大气CO_2浓度升高对植物影响的研究进展

CO2是植物光合作用的底物,也是构成植物生境的一种环境因子。随着现代工农业的发展与人类活动的增加,大气中CO2浓度将会不可避免的呈升高趋势。这种趋势将对植物生长发育及生物产量产生影响,最终将影响人类的生存环境,因此,近年来对其研究逐渐增多。综述了国内外学者对植物的形态学特征、生物量的变化、生理生化机制、根系分泌物及根际微生物等受大气中CO2浓度升高的影响等方面的研究。结果表明,CO2的浓度升高会增加植物的根系面积、增加叶片的厚度、促进根系分泌物的释放,改变根际微生物的多样性,也可能会提高植物的呼吸作用和光合速率。     

大气CO2浓度上升对植物地下竞争的影响

植物地下竞争是影响农业生态系统和自然生态系统中植物群落结构变化的重要因素,而大气CO2浓度升高直接或间接影响了植物的生长及其地下环境,可能需重新评估植物地下竞争的状况。本文从大气CO2浓度升高对植物根系形态结构、生理吸收能力、根系共生真菌、叶片蒸腾速率的影响以及对地下竞争因子的土壤环境方面的影响,探讨未来CO2浓度升高条件下地下竞争的变化。     

大气CO2浓度升高对植物 土壤系统地下过程影响的研究

综述了大气CO2浓度升高对根系、根际、根系分泌物、土壤呼吸和土壤物质转化和C、N循环影响的研究进展,阐述了有关实验的研究情况,以及它们在整个生态系统响应大气CO2浓度升高中的重要作用、目前研究中存在的争论、以及还需要研究的领域和方向及其研究的重要性。     

大气CO2与植物氮素营养的关系

大气CO2浓度升高对植物吸收氮素，以及对植物和土壤中的氮浓度，C／N比和氮循环都存在着影响。大气CO2浓度与植物氮素营养之间存在着交互作用。大气CO2浓度升高对植物氮素营养物结果与氮浓度，氮形态等因素有关。     

土壤微生物生物量和呼吸强度对大气CO2浓度升高的响应

随着全球环境变化对陆地生态系统的影响逐渐成为公众和科学界关注的热点,CO2作为一种重要的温室气体受到格外重视.大气CO2浓度升高将直接影响陆地植物的光合作用[1].植物的光合产物约有20% ～ 50%被运送到地下,通过根系分泌及死亡输入土壤[2],因此大气CO2浓度升高将会间接影响土壤生态系统.长期以来,关于大气CO2浓度升高对农作物地上部分的研究较多,但关于大气CO2浓度升高对土壤特别是土壤微生物的影响的研究报道较少.     

高CO2浓度下根系分泌物的研究进展

根系分泌物是植物对大气CO2浓度升高响应的调节器,它能活化土壤养分元素、调节微生物区系组成,在根际微生态系统中扮演重要角色。本文综述了大气CO2浓度升高及其导致的温度上升引起根系分泌物数量和组成上的变化,讨论了这种变化与土壤微生物的关系和对全球C循环的贡献,并根据现状提出了今后的研究方向。     

CO2浓度升高对2个品种水稻Cu吸收、根形态和植物络合素合成的影响

为探讨CO2浓度升高条件下不同水稻品种粤杂889 (YZ)和荣优398(RY)对耐Cu胁迫性的变化特征,利用水培试验研究不同Cu浓度下CO2浓度升高对2种水稻幼苗生物量、Cu含量、根形态及植物络合素(GSH和PCs)的影响.结果表明,低铜处理对水稻生长具有促进作用,增加2种水稻生物量及根系根毛数、总根长、表面积和体积.随着Cu处理浓度升高,根系GSH和PCs含量分别呈现渐减和渐增趋势.CO2浓度升高条件下,2种水稻生物量显著增加,600 μmol/L Cu处理时增加比例最大,YZ和RY分别增加59.8％和49.0％;水稻根、茎叶Cu含量降低,但根系形态各个指标明显增加,且在高Cu处理下其增加比例较大.CO2浓度升高显著增加根系PCs合成,50 μmol/L Cu处理时增加比例最大,YZ和RY分别增加121.6％,78.7％.在CO2浓度正常与升高条件下,根系GSH、PCs含量与Cu浓度都具有显著相关性.CO2浓度升高通过增加根系形态和PCs含量以增强水稻对Cu的抗逆性,但存在着品种差异,YZ的增加比例大于RY.     

CO_2浓度升高对土壤微生物及土壤酶影响的研究进展

概述了大气CO2浓度条件下,土壤微生物生物量C、土壤微生物区系、土壤细菌群落结构以及土壤酶的变化等方面的研究进展。并提出在CO2浓度升高条件下,根际微生物的优势种群的变化趋势和根系分泌物与根际微生物之间相互影响的研究是今后的研究趋向。     

CO2浓度与氮磷供应水平对黄瓜根系生长及各组织矿质养分含量的影响

在开顶式生长箱内，以黄瓜为试验材料，采用营养液培养方法，研究了不同氮水平、磷水平条件下大气CO2浓度对黄瓜植株内矿质养分含量以及根系形态的影响。结果表明：黄瓜植株各部位氮素含量随供氮水平提高而增加，磷水平提高，也能促进各部位氮含量的提高。植株各部位磷含量随供磷水平的提高而升高，在相同磷水平下，缺氮会使各部位磷含量升高。大气CO2浓度升高会使黄瓜植株各部位氮及特定部位的磷含量降低。黄瓜根部的Ca含量随CO2浓度的升高而显著降低，氮和磷水平的升高极显著地增加了其含量，且CO2浓度与供磷水平、供氮与供磷水平以及这三者之间存在明显的交互作用。供氮、供磷水平的升高极显著的提高了黄瓜叶片Ca的含量以及茎部Mg的含量，且两者存在明显的交互作用。黄瓜总根长和总根表面积随CO2浓度的增加有增大的趋势；在缺磷条件下，总根长和总根表面积随氮水平的提高而增大；而同一氮水平和CO2浓度下，磷水平的降低会增加总根长和总根表面积。总体看来，大气CO2浓度的升高能促进黄瓜根系的生长，但会使得黄瓜植株某些部位氮、磷、钙、镁等矿质元素含量降低，而供氮、供磷水平的提高可以通过增强黄瓜的生长与活力促进黄瓜根系对矿质养分的吸收，从而缓解由于CO2浓度升高带来的矿质元素含量降低的风险。这启示我们在对设施蔬菜CO2施肥的同时，也要注重适量提高合理配比下矿质元素的供应。     

CO2浓度升高对不同水稻品种幼苗养分吸收和根系形态的影响

本文利用水培试验研究了CO2浓度升高对水稻幼苗生物量、养分含量和根形态的影响,探讨了CO2浓度升高下粤杂889(YZ)和荣优398 (RY)幼苗养分吸收和根系形态的差异性.结果表明,与CO2浓度正常水平(对照)相比,CO2浓度升高显著增加了2个水稻品种幼苗根系、茎叶和总生物量,YZ分别增加58.33％、27.96％、33.16％;RY分别增加45.87％、34.17％、36.07％.同时,CO2浓度升高增加了2个水稻品种的根冠比.CO2浓度升高显著降低了2个水稻品种茎叶中的N、P、K、Ca、Mg和Fe含量,这是“稀释效应”的结果;但YZ幼苗中S含量显著增加,2个品种幼苗Mn含量均显著增加.CO2浓度升高显著增加了2个水稻品种的幼苗根系根毛数、总根长、表面积,降低幼苗粗根比例,增加了细根比例.CO2浓度升高增加了细根在总根长中的比例,有利于水稻对养分的吸收,导致部分营养元素含量增加;但CO2浓度升高条件下水稻生物量的增加使大部分营养元素含量降低.同时,CO2浓度升高对水稻幼苗生物量、养分吸收和根形态的影响存在显著的品种差异.     

高低应答CO2水稻品种苗期根系对高C环境的响应

本文利用水培试验和琼脂板培养试验研究了高CO₂条件下产量响应存在显著差异的两个水稻品种：II优084(高响应)和武运粳23(低响应),在幼苗期根系形态对高C的响应差异。水培试验结果表明,在幼苗时期,高应答品种II优084在低氮条件下地上部生物量在高CO₂下增加28.5%,根系干物质量对高CO₂响应显著,增幅为28.5%,而其不定根数目没有显著增加,对干物质量响应贡献较大的为总根长。II优084的总根长在高CO₂下增幅为26.3%,不同根粗的根长均有高响应。低应答品种武运粳23低氮下地上部和根系响应不显著,而在正常氮和高氮下则不同。正常氮条件下,地上部对高CO₂响应不显著,而根系生物量在高CO₂下显著增加76.0%,不定根数目增加25.8%,同时总根长增加45.0%,不同根粗的根长均有高响应,II优084则没有显著响应。在高氮条件下,武运粳23地上部生物量在高CO₂下增加35.5%,根系生物量增加80.3%,不定根数目增加38.5%,根系平均直径增加16.7%,总根长无响应,而II优084生物量在高氮下无显著差异。同时,武运粳23在正常氮和高氮下的根系表面积和体积对高CO₂响应也较II优084显著。琼脂板培养试验的结果与水培结果一致,武运粳23根系形态对高浓度蔗糖的响应普遍高于II优084。试验结果说明品种对高C环境的响应特征不随培养条件的变化而变化。与植株生长后期不同,在幼苗期正常氮条件下低应答品种武运粳23的根系生物量和各形态指标对高C的响应明显高于II优084,说明水稻苗期生长响应参数与后期产量响应参数不一定一致,可能是由于苗期生长高响应的品种在营养生长期旺长,反而不利于后期生殖生长,从而导致后期产量的低响应。     

Elevated CO2 and plant species diversity interact to slow root decomposition

Changes in plant species diversity can result in synergistic increases in decomposition rates, while elevated atmospheric CO₂ can slow the decomposition rates; yet it remains unclear how diversity and changes in atmospheric CO₂ may interact to alter root decomposition. To investigate how elevated CO₂ interacts with changes in root-litter diversity to alter decomposition rates, we conducted a 120-day laboratory incubation. Roots from three species (Trifolium repens, Lespedeza cuneata, and Festuca pratense) grown under ambient or elevated CO₂ were incubated individually or in combination in soils that were exposed to ambient or elevated CO₂ for five years. Our experiment resulted in two main findings: (1) Roots from T. repens and L. cuneata, both nitrogen (N) fixers, grown under elevated CO₂ treatments had significantly slower decomposition rates than similar roots grown under ambient CO₂ treatments; but the decomposition rate of F. pratense roots (a non-N-fixing species) was similar regardless of CO₂ treatment. (2) Roots of the three species grown under ambient CO₂ and decomposed in combination with each other had faster decomposition rates than when they were decomposed as single species. However, roots of the three species grown under elevated CO₂ had similar decomposition rates when they were incubated alone or in combination with other species. These data suggest that if elevated CO₂ reduces the root decomposition rate of even a few species in the community, it may slow root decomposition of the entire plant community.     

Effects of Atmospheric CO2 Enrichment on Crop Nutrient Dynamics under No-Till Conditions

Increasing atmospheric CO₂ concentration could increase crop productivity and alter crop nutrient dynamics. This study was conducted (3 yrs) with two crops ([Glycine max (L.) Merr.] and grain sorghum [Sorghum bicolor (L.) Moench.]) grown under two CO₂ levels (ambient and twice ambient) using open top field chambers on a Blanton loamy sand under no-tillage. Macronutrient and micronutrient concentrations and contents were determined for grain, stover, and roots. Although elevated CO₂ tended to reduce nutrient concentrations, high CO₂ consistently increased nutrient content especially in grain tissue; this response pattern was more notable with macronutrients. The CO₂ effect was observed primarily in soybean. The consistent CO₂-induced increases in grain macronutrient contents favors reliable predictions of system outputs, however, predictions of crop nutrient inputs (i.e., stover and root contents) to the soil are less robust due to observed variability. Again, this is particularly true in regards to micronutrient dynamics in CO₂-enriched cropping systems.     

Effects of elevated CO2 and O3 on soil respiration under ponderosa pine

Soil respiration represents the integrated response of plant roots and soil organisms to environmental conditions and the availability of C in the soil. A multi-year study was conducted in outdoor sun-lit controlled-environment chambers containing a reconstructed ponderosa pine/soil-litter system. The study used a 2×2 factorial design with two levels of CO₂ and two levels of O₃ and three replicates of each treatment. The objectives of our study were to assess the effects of long-term exposure to elevated CO₂ and O₃, singly and in combination, on soil respiration, fine root growth and soil organisms. Fine root growth and soil organisms were included in the study as indicators of the autotrophic and heterotrophic components of soil respiration. The study evaluated three hypotheses: (1) elevated CO₂ will increase C assimilation and allocation belowground increasing soil respiration; (2) elevated O₃ will decrease C assimilation and allocation belowground decreasing soil respiration and (3) as elevated CO₂ and O₃ have opposing effects on C assimilation and allocation, elevated CO₂ will eliminate or reduce the negative effects of elevated O₃ on soil respiration. A mixed-model covariance analysis was used to remove the influences of soil temperature, soil moisture and days from planting when testing for the effects of CO₂ and O₃ on soil respiration. The covariance analysis showed that elevated CO₂ significantly reduced the soil respiration while elevated O₃ had no significant effect. Despite the lack of a direct CO₂ stimulation of soil respiration, there were significant interactions between CO₂ and soil temperature, soil moisture and days from planting indicating that elevated CO₂ altered soil respiration indirectly. In elevated CO₂, soil respiration was more sensitive to soil temperature changes and less sensitive to soil moisture changes than in ambient CO₂. Soil respiration increased more with days from planting in elevated than in ambient CO₂. Elevated CO₂ had no effect on fine root biomass but increased abundance of culturable bacteria and fungi suggesting that these increases were associated with increased C allocation belowground. Elevated CO₂ had no significant effect on microarthropod and nematode abundance. Elevated O₃ had no significant effects on any parameter except it reduced the sensitivity of soil respiration to changes in temperature.     

Changes in soil microbial biomass carbon and enzyme activities under elevated CO2 affect fine root decomposition processes in a Mongolian oak ecosystem

The relationships between soil microbial properties and fine root decomposition processes under elevated CO₂ are poorly understood. To address this question, we determined soil microbial biomass carbon (SMB-C) and nitrogen (SMB-N), enzymes related to soil carbon (C) and nitrogen (N) cycling, the abundance of cultivable N-fixing bacteria and cellulolytic fungi, fine root organic matter, lignin and holocellulose decomposition, and N mineralization from 2006 to 2007 in a Mongolian oak (Quercus mongolica Fischer ex Ledebour) ecosystem in northeastern China. The experiment consisted of three treatments: elevated CO₂ chambers, ambient CO₂ chambers, and chamberless plots. Fine roots had significantly greater organic matter decomposition rates under elevated CO₂. This corresponded with significantly greater SMB-C. Changes in the activities of protease and phenol oxidase under elevated CO₂ could not explain the changes in fine root N release and lignin decomposition rates, respectively, while holocellulose decomposition rate had the same response to experimental treatments as did cellulase activity. Changes in cultivable N-fixing bacterial and cellulolytic fungal abundances in response to experimental treatments were identical to those of N mineralization and lignin decomposition rates, respectively, suggesting that the two indices were closely related to fine root N mineralization and lignin decomposition. Our results showed that the increased fine root organic matter, lignin and holocellulose decomposition, and N mineralization rates under elevated CO₂ could be explained by shifts in SMB-C and the abundance of cellulolytic fungi and N-fixing bacteria. Enzyme activities are not reliable for the assessment of fine root decomposition and more attention should be given to the measurement of specific bacterial and fungal communities.     

Elevation of atmospheric CO2 and N-nutritional status modify nodulation, nodule-carbon supply, and root exudation of Phaseolus vulgaris L.

Increased root exudation and a related stimulation of rhizosphere-microbial growth have been hypothesised as possible explanations for a lower nitrogen- (N-) nutritional status of plants grown under elevated atmospheric CO₂ concentrations, due to enhanced plant-microbial N competition in the rhizosphere. Leguminous plants may be able to counterbalance the enhanced N requirement by increased symbiotic N₂ fixation. Only limited information is available about the factors determining the stimulation of symbiotic N₂ fixation in response to elevated CO₂.In this study, short-term effects of elevated CO₂ on quality and quantity of root exudation, and on carbon supply to the nodules were assessed in Phaseolus vulgaris, grown in soil culture with limited (30 mg N kg⁻¹ soil) and sufficient N supply (200 mg N kg⁻¹ soil), at ambient (400 μmol mol⁻¹) and elevated (800 μmol mol⁻¹) atmospheric CO₂ concentrations.Elevated CO₂ reduced N tissue concentrations in both N treatments, accelerated the expression of N deficiency symptoms in the N-limited variant, but did not affect plant biomass production. ¹⁴CO₂ pulse-chase labelling revealed no indication for a general increase in root exudation with subsequent stimulation of rhizosphere microbial growth, resulting in increased N-competition in the rhizosphere at elevated CO₂. However, a CO₂-induced stimulation in root exudation of sugars and malate as a chemo-attractant for rhizobia was detected in 0.5-1.5 cm apical root zones as potential infection sites. Particularly in nodules, elevated CO₂ increased the accumulation of malate as a major carbon source for the microsymbiont and of malonate with essential functions for nodule development. Nodule number, biomass and the proportion of leghaemoglobin-producing nodules were also enhanced. The release of nod-gene-inducing flavonoids (genistein, daidzein and coumestrol) was stimulated under elevated CO₂, independent of the N supply, and was already detectable at early stages of seedling development at 6 days after sowing.     

Decomposition of C-labeled roots in a pasture soil exposed to 10 years of elevated CO2

The net flux of soil C is determined by the balance between soil C input and microbial decomposition, both of which might be altered under prolonged elevated atmospheric CO₂. In this study, we determined the effect of elevated CO₂ on decomposition of grass root material (Lolium perenne L.). ¹⁴C-labeled root material, produced under ambient (35 Pa pCO₂) or elevated CO₂ (70 Pa pCO₂) was incubated in soil for 64 days. The soils were taken from a pasture ecosystem which had been exposed to ambient (35 Pa pCO₂) or elevated CO₂ (60 Pa pCO₂) under FACE-conditions for 10 years and two fertilizer N rates: 140 and 560 kg N ha⁻¹ year⁻¹. In soil exposed to elevated CO₂, decomposition rates of root material grown at either ambient or elevated CO₂ were always lower than in the control soil exposed to ambient CO₂, demonstrating a change in microbial activity. In the soil that received the high rate of N fertilizer, decomposition of root material grown at elevated CO₂ decreased by approximately 17% after incubation for 64 days compared to root material grown at ambient CO₂. The amount of ¹⁴CO₂ respired per amount of ¹⁴C incorporated in the microbial biomass (q¹⁴CO₂) was significantly lower when roots were grown under high CO₂ compared to roots grown under low CO₂. We hypothesize that this decrease is the result of a shift in the microbial community, causing an increase in metabolic efficiency. Soils exposed to elevated CO₂ tended to respire more native SOC, both with and without the addition of the root material, probably resulting from a higher C supply to the soil during the 10 years of treatment with elevated CO₂. The results show the importance of using soils adapted to elevated CO₂ in studies of decomposition of roots grown under elevated CO₂. Our results further suggest that negative priming effects may obscure CO₂ data in incubation experiments with unlabeled substrates. From the results obtained, we conclude that a slower turnover of root material grown in an ‘elevated-CO₂ world’ may result in a limited net increase in C storage in ryegrass swards.     

CO2浓度升高对番茄苗根系生长及其吲哚乙酸和乙烯含量的影响

A hydroponic experiment was carried out to study the effect of elevated carbon dioxide (CO2) on root growth of tomato seedlings. Compared with the control (350 μL L-1), CO2 enrichment (800 μL L-1) significantly increased the dry matter of both shoot and root, the ratio of root to shoot, total root length, root surface area, root diameter, root volume, and root tip numbers, which are important for forming a strong root system. The elevated CO2 treatment also significantly improved root hair development and elongation, thus enhancing nutrient uptake. Increased indole acetic acid concentration in plant tissues and ethylene release in the elevated CO2 treatment might have resulted in enhanced root growth and root hair development and elongation.     

增施CO2降低小白菜硝酸盐积累的机理研究

以低硝酸盐积累基因型（东妃）和高硝酸盐积累基因型（高雄甜脆）两种小白菜为材料,采用溶液培养法研究了增施CO2降低蔬菜硝酸盐积累的生理机制。结果表明,CO2浓度升高能显著提高2种基因型小白菜的生物量和硝酸还原酶活性,并降低根、茎叶各部位的硝酸盐含量。CO2浓度升高不仅促进了植株对硝态氮的吸收,而且植株吸收硝酸盐的累积量增幅均高于鲜重的增幅。由此可见,除了鲜重增加的稀释作用,处理后生理机制的变化也可能是CO2浓度升高引起硝酸盐含量降低的重要原因。研究还表明,增施CO2后“东妃”的硝酸盐含量降低百分率与硝酸还原酶活性的增加百分率呈极显著相关,而“高雄甜脆”的硝酸盐含量降低百分率则与鲜重的增加百分率的相关性达极显著水平。说明增施CO2后植株各部位硝酸还原酶活性提高及鲜重的增加均为引起硝酸盐含量降低的重要原因,但贡献率具有明显的基因型差异。