开放式空气CO2浓度增高对水稻生长发育影响的研究进展

地球大气中CO₂浓度不断升高已是不争的事实.CO₂浓度升高势必对植物的生长发育过程产生深刻的影响.水稻是世界上最重要的作物之一,也是中国第一大作物.结合气室条件下的研究结果,从光合作用、水分关系、生育期、叶片和根系生长、物质生产与分配、化学组分以及产量和品质等方面,重点收集和整理了开放式空气中CO₂浓度增高 (FACE) 对水稻生长发育影响的研究进展,并讨论了该领域有待深入研究的方向.     

大气CO2浓度升高对森林食叶昆虫的潜在影响

评述了大气CO₂浓度升高对森林食叶昆虫的影响,昆虫对森林取食为害水平的潜在变化,以及研究中的主要实验方法.大气CO₂浓度升高通过引起叶片化学变化进而影响食叶昆虫个体的取食和生长;但物种对环境变化反应的特异性、植物化学对高浓度CO₂的反应强度、昆虫对植物生理变化的敏感性和适应性、研究周期的长短、其它环境因子的协同效应以及不同实验中植物生长条件和研究方法的差异均将影响昆虫反应的方向和强度;CO₂气体浓度增高本身可能不足以对食叶昆虫个体的新陈代谢构成影响;大气CO₂浓度升高也可能影响森林食叶昆虫种群的大小.     

大气CO2浓度升高对不同施氮土壤酶活性的影响

利用中国唯一的无锡FACE（Free-air CO₂ enrichment,开放式空气CO₂浓度升高）平台,研究了大气CO₂浓度升高对土壤β-葡糖苷酶、转化酶、脲酶、酸性磷酸酶、β-氨基葡糖苷酶的影响。研究发现,不同氮肥处理下大气CO₂浓度升高对某些土壤酶活性的影响不同。在低氮施肥处理中,大气CO₂浓度升高显著降低β-葡糖苷酶活性,但是在高氮施肥处理下,大气CO₂浓度升高显著增加β-葡糖苷酶活性。在低氮和常氮施肥处理中大气CO₂浓度升高显著增加了土壤脲酶活性,但在高氮水平下影响不显著。在低氮、常氮施肥处理中,大气CO₂浓度升高对土壤酸性磷酸酶活性没有影响,而在高氮施肥处理中显著增强了土壤中磷酸酶活性。大气CO₂浓度升高对土壤转化酶活性和β-氨基葡糖苷酶的活性有增加趋势,但影响不显著。研究还发现,在不同的CO₂浓度下,土壤酶活性对不同氮肥处理的响应也不同。在正常CO₂浓度下,土壤中β-葡糖苷酶活性随着氮肥施用量的增加而降低,而在大气CO₂浓度升高条件下,却随着氮肥施用量的增加而增加。在大气CO₂浓度升高条件下,高氮施肥显著增加了转化酶和酸性磷酸酶活性,而在正常CO₂浓度下,影响不显著。在大气CO₂浓度升高条件下,氮肥处理对脲酶活性的影响不大,但在正常CO₂浓度下,脲酶活性随着氮肥施用量的增加而增加。氮肥对β-氨基葡糖苷酶活性的影响不明显。     

转PEPC基因水稻具有初级CO2浓缩机制生理特点

以原种粳稻Kitaake为对照, 研究了转玉米PEPC基因水稻的PEPC的高表达和碳同化特性的关系. 结果显示: 与原种相比, 转PEPC基因水稻气孔导度和光合速率显著增加, 经统计分析, 气孔导度的增加与光合速率的增加并无相关性. 而在高光强 下, 与CO₂浓缩有关的PEPC, CA酶蛋白的表达显著增加. 因此在大气CO₂浓度下可显著增加光合能力(50%); 无CO₂条件下, 可减少叶内CO₂释放量, 从而降低了CO₂补偿点. 用专一的抑制剂DCDP处理, 证明转PEPC基因水稻叶内PEPC的高表达与碳同化能力的提高和Fv/Fm的稳定性有关. 用14C示踪20 s, 转PEPC基因水稻14C较多的分配在C4光合原初产物天冬氨酸中, 意味着叶内存在着一定的C4光合代谢途径. 上述结果说明, 用代谢工程可以在叶内构建初级的CO₂浓缩机制, 为转基因的高光效育种技术提供了生理依据.     

林冠与大气间二氧化碳交换过程的模拟

以长白山阔叶红松林为研究对象,利用Raupach提出的局地近场理论(localized near field, LNF)耦合垂直风速标准差σ_w（z）和拉格朗日时间尺度T_L （z）,建立林冠内CO₂源汇强度和平均浓度廓线之间的关系.结果表明,拉格朗日模型能准确、稳定地模拟林冠与大气之间CO₂的交换特征.模拟值比涡动相关系统实测值高出约15%,与实测值的相关性为89%,这种差异可能主要来自于输入的浓度廓线的波动以及大气稳定层结造成的涡动相关观测系统误差.在近地面层,由于土壤呼吸作用,整个时间段都为CO₂源.林冠层的CO₂源汇强度变化较为复杂,其日变化经历了源－汇－源的转变过程.林冠与大气间CO₂通量交换明显受大气稳定度影响.     

FACE条件下水稻冠层蒸散和水分利用率的模拟

利用开放式CO₂浓度增高（FACE）系统平台,通过在水稻拔节期至成熟期对水稻冠层微气候及相关生理指标的连续观测,并结合能量平衡分析,模拟研究了FACE对水稻冠层蒸散和水分利用率的影响.结果表明：将水稻叶片气孔导度与光合有效辐射、饱和水气压差的定量关系与Penman-Monteith方程相结合,可以较好地模拟FACE和对照条件下的水稻蒸散量;观测期间,CO₂浓度升高使水稻的水分利用比对照减小约10 mm,结合水稻生物量增加12%,FACE条件下水稻水分利用率（WUE）增加约12%.     

大豆对臭氧、二氧化碳及其复合效应的响应

以大豆‘中黄14'为试验材料,首次模拟研究大气中O₃、CO₂浓度增加,及其逐渐和持续增加O₃、CO₂浓度复合效应对大豆的影响.结果表明,CO₂浓度增加可缓解O₃对叶片的伤害程度,受害时间推迟,受害症状无实质性变化.熏气20 d测定各处理叶片生理参数发现,在本底大气环境下,叶片气孔阻力和蒸腾速率与对照差异较小,熏气时O₃、CO₂浓度增加诱导叶片气孔关闭,气孔阻力明显增加,蒸腾速率显著降低.与对照相比,O₃浓度增加,大豆干物质积累、产量和收获指数明显降低,籽粒粗脂肪含量明显减少,粗蛋白含量显著增加;CO₂浓度增加,干物质积累和产量显著提高,收获指数无明显差异,籽粒粗脂肪和粗蛋白含量均明显减少;逐渐和持续增加O₃和CO₂浓度复合效应处理下,大豆干物质积累、产量和收获指数差异不明显,籽粒粗蛋白含量不同程度地减少,粗脂肪含量显著增加.     

CO2浓度升高对红松和长白松土壤呼吸作用的影响

以开顶箱法研究了CO₂浓度升高对红松和长白松土壤呼吸作用的影响.结果表明,500 μmol CO₂·mol^-1使红松和长白松土壤呼吸速率明显降低,土壤表面CO₂浓度升高导致CO₂扩散受阻可能是土壤呼吸受到抑制的主要原因.500 μmol CO₂·mol^-1下两树种土壤表面CO₂浓度明显高于对照箱和裸地条件下的CO₂浓度,增加幅度在40～150 μmol·mol^-1之间;对照箱内长白松土壤表面CO₂浓度略高于裸地,差异不显著,红松差异显著500 μmol CO₂·mol^-1下的长白松土壤全氮及总有机碳含量略高于对照组,差异不显著,红松裸地的碳氮含量明显低于500 μmol CO₂·mol^-1 及对照箱内土壤碳氮含量;500 μmol CO₂·mol^-1 及开顶箱的微环境对地下3 cm处土壤温度没有明显影响.     

开放式空气二氧化碳浓度增高对小麦产量形成的影响

利用农田开放式空气CO₂浓度增高(FACE)系统平台,以弱筋小麦宁麦9号为供试品种,研究大气CO₂浓度增高和不同施氮水平对小麦生育期、株高、产量和产量构成因素的影响.结果表明：FACE处理的小麦播种至抽穗期、抽穗至成熟期及全生育期天数分别比对照缩短1.3、1.3和2.6 d,但均未达到显著水平;FACE处理的小麦穗长、穗下第1和第2节间长度显著变长,成熟期株高显著增加,比对照增加4.0%;低、中、高氮条件下,FACE处理小麦的籽粒产量分别比对照提高15.2%、21.4%和35.4%,平均增产24.6%,均达极显著水平;FACE处理小麦的单位面积穗数极显著增加,比对照增加17.8%,使穗粒数和粒重显著增加,分别比对照增加了2.9%和4.8%.FACE处理使小麦显著增产主要是由于单位面积穗数显著增加,而单位面积穗数的增加主要是由于小麦的分蘖能力明显增强所致.     

春小麦对大气CO2浓度升高的响应及其对麦长管蚜生长发育和繁殖的影响

通过开顶式气室研究了春小麦对大气CO₂浓度升高(542.1±24.8和738.8±25.7 μl·L^-1 vs.382.4±248 μl·L^-1)的响应及其对麦长管蚜生长发育和繁殖的影响.结果表明,大气CO₂浓度升高有利于春小麦的生长.与对照相比,5月5日～6月14日,550和750 μl·L^-1 CO₂浓度处理春小麦的株高分别增加2.80%～14.92%和6.30%～17.56%;4月30日～6月9日,叶面积分别增加5.68%～50.52%和6.14%～83.45%;DC₅₀分别提前了0.39和0.90 d,DC₇₅也分别提前了0.53和1.02 d;茎、叶、穗以及整个地上部组织的鲜、干重均有不同程度的增加.大气CO₂浓度升高可显著提高春小麦的穗长和穗粒数,降低千粒重.与对照相比,550和750 μl·L^-1 CO₂浓度处理的麦穗长分别增加0.56%和3.20%;单株穗粒数分别增加12.5%和18%;而千粒重分别降低了2.23%和6.34%.随着大气中CO₂浓度增加,麦穗中葡萄糖、二糖、多糖、总糖、总糖与总氮的比值都显著增加,而果糖、三糖和总氮含量都显著降低.大气CO₂浓度升高可缩短麦长管蚜的产卵前期和世代历期,提高繁殖量和平均相对生长率.与对照相比,550和750 μl·L^-1 CO₂浓度处理麦长管蚜的平均相对生长率分别提高33.26%和74.34%.麦长管蚜种群的平均相对生长率与寄主麦穗中总糖和总氮的比值相关显著.     

Climate change impacts on crop yield and quality with CO2 fertilization in China

A regional climate change model (PRECIS) for China, developed by the UK's Hadley Centre, was used to simulate China's climate and to develop climate change scenarios for the country. Results from this project suggest that, depending on the level of future emissions, the average annual temperature increase in China by the end of the twenty-first century may be between 3 and 4 degrees C. Regional crop models were driven by PRECIS output to predict changes in yields of key Chinese food crops: rice, maize and wheat. Modelling suggests that climate change without carbon dioxide (CO2) fertilization could reduce the rice, maize and wheat yields by up to 37% in the next 20-80 years. Interactions of CO2 with limiting factors, especially water and nitrogen, are increasingly well understood and capable of strongly modulating observed growth responses in crops. More complete reporting of free-air carbon enrichment experiments than was possible in the Intergovernmental Panel on Climate Change's Third Assessment Report confirms that CO2 enrichment under field conditions consistently increases biomass and yields in the range of 5-15%, with CO2 concentration elevated to 550 ppm Levels of CO2 that are elevated to more than 450 ppm will probably cause some deleterious effects in grain quality. It seems likely that the extent of the CO2 fertilization effect will depend upon other factors such as optimum breeding, irrigation and nutrient applications.     

基于CTGC试验系统下面包小麦主要品质性状的研究

 当前,全球气候变化背景下,CO2浓度升高和气候变暖可能已经并将持续对小麦（Triticum aestivum）品质产生影响。为此依据自行设计的模拟气候变化的试验装置系统 （CTGC）,研究大田条件下CO2浓度和温度增加对面包小麦主要品质性状的交互作用。 结果表明：在433.3～610.2μmol•mol-1范围内,CO2浓度增加对面包小麦的籽粒蛋白质含量、湿面筋含量和沉降值的影响不利;增温（+2 ℃多）表现为有利。当CO2浓度从433.3 mol•mol-1逐渐增加到551.5 mol•mol-1且温度增幅逐渐为+2 ℃时,CO2和温度对面包小麦的籽粒蛋白质含量、湿面筋含量和沉降值的交互作用表现为增加;而当CO2浓度增幅较大（达到610.2 mol•mol-1）,温度增幅不大（白天平均温度仅增加2℃多）时,交互作用则 表现为减少。此外,CO2浓度增加使面包小麦的醎淀粉酶活性降低,温度上升则使之提高,两因素的交互作用则表现为醎淀粉酶活性提高。     

Effects of elevated CO2 concentration on the quality of agricultural products: a review

The increasing concentration of atmospheric CO2 and the nutritional quality of human diets are the two important issues we are facing. At present, the atmospheric CO2 concentration is about 380 micromol mol(-1), and to be reached 550 micromol mol(-1) by 2050. A great deal of researches indicated that the quality of agricultural products is not only determined by inherited genes, but also affected by the crop growth environmental conditions. This paper summarized the common methods adopted at home and abroad for studying the effects of CO2 enrichment on the quality of agricultural products, and reviewed the research advances in evaluating the effects of elevated CO2 on the quality of rice, wheat, soybean, and vegetables. Many experimental results showed that elevated CO2 concentration causes a decrease of protein content in the grains of staple food crops and an overall decreasing trend of trace elements contents in the crops, but improves the quality of vegetable products to some extent. Some issues and future directions regarding the effects of elevated CO2 concentration on the quality of agricultural products were also discussed, based on the present status of related researches.     

Responses of wheat and rice to factorial combinations of ambient and elevated CO2 and temperature in FACE experiments

Elevated CO₂ and temperature strongly affect crop production, but understanding of the crop response to combined CO₂ and temperature increases under field conditions is still limited while data are scarce. We grew wheat (Triticum aestivum L.) and rice (Oryza sativa L.) under two levels of CO₂ (ambient and enriched up to 500 μmol mol^?1) and two levels of canopy temperature (ambient and increased by 1.5–2.0 °C) in free‐air CO₂ enrichment (FACE) systems and carried out a detailed growth and yield component analysis during two growing seasons for both crops. An increase in CO₂ resulted in higher grain yield, whereas an increase in temperature reduced grain yield, in both crops. An increase in CO₂ was unable to compensate for the negative impact of an increase in temperature on biomass and yield of wheat and rice. Yields of wheat and rice were decreased by 10–12% and 17–35%, respectively, under the combination of elevated CO₂ and temperature. The number of filled grains per unit area was the most important yield component accounting for the effects of elevated CO₂ and temperature in wheat and rice. Our data showed complex treatment effects on the interplay between preheading duration, nitrogen uptake, tillering, leaf area index, and radiation‐use efficiency, and thus on yield components and yield. Nitrogen uptake before heading was crucial in minimizing yield loss due to climate change in both crops. For rice, however, a breeding strategy to increase grain number per m² and % filled grains (or to reduce spikelet sterility) at high temperature is also required to prevent yield reduction under conditions of global change.     

Effects of atmospheric CO2 concentration enhancement and nitrogen application rate on wheat grain yield and quality

FACE platform was applied to study the effects of elevated atmospheric CO2 concentration on wheat grain yield and quality under two nitrogen (N) application rates. Elevated atmospheric CO2 concentration and applying N increased the grain yield, spike number, grain number per spike, and biomass significantly, but elevated CO2 concentration had no significant effects on harvest index (HI). Under elevated CO2 concentration, there was a significant decrease in the protein, gliadin, gluteinin, and glutein contents of the grain and the sedimentation value of the flour, and a significant increase in the starch and its components contents of the grain; under N application, an inverse was observed. The dough stability time and the dough viscosity characteristics, such as peak viscosity, final viscosity, and setback value, increased significantly under elevated CO2 concentration and high N application rate. The interaction of atmospheric CO2 concentration and N application rate had significantly positive effects on wheat grain yield and biomass, but less effect on grain quality. Therefore, with elevated atmospheric CO2 concentration in the future, maintaining a higher N application level would benefit wheat grain yield and paste characteristics, and mitigate the decline of grain quality.     

Understanding the regulation of cereal grain filling: The way forward

During grain filling, starch and other nutrients accumulate in the endosperm; this directly determines grain yield and grain quality in crops such as rice (Oryza sativa), maize (Zea mays), and wheat (Triticum aestivum). Grain filling is a complex trait affected by both intrinsic and environmental factors, making it difficult to explore the underlying genetics, molecular regulation, and the application of these genes for breeding. With the development of powerful genetic and molecular techniques, much has been learned about the genes and molecular networks related to grain filling over the past decades. In this review, we highlight the key factors affecting grain filling, including both biological and abiotic factors. We then summarize the key genes controlling grain filling and their roles in this event, including regulators of sugar translocation and starch biosynthesis, phytohormone-related regulators, and other factors. Finally, we discuss how the current knowledge of valuable grain filling genes could be integrated with strategies for breeding cereal varieties with improved grain yield and quality.     

大气CO2浓度和氮肥水平对小麦籽粒产量和品质的影响

应用FACE平台,研究了不同氮肥水平下大气CO₂浓度升高对小麦籽粒产量和品质的影响.结果表明： 大气CO₂浓度升高（FACE）和增施氮肥显著提高了小麦籽粒产量、穗数、穗粒数和生物量.但FACE处理对收获指数无显著影响.FACE处理显著降低了籽粒蛋白质、醇溶蛋白、谷蛋白、面筋含量和沉降值,显著提高了小麦籽粒淀粉及其组分含量,而氮肥处理具有相反的效应.面团稳定时间及峰值黏度、最终黏度、反弹值等黏度特征参数在FACE和高氮水平下显著增加.此外,CO₂浓度与氮肥水平互作对小麦籽粒产量和生物量有显著的正效应,但对籽粒品质无显著影响.在未来大气CO₂浓度升高的情况下,维持较高的施氮量有利于提高小麦籽粒产量,改善淀粉糊化特性,缓解小麦品质特性的下降.     

气候变化对我国干旱/半干旱区小麦生产影响的模拟研究

利用随机天气模型,将气候模式对大气中CO₂倍增时预测的气候情景与CERES-小麦模式相连接,研究了气候变化对我国冬小麦和春小麦生产的可能影响。并对水分、温度、CO₂综合对小麦的作用进行初步模拟分析。所得结论为:①气候变化后小麦发育将加快,生育期缩短,春小麦生育期缩短的绝对数和相对数均小于冬小麦。②北方十个站点小麦生产的最适水分条件在不同站点、不同气候情景下都有所不同。最适水分条件变幅在40%～80%。③在不考虑CO₂对小麦影响的情况下,由于热量充足,只要水分条件适宜,未来我国北方干旱、半干旱地区小麦产量整体都有增产趋势。如果考虑CO₂,增产效果更加明显。     

播期和种植密度对强、中筋冬小麦蛋白质组分及品质性状的影响

采取裂裂区试验设计,研究了播期和种植密度对强筋小麦临优145和中筋小麦临优2018蛋白质组分和品质性状的影响.结果表明：适期播种的小麦籽粒蛋白质含量和蛋白质产量均最高;推迟播期,强筋品种的醇溶蛋白和谷蛋白含量明显增加,而中筋品种变化不明显;强筋品种的品质性状受播期影响程度高于中筋品种.适期播种,小麦籽粒蛋白质、麦谷蛋白与湿面筋、沉降值、稳定时间、软化度和评价值呈显著或极显著正相关;推迟播期,醇溶蛋白与湿面筋含量呈显著正相关.播期变化引起的蛋白质各组分所占比例的改变是改善小麦品质性状的重要原因.在试验种植密度范围内（225万株·hm-2、300万株·hm^-2和375万株·hm^-2）,小麦籽粒蛋白质含量变化不明显,密度对强筋品种的品质性状有一定调节作用;在低密度条件下（225万株·hm^-2）中筋品种的品质性状最佳.     

籼型黑米粒形性状的遗传效应及其与矿质元素含量的遗传相关性

采用禾谷类作物种子数量性状的遗传模型,分析了灿型黑米稻品种双列杂交F1和F2种子的粒重,粒长,粒宽和粒长／粒宽等粒形性状的遗传效应及其与米粒中矿质元素Fe,Zn,Mn和P含量的遗传相关性,结果表明：4种粒形性状同时受制于种子直接遗传效应,母体效应和细胞质作用影响,其中种子直接遗传效应比母体效应和细胞质效应的作用更大,且种子直接遗传效应以加性效应占主导,粒重,粒宽和粒长／粒宽的种子直接遗传率较高,杂种早代单粒选择效果较好,粒长的种子直接遗传率和母体遗传率均属中等,较高世代的杂种进行单株选择和单粒选择均有一定效果,4种粒形性状与其米粒中矿质元素Fe,Zn,Mn和P含量表现较强的种子直接加性相关,直接显性相关,细胞质相关,母体加性相关和母体显性相关,在特种稻育种实践中,可以通过粒形性的间接选择,达到改良其矿质元素含量等营养品质性状的目标。