevated CO2

Free-air CO₂ enrichment (FACE) technology was used to expose a loblolly pine (Pinus taeda L.) forest to elevated atmospheric CO₂ (ambient + 200 µl l^-1). After 4 years, basal area of pine trees was 9.2% larger in elevated than in ambient CO₂ plots. During the first 3 years the growth rate of pine was stimulated by ~26%. In the fourth year this stimulation declined to 23%. The average net ecosystem production (NEP) in the ambient plots was 428 gC m^-2 year^-1, indicating that the forest was a net sink for atmospheric CO₂. Elevated atmospheric CO₂ stimulated NEP by 41%. This increase was primarily an increase in plant biomass increment (57%), and secondarily increased accumulation of carbon in the forest floor (35%) and fine root increment (8%). Net primary production (NPP) was stimulated by 27%, driven primarily by increases in the growth rate of the pines. Total heterotrophic respiration (R_h) increased by 165%, but total autotrophic respiration (R_a) was unaffected. Gross primary production was increased by 18%. The largest uncertainties in the carbon budget remain in separating belowground heterotrophic (soil microbes) and autotrophic (root) respiration. If applied to temperate forests globally, the increase in NEP that we measured would fix less than 10% of the anthropogenic CO₂ projected to be released into the atmosphere in the year 2050. This may represent an upper limit because rising global temperatures, land disturbance, and heterotrophic decomposition of woody tissues will ultimately cause an increased flux of carbon back to the atmosphere.     

Temperature dependence of growth,development, and photosynthesis in maize under elevated CO2

Global atmospheric carbon dioxide concentrations (C_a) are rising. As a consequence, recent climate models have projected that global surface air temperature may increase 1.4–5.8 °C with the doubling of C_a by the end of the century. Because, changes in C_a and temperature are likely to occur concomitantly, it is important to evaluate how the temperature dependence of key physiological processes are affected by rising C_a in major crop plants including maize (Zea mays L.), a globally important grain crop with C₄ photosynthetic pathway. We investigated the temperature responses of photosynthesis, growth, and development of maize plants grown at five temperature regimes ranging from 19/13 to 38.5/32.5 °C under current (370 μmol mol⁻¹) and doubled (750 μmol mol⁻¹) C_a throughout the vegetative stages using sunlit controlled environmental chambers in order to test if the temperature dependence of these processes was altered by elevated C_a. Leaf and canopy photosynthetic rates, C₄ enzyme activities, leaf appearance rates, above ground biomass accumulation and leaf area were measured. We then applied temperature response functions (e.g., Arrhenius and Beta distribution models) to fit the measured data in order to provide parameter estimates of the temperature dependence for modeling photosynthesis and development at current and elevated C_a in maize. Biomass, leaf area, leaf appearance rate, and photosynthesis measured at growth C_a was not changed in response to CO₂ enrichment. Carboxylation efficiency and the activities of C₄ enzymes were reduced with CO₂ enrichment indicating possible photosynthetic acclimation of the C₄ cycle. All measured parameters responded to growth temperatures. Leaf appearance rate and leaf photosynthesis showed curvilinear response with optimal temperatures near 32 and 34 °C, respectively. Total above ground biomass and leaf area were negatively correlated with growth temperature. The dependence of leaf appearance rate, biomass, leaf area, leaf and canopy photosynthesis, and C₄ enzyme activities on growth temperatures was comparable between current and elevated C_a. The results of this study suggest that the temperature effects on growth, development, and photosynthesis may remain unchanged in elevated C_a compared with current C_a in maize.     

Tree species diversity interacts with elevated CO2 to induce a greater root system response

As a consequence of land‐use change and the burning of fossil fuels, atmospheric concentrations of CO₂ are increasing and altering the dynamics of the carbon cycle in forest ecosystems. In a number of studies using single tree species, fine root biomass has been shown to be strongly increased by elevated CO₂. However, natural forests are often intimate mixtures of a number of co‐occurring species. To investigate the interaction between tree mixture and elevated CO₂, Alnus glutinosa, Betula pendula and Fagus sylvatica were planted in areas of single species and a three species polyculture in a free‐air CO₂ enrichment study (BangorFACE). The trees were exposed to ambient or elevated CO₂ (580 μmol mol^?1) for 4 years. Fine and coarse root biomass, together with fine root turnover and fine root morphological characteristics were measured. Fine root biomass and morphology responded differentially to the elevated CO₂ at different soil depths in the three species when grown in monocultures. In polyculture, a greater response to elevated CO₂ was observed in coarse roots to a depth of 20 cm, and fine root area index to a depth of 30 cm. Total fine root biomass was positively affected by elevated CO₂ at the end of the experiment, but not by species diversity. Our data suggest that existing biogeochemical cycling models parameterized with data from species grown in monoculture may be underestimating the belowground response to global change.     

CO2浓度升高对不同品种水稻镉吸收和根形态的影响

为了探讨CO2浓度升高下不同水稻品种荣优398 (RY)和粤杂889(YZ)吸收重金属Cd差异性的原因,利用水培试验研究了不同浓度Cd处理下两种水稻吸收Cd的差异及根形态的变化特征.结果表明:低Cd处理(5、10、20 μmol·L-1)显著增加水稻生物量;当Cd浓度高于50 μmol·L-1时,Cd胁迫效果开始显现,水稻生物量减少.CO2浓度升高显著增加了水稻的生物量,增加了YZ茎Cd含量而降低了RY茎Cd含量.在5～200 μmol·L-1的Cd浓度下,CO2浓度升高增加了YZ活性根在总根长中的比例,降低了RY活性根的比例.CO2浓度升高下不同水稻品种根形态的变化是导致其对Cd吸收差异性的原因之一.