[CO2]升高对粮食作物影响的研究进展

自工业革命以来,由人类活动引起的大气CO_2浓度([CO_2])不断攀升,正驱动着全球气候变化,对全球农业产生重大影响。本文归纳总结了目前作物对高[CO_2]响应的主要研究技术手段,以及作物对高[CO_2]响应的机理研究,并进一步梳理了当前全球关于[CO_2]升高对作物产量和营养品质影响的研究。结果表明:相比封闭式或半封闭式环境控制试验系统,开放式试验系统(如开放式CO_2控制系统FACE)由于其能更加真实地模拟自然条件下作物对未来高[CO_2]的响应和适应情况,被公认为是目前研究作物对高[CO_2]响应的最理想手段。[CO_2]增高会增加C3作物光合速率、生物量和产量,在一定程度上缓解气候变化对农作物产生的负面影响,但是作物对大气[CO_2]的升高存在光合适应现象,当作物长期暴露在高[CO_2]条件下时,高[CO_2]对作物的促进作用会逐渐减缓。近10年的FACE试验发现,对高[CO_2]出现高应答的水稻品种,其光合速率和产量在高[CO_2]下的增加幅度比早期的主要粮食作物FACE试验结果平均高出两倍。此外,高[CO_2]会明显降低大部分非豆科C3作物中蛋白质和矿物质(如锌、铁)以及部分维生素的含量,加剧目前全球约2亿人由于维生素和矿物质元素等营养缺乏导致的健康问题。如何充分利用未来高[CO_2]实现高增产的同时,减缓粮食养分下降的负面影响,是迫切需要解决的科学问题。     

A possible mechanism of mineral responses to elevated atmospheric CO2 in rice grains

Increasing attentions have been paid to mineral concentration decrease in milled rice grains caused by CO2 enrichment, but the mechanisms still remain unclear. Therefore, mineral(Ca, Mg, Fe, Zn and Mn) translocation in plant-soil system with a FACE(Free-air CO2 enrichment) experiment were investigated in Eastern China after 4-yr operation. Results mainly showed that:(1) elevated CO2 significantly increased the biomass of stem and panicle by 21.9 and 24.0%, respectively, but did not affect the leaf biomass.(2) Elevated CO2 significantly increased the contents of Ca, Mg, Fe, Zn, and Mn in panicle by 61.2, 28.9, 87.0, 36.7, and 66.0%, respectively, and in stem by 13.2, 21.3, 47.2, 91.8, and 25.2%, respectively, but did not affect them in leaf.(3) Elevated CO2 had positive effects on the weight ratio of mineral/biomass in stem and panicle. Our results suggest that elevated CO2 can favor the translocation of Ca, Mg, Fe, Zn, and Mn from soil to stem and panicle. The CO2-led mineral decline in milled rice grains may mainly attribute to the CO2-led unbalanced stimulations on the translocations of minerals and carbohydrates from vegetative parts(e.g., leaf, stem, branch and husk) to the grains.     

Intra-specific variation of wheat grain quality in response to elevated [CO2] at two sowing times under rain-fed and irrigation treatments

In order to investigate the intra-specific variation of wheat grain quality response to elevated atmospheric CO₂ concentration (e[CO₂]), eight wheat (Triticum aestivum L.)cultivars were grown at two CO₂ concentrations ([CO₂]) (current atmospheric, 389 CO₂ μmol mol⁻¹vs. e[CO₂], FACE (Free-Air CO₂ Enrichment), 550  ±  10% CO₂ μmol mol⁻¹), at two water levels (rain-fed vs. irrigated) and at two times of sowing (TOS₁, vs. TOS₂). The TOS treatment was mainly imposed to understand whether e[CO₂] could modify the effects of timing of higher grain filling temperatures on grain quality. When plants were grown at TOS₁, TKW (thousand kernel weight), grain test weight, hardness index, P, Ca, Na and phytate were not significantly changed under e[CO₂]. On the other hand, e[CO₂] increased TKW (16%), hardness index (9%), kernel diameter (6%), test weight (2%) but decreased grain protein (10%) and grain phytate (11%) at TOS₂. In regard to grain Zn, Mn and Cu concentrations and some flour rheological properties, cultivar specific responses to e[CO₂] were observed at both sowing times. Observed genetic variability in response to e[CO₂] in terms of grain minerals and flour rheological properties could be easily incorporated into future wheat breeding programs to enable adaptation to climate change.     

Changing climate and potential impacts on potato yield and quality ‘CHIP’: introduction, aims and methodology

Climate change effects caused by an increasing concentration of CO₂ and ozone represent an issue of major concern both for scientists and policy-makers. In a concerted program funded by the Commission of the European Union, a European network of experiments (in open-top chambers (OTC), and free air carbon dioxide enrichment systems (FACE)) and modelling was carried out to investigate the effects of increasing atmospheric CO₂ and ozone concentrations, under different climatic conditions, on potato (Solanum tuberosum L. cv. Bintje). This contribution describes the experimental network and the standard protocol set-up for the assessments that served to improve and to validate process oriented potato growth simulation models leading to scenarios of future productivity of potato in Europe.     

FACE对水稻产量形成的影响及其原因分析

2001～2003年,在大田条件下以武香粳14号为供试品种,设计高、中、低氮处理,研究开放式空气CO2浓度增加(FACE)200μmolmol-1对水稻产量形成的影响及其原因。结果表明,FACE处理使水稻产量平均比对照增加12.8%,其中2001、2002和2003年增幅分别为10.8、14.1和13.6%,高、中、低氮处理的增幅分别为17.6、12.4、10.9%,均达到显著或极显著水平;FACE和氮处理对水稻产量的互作效应,2001、2002年达到显著水平,产量最高的处理组合为FACE×NN;FACE处理使水稻穗数平均比对照增加18.8%,每穗颖花数平均比对照减少7.6%,均达极显著水平;FACE处理使水稻生物产量平均提高16.2%,达极显著水平,使经济系数下降但未达到显著水平。进一步研究表明,FACE处理使水稻穗数极显著多于对照是因其分蘖发生速度快,最高分蘖数多所致,而不是其分蘖成穗率高的缘故;FACE处理使水稻每穗颖花数极显著少于对照是因其分化颖花的大量退化所致,而不是分化颖花数少的缘故。     

FACE环境下不同秸秆与氮肥管理对稻田土壤硝化和反硝化菌的影响

利用位于江都市小记镇的中国稻-麦轮作FACE平台,采用最大可能(MPN)法,在2004年水稻生长季研究了不同施肥情况(施常规N量UN和低N量LN)、不同秸秆还田情况(秸秆全还田HR和秸秆不还田NR)下,土壤中硝化和反硝化细菌数量在FACE条件下随时间的变化。结果表明,FACE条件下,土壤硝化菌数普遍在抽穗期或乳熟期达到最大值,而对照土壤的硝化菌数普遍到成熟期才达到最大值,并且显著高于FACE处理的相应值(P<0.05)。在HR条件下,LN和UN小区FACE处理的土壤硝化细菌数量较对照减少6%~10%。FACE条件LN小区的反硝化菌数在成熟期达到最大值,而对照处理则在乳熟期达到最大值,并显著高于FACE处理(P<0.05);而UN小区的反硝化菌数二者均在抽穗期达到最大值。在LN小区HR和NR情况下,FACE处理土壤反硝化细菌数量分别低于对照处理的相应值8%和13%。在HR情况下,土壤反硝化潜势FACE处理显著低于对照。在LN和UN小区,FACE处理土壤的反硝化作用潜势分别是对照的83.7%和95.4%。     

杂交水稻光合和荧光特性对高CO2浓度的动态响应

为探究大气二氧化碳 (CO₂) 浓度增高对杂交水稻光合和荧光特性的动态影响,利用开放式空气中CO₂浓度增高(FACE)系统,研究高CO₂浓度对大田水稻不同生育期叶片光合和荧光参数的动态影响及其种间差异。以高产杂交组合甬优2640和Y两优2号为试验材料,设置环境CO₂和高CO₂浓度2个水平,测定2个品种气体交换和荧光参数。结果表明,大气CO₂浓度升高使2个品种平均叶片净光合速率(Pn)在移栽后30、59、76、91和108 d分别增加25%、16%、25%、18%和8%,除移栽后108 d外,均达极显著水平(P<0.01);甬优2640移栽后76和91 d叶片Pn对CO₂的响应明显大于Y两优2号。除灌浆末期外,CO₂浓度升高使2个品种叶片气孔导度(Gs)和蒸腾速率(Tr)均呈下降趋势,且Y两优2号的降幅多大于甬优2640。CO₂熏蒸下叶片气孔部分关闭,但叶片胞间与周围空气中CO₂浓度之比无显著变化,甚至明显增加。大气CO₂浓度升高使2个品种灌浆末期PSⅡ最大光化学量子产量均下降4%,但对其他各期均无显著影响;除灌浆末期外,CO₂浓度升高使PSⅡ实际光量子产量和光化学猝灭均显著增加。两杂交组合叶片光合参数对高CO₂浓度的动态响应多存在种间差异,但荧光参数的响应品种间差异较小。本研究结果为增强气候假定情景下水稻响应的预测能力并制订应对策略提供了依据。     

开放式空气CO2浓度升高对小麦冠层微气候的影响

利用农田开放式CO2 浓度升高 (Free-air CO2 Enrichment,简称FACE)系统平台,于小麦拔节-成熟期进行作物冠层微气候要素的连续观测.结果表明:FACE条件下叶片气孔导度低于对照,倒一、倒二、倒三叶的气孔导度比对照平均分别减少了28%、32%和26%,均达极显著水平(p<0.01);由于叶片气孔导度降低使得蒸腾降温作用减弱,导致白天FACE条件下小麦的冠层温度升高,开花-蜡熟期平均升高0.77℃,抽穗后最高达1.58℃;FACE系统中,白天小麦冠层内部空气温度比对照高0.12～0.98℃(最大差值),冠层顶部空气温度高0.03～0.7℃(最大差值),但冠层空气湿度均低于对照,冠层中部空气湿度最低差值在-0.3～-7个百分点;FACE处理对小麦冠层上方净辐射的影响不大.     

大气CO2浓度升高对不同类型水稻灌浆期有机物合成与分配的影响

为明确不同类型水稻对高浓度CO2的响应差异,应用先进的FACE(Free air CO_2 enrichment)试验平台,以粳稻(Japonica)武运粳21(WYJ21)和籼稻(Indica)扬稻6(YD6)为试材,研究灌浆期两种类型水稻体内有机物含量的变化。与对照相比,WYJ21籽粒中直链和支链淀粉含量在FACE处理下略有增加,YD6籽粒中直链淀粉含量在完熟期显著高于对照8.9%;WYJ21籽粒中α淀粉酶、β淀粉酶活性变化不显著,YD6籽粒中β淀粉酶活性在完熟期显著低于对照28.1%;WYJ21籽粒蔗糖含量在乳熟期高于对照11.5%,YD6籽粒中蔗糖含量在蜡熟期低于对照14.4%,WYJ21和YD6茎中蔗糖含量分别在蜡熟和乳熟期显著下降。FACE下两水稻品种茎中蔗糖合成酶(SS)活性变化自乳熟至完熟期呈现先降后升的趋势,且均在中后期达到显著水平,WYJ21在灌浆前期茎中磷酸蔗糖合成酶(SPS)活性显著增加,YD6茎中SPS活性在灌浆后期显著增加;FACE处理降低了两水稻茎中蛋白质含量,其中WYJ21茎中蛋白质含量在蜡熟期显著下降38.5%,YD6茎中蛋白质含量在蜡熟和完熟期分别下降33.9%和60.5%。CO_2、时期、品种对水稻茎中蔗糖含量、SS和SPS活性的影响达到显著水平;CO_2、时期、品种以及它们的交互作用显著影响直链和支链淀粉含量、α与β淀粉酶活性以及蛋白质含量。上述结果表明,不同类型水稻在不同灌浆时期对高浓度CO_2的响应程度不同,籼稻YD6的响应程度要大于粳稻WYJ21。     

高CO2浓度和剪叶疏花对水稻‘Y两优2号’ 产量形成的影响

利用稻田FACE(Free Air CO_2 Enrichment)平台,以创造世界高产纪录的超级稻组合‘Y两优2号’为试验材料,CO_2处理设环境CO_2浓度[(382.5±2.0)μmol·mol-1]和高CO_2浓度(增200μmol·mol-1)两个水平,齐穗期源库改变设对照、剪除剑叶(剪1叶)、剪除所有功能叶(剪3叶)以及相间剪除一次枝梗(疏花),研究开放条件下高CO_2浓度对不同源库处理水稻产量及其构成因子的影响。结果表明,对没有进行剪叶疏花处理的水稻(即对照)而言,高CO_2浓度使‘Y两优2号’籽粒产量平均增加12%,这主要与每穗颖花数和结实能力均略有增加有关。高CO_2浓度使剪1叶、剪3叶处理水稻的产量分别增加26%和57%,这主要与饱粒率和所有籽粒平均粒重均大幅增加有关。对齐穗期疏花处理水稻而言,高CO_2浓度导致的产量增幅与对照水稻接近。与对照相比,齐穗期剪1叶、剪3叶处理使水稻籽粒产量分别降低17%和52%,均达极显著水平,这主要与饱粒率和所有籽粒平均粒重均显著下降有关;尽管齐穗期疏花处理使水稻结实能力显著增加,但因每穗颖花数减半,产量大幅下降(-29%)。籽粒最终产量对高CO_2浓度的响应与饱粒率和所有籽粒平均粒重的响应呈显著正相关。以上结果表明,水稻齐穗期人为改变源库比例(特别是剪叶)可以改变籽粒结实能力和最终产量对高CO_2浓度的响应。