开放系统中农作物对空气CO2浓度增加的响应

FACE试验(free-air CO₂ enrichment)开展的10多年中,供试农作物主要有:C₃禾本科作物小麦(Triticum aestivum L.)、多年生黑麦草(Lolium perenne)和水稻(Oryza sativa L.),C₄禾本科类高粱(Sorghum bicolor(L.)Mench),C₃豆科植物白三叶草(Trifolium repens),C₃非禾本科块茎状作物马铃薯(Solanum tuberosum L.),以及多年生C₃类木本作物棉花(Gossypium hirsutumL.)和葡萄(Vitis viniferaL.).本文系统整理和分析了以下各项参数的结果:光合作用、气孔导度、冠层温度、水分利用、水势、叶面积指数、根茎生物量累积、作物产量、辐射利用率、比叶面积、N含量、N收益、碳水化合物含量、物候变化、土壤微生物、土壤呼吸、痕量气体交换以及土壤碳固定.CO₂浓度升高对农作物的影响作用主要表现在以下方面:(1)促进了植物光合作用、增加了其生物量累积;(2)显著提高C₃作物产量,但对C₄作物产量的影响很小;(3)降低了C₃和C₄作物气孔导度,非常显著地提高了所有作物的水分利用率;(4)对植物生长的促进作用在水分不足与水分充足时二者相当或前者大于后者;(5)对非豆科植物生长的促进作用要受到土壤低N水平限制,而对豆科植物则不受氮肥水平限制;(6)对根系生长的促进作用要大于地上部分;(7)对多年     

28种园林植物对大气CO2浓度增加的生理生态反应

通过对28种园林植物在不同CO2浓度水平下的气体交换参数的观测,分析了净光合速率、气孔导度、蒸腾速率和水分利用效率等生理生态指标的变化趋势与规律.结果表明,所测植物净光合速率和水分利用效率随CO2浓度升高而线性增加,但不同植物种类对高CO2浓度的反应存在较大差异.气孔导度和蒸腾速率与CO2浓度呈线性负相关关系.当CO2浓度倍增(350～700 μmol·mol-1)时,28种园林植物净光合速率平均提高31.2%,气孔导度降低16.5%,蒸腾速率下降11.7%,而水分利用效率则提高了49.2%.不同光合途径的植物净光合速率和水分利用效率受CO2浓度增加的影响程度为C3植物较大,C4植物较小, CAM植物介于两者之间.对不同生活型植物而言,影响程度则为草本C3植物较大,乔木C3植物较小,灌木C3植物居于两者之间.     

狗尾草蒸腾特性与水分利用效率对模拟光辐射增强和CO2浓度升高的响应

 在人工控制光照强度和CO2浓度条件下,测量了禾本科C4植物狗尾草（Setaria viridis）的光合速率（Pn）,蒸腾速率（Tr）,胞间CO2浓度（Ci）,气孔导度（Gs）和叶面饱和水汽压亏缺（Vpdl）对不同模拟光辐射（SPR）强度与CO2浓度的响应。结果表明：Pn, Tr 及Gs均随SPR的升高而增大,增幅趋缓,最终趋于动态平衡。SPR增强的起始阶段,水分利用率（WUE）逐渐增大,在SPR为1200 μmol·m-2·s-1时达到最大值,然后逐渐降低。Ci与Vpdl则随SPR的增强而减小,SPR高于600 μmol·m-2·s-1之后,两者均达到平衡状态。CO2浓度从300增至600 μmol·mol-1的过程中,狗尾草Pn逐渐增大,从600增至1 000 μmol·mol-1过程中,其Pn逐渐降低。Ci、Vpdl和WUE随CO2浓度的升高而增大,Gs和Tr则随CO2浓度的升高而减小。即禾本科一年生C4植物的光合作用对CO2浓度升高响应不敏感,水分蒸腾消耗的减少和WUE的提高对CO2浓度升高的响应极显著。可见,CO2浓度升高对C4植物光合作用的直接促进作用有限,但是却能从提高现有水分利用效率途径促进植物的第一性生产。     

C4作物FACE(free-air CO2 enrichment)研究进展

持续迅速上升的大气二氧化碳浓度([CO₂])是全球变暖最大的驱动因子,但其作为光合作用底物直接增加了作物的生产力。相比C₃作物,人们对未来高浓度CO₂情形下C₄作物的响应规律认识较少。与封闭或半封闭气室研究相比,FACE(free-air CO₂ enrichment)试验在空气自由流动的大田条件下对作物表现进行研究,它提供了对未来作物生长环境的真实模拟,因此提供了评估CO₂肥料效应以及揭示植物响应机制的最好机会。作为人类重要的粮食和饲料来源,高粱和玉米是最重要的C₄作物。在简介美国玉米和高粱FACE系统的基础上,综述了FACE情形下高浓度CO₂(模拟本世纪中叶大气CO₂浓度,即550 μmol/mol)对两大作物生理、生长和产量以及土壤特性等方面的影响,同时比较了与气室研究结果的异同点。(1)FACE使干旱条件下两作物光合作用显著增强,但湿润条件下没有影响;FACE条件下高粱出现光合适应现象,而玉米没有;(2)FACE使两作物气孔导度大幅下降,导致叶温升高、蒸腾速率下降、蒸发蒸腾总量减少或没有变化、叶片总水势和水分利用效率增加或没有变化;(3)FACE对两作物物候期和化学组分影响很少;(4)FACE使干旱条件下两作物生长和产量略有增加,但湿润条件下没有影响;(5)FACE使高粱田土壤丛枝状菌根真菌的长度和易提取胶状物质浓度显著增加,导致水稳性土壤团聚体增加;FACE对高粱田N₂O或含氮气体(N₂O+N₂)的排放没有影响;(6)高浓度CO₂对两作物气孔导度的影响FACE试验明显大于气室试验,而对生长和产量的影响呈相反趋势。阐明CO₂与基因型、土壤湿度和大气温度间的互作效应及其机制是下一轮C₄作物FACE研究优先考虑的方向,技术的不断进步已为利用大型FACE系统来研究这些互作效应提供了可能。     

开放式空气CO2浓度增高条件下C3作物(水稻)与C4杂草(稗草)的竞争关系

通过田间试验,研究了FACE（开放式空气CO2浓度升高）条件下C3作物水稻（Oryza sativa）和C4杂草稗草（Echinochloa crusgalli）的生长和竞争关系,结果表明,FACE条件下C3植物水稻生物量和产量增加,吉片数增加,分蘖数增加,叶面积系数（LAI）增大;而C4植物稗草相反,FACE条件下水稻和稗草中面积均减少,而净同化率（NAR）均增加;FACE条件下水稻-稗草比例为1：1时,水稻与稗草的生物量比率、产量比率、LAI比率、茎蘖比率和NAR比率均增加,水稻-稗草的竞争关系发生变化,水稻（C3植物）竞争能力增加,稗草（C4植物）竞争能力下降。     

不同尺度上植物叶气孔导度对升高CO2的响应

植物叶气孔导度对大气CO2浓度升高的响应可表现在以下几个层面：在叶水平上,叶气孔导度和气孔密度下降;在植物个体水平上,单位叶面积蒸腾下降,植株的水分利用率升高;在生态系统水平上,蒸散降低,土壤泾流和土壤水分含量增加;在全球尺度上,扩大了温室气体的增温效应,同时也降低了全球降雨量增加的趋势。正是因为植物叶气孔导度的变化会影响全球水循环,所以它在全球变化中起着非常重要的作用。但目前的研究结果还不能外推到更大的尺度上去。     

水稻叶片光合作用对开放式空气CO2浓度增高(FACE)的响应与适应

利用便携式光合气体分析系统 (LI 6 4 0 0 ) ,比较测定了高CO2 浓度 (FACE ,free airCO2 enrich ment)和普通空气CO2 浓度下生长的水稻叶片的净光合速率、水分利用率、表观量子效率和RuBP羧化效率等光合参数 .在各自生长CO2 浓度 (380vs 5 80 μmol·mol-1)下测定时 ,高CO2 浓度 (5 80 μmol·mol-1)下生长的水稻叶片的净光合速率、碳同化的表观量子效率和水分利用率明显高于普通空气 (380 μmol·mol-1)下生长的水稻叶片 .但是 ,随着FACE处理时间的延长 ,高CO2 浓度对净光合速率的促进作用逐渐减小 .在相同CO2 浓度下测定时 ,FACE条件下生长的水稻叶片净光合速率和羧化效率明显比普通空气下生长的对照低 .尽管高CO2 浓度下生长的水稻叶片的气孔导度明显低于普通空气中生长的水稻叶片 ,但两者胞间CO2 浓度差异不显著 ,因此高CO2 浓度下生长的水稻叶片光合下调似乎不是由气孔导度降低造成的 .     

羊草叶片气孔导度对环境因子的响应模拟

准确定量描述植物气孔对环境的响应是了解植物光合作用机理、预测植物生产力及其大气－植被－土壤系统中水分和热量交换的关键。利用松嫩平原盐碱化草地羊草光合生理特征的野外观测数据,分析了羊草叶片气孔导度对环境因子的反应,结果表明：羊草叶片气孔导度对环境因子变化敏感,尤其对瞬时光合有效辐射（ＰＡＲ）、叶片与空气间的水汽压亏损（ＶＰＤ）和空气温度（Ｔａ）反应十分明显。依据野外实测资料对国际上两类代表性气孔导度     

土壤水分变化对长白山主要树种蒙古栎幼树生长的影响

选择长白山红松阔叶林主要优势树种蒙古栎为研究对象,人工控制3种施水量研究蒙古栎幼树形态、生物量效应和光合生理特征对土壤含水量变化的响应．结果表明,不同土壤含水量变化显著影响蒙古栎叶片、枝、根的生物量及其分配格局和叶片光合气体交换特征．水分胁迫改变幼树树冠结构,抑制幼树树高、地径、叶片大小、地上和地下生物量;同时,蒙古栎幼树根冠生物量比随着土壤水分含量的减少显著提高;供水量减少对幼树净光合速率、CO2利用率和碳利用率等特征有显著的负向影响;而叶片气孔导度、蒸腾速率和水分利用率对不同土壤含水量反应较复杂,只在土壤含水量较低时,幼树气孔导度、蒸腾速率明显降低,叶片水分利用率升高,表现出蒙古栎树种是干旱可变植物,长期水分胁迫可提高树种的耐旱能力．     

干旱生境中接种丛枝菌根真菌对三叶鬼针草

为阐明丛枝菌根真菌对石灰岩地区适生植物三叶鬼针草(Bidens pilosa L.)光合作用的影响,设置正常浇水(A)、中度干旱胁迫(B)和重度干旱胁迫(C)3个水分处理梯度,比较了不同水分处理条件下接种丛枝菌根真菌Glomus mosseae和未接种三叶鬼针草之间净光合速率、气孔导度、蒸腾速率、胞间CO2浓度、羧化效率、水分利用效率等特征的差异.结果表明,水分胁迫显著降低三叶鬼针草的净光合速率、气孔导度、蒸腾速率和羧化效率.胞间CO2浓度在处理的前期(7d)因干旱胁迫而降低,在后期随土壤含水量的降低而升高;水分利用效率则是中度胁迫的植株、正常浇水处理植株、重度胁迫植株依次降低.在正常浇水条件下接种G. mosseae 对三叶鬼针草光合参数没有产生显著性影响;在中度胁迫条件下,接种植株较未接种植株在水分处理的前28d有更高的净光合速率、气孔导度、蒸腾速率和羧化效率;在重度胁迫条件下,虽然净光合速率、气孔导度、蒸腾速率和羧化效率接种植株高于未接种植株,但是二者并不显著.研究认为,干旱胁迫对三叶鬼针草光合作用的影响在水分处理的前期表现为气孔因素制约,在后期则主要是非气孔因素的影响;在正常浇水条件下接种G. mosseae 对三叶鬼针草的光合作用没有显著性影响,在干旱胁迫条件下,丛枝菌根真菌通过改善三叶鬼针草气孔导度和羧化效率等减弱干旱胁迫对植株的伤害,但是这种保护作用因为土壤水分的严重匮乏以及土壤干旱的时间延长而受到限制.     

植物对开放式CO2 浓度增高(FACE)的响应与适应研究进展

开放式CO2浓度增高(FACE)系统是近年研究植物对高CO2浓度响应和适应的新手段,它比以往密闭和半密闭系统对实验植物生长环境的干扰少.利用FACE系统进行研究更有助于正确地预测未来大气CO2浓度增高对植物的影响.该文结合作者的研究工作简要评介了FACE系统与以往密闭和半密闭式CO2浓度增高实验系统的不同之处以及近年来利用FACE系统所作的最新研究进展.     

The response of photosynthesis and stomatal conductance to rising [CO2]: mechanisms and environmental interactions

This review summarizes current understanding of the mechanisms that underlie the response of photosynthesis and stomatal conductance to elevated carbon dioxide concentration ([CO2]), and examines how downstream processes and environmental constraints modulate these two fundamental responses. The results from free-air CO2 enrichment (FACE) experiments were summarized via meta-analysis to quantify the mean responses of stomatal and photosynthetic parameters to elevated [CO2]. Elevation of [CO2] in FACE experiments reduced stomatal conductance by 22%, yet, this reduction was not associated with a similar change in stomatal density. Elevated [CO2] stimulated light-saturated photosynthesis (Asat) in C3 plants grown in FACE by an average of 31%. However, the magnitude of the increase in Asat varied with functional group and environment. Functional groups with ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco)-limited photosynthesis at elevated [CO2] had greater potential for increases in Asat than those where photosynthesis became ribulose-1,5-bisphosphate (RubP)-limited at elevated [CO2]. Both nitrogen supply and sink capacity modulated the response of photosynthesis to elevated [CO2] through their impact on the acclimation of carboxylation capacity. Increased understanding of the molecular and biochemical mechanisms by which plants respond to elevated [CO2], and the feedback of environmental factors upon them, will improve our ability to predict ecosystem responses to rising [CO2] and increase our potential to adapt crops and managed ecosystems to future atmospheric [CO2].     

What have we learned from 15 years of free-air CO2 enrichment (FACE)? A meta-analytic review of the responses of photosynthesis, canopy properties and plant production to rising CO2

Free-air CO(2) enrichment (FACE) experiments allow study of the effects of elevated [CO(2)] on plants and ecosystems grown under natural conditions without enclosure. Data from 120 primary, peer-reviewed articles describing physiology and production in the 12 large-scale FACE experiments (475-600 ppm) were collected and summarized using meta-analytic techniques. The results confirm some results from previous chamber experiments: light-saturated carbon uptake, diurnal C assimilation, growth and above-ground production increased, while specific leaf area and stomatal conductance decreased in elevated [CO(2)]. There were differences in FACE. Trees were more responsive than herbaceous species to elevated [CO(2)]. Grain crop yields increased far less than anticipated from prior enclosure studies. The broad direction of change in photosynthesis and production in elevated [CO(2)] may be similar in FACE and enclosure studies, but there are major quantitative differences: trees were more responsive than other functional types; C(4) species showed little response; and the reduction in plant nitrogen was small and largely accounted for by decreased Rubisco. The results from this review may provide the most plausible estimates of how plants in their native environments and field-grown crops will respond to rising atmospheric [CO(2)]; but even with FACE there are limitations, which are also discussed.     

Mechanisms underlying the amelioration of O3-induced damage by elevated atmospheric concentrations of CO2

There is growing evidence that rising atmospheric CO2 concentrations will reduce or prevent reductions in the growth and productivity of C3 crops attributable to ozone (O3) pollution. In this study, the role of pollutant exclusion in mediating this response was investigated through growth chamber-based investigations on leaves 4 and 7 of spring wheat (Triticum aestivum cv. Hanno). In the core experiments, plants were raised at two atmospheric CO2 concentrations (ambient [350 micro l l(-1)] or elevated CO2 [700 micro l l(-1)] under two O3 regimes (charcoal/Purafil-filtered air [<5 nl l(-1) O3] or ozone-enriched air [75 nl l(-1) 7 h d(-1)]). A subsequent experiment used an additional O3 treatment where the goal was to achieve equivalent daily O3 uptake over the life-span of leaves 4 and 7 under ambient and CO2-enriched conditions, through daily adjustment of exposures based on measured shifts in stomatal conductance. Plant growth and net CO2 assimilation were stimulated by CO2-enrichment and reduced by exposure to O3. However, the impacts of O3 decreased with plant age (i.e. leaf 7 was more resistant to O3 injury than leaf 4); a finding consistent with ontogenic shifts in the tolerance of plant tissue and/or acclimation to O3-induced oxidative stress. In the combined treatment, elevated CO2 protected against the adverse effects of O3 and reduced cumulative O3 uptake (calculated from measurements of stomatal conductance) by c. 10% and 35% over the life-span of leaves 4 and 7, respectively. Analysis of the relationship between O3 uptake and the decline in the maximum in vivo rate of Rubisco carboxylation (Vcmax) revealed the protection afforded by CO2-enrichment to be due, to a large extent, to the exclusion of the pollutant from the leaf interior (as a consequence of the decline in stomatal conductance triggered by CO2-enrichment), but there was evidence (especially from flux-response relationships constructed for leaf 4) that CO2-enrichment resulted in additional effects that alleviated the impacts of ozone-induced oxidative stress on photosynthesis.     

冬小麦旺盛生长期间CO2浓度升高对根际呼吸的影响

依托FACE（free air carbon dioxide enrichment）技术平台,利用阻断根法,采用H6400红外气体分析仪（IRGA）-田间原位测定的方法,研究了大气CO2浓度升高和不同氮肥水平对水稻／小麦轮作制中冬小麦旺盛生长期间根际呼吸的影响。结果表明,在整个测定期间,大气CO2浓度升高增强了根际呼吸速率,提高了根际呼吸排放量。在高N和低N处理中,高CO2浓度下的根际呼吸总排放量分别比Ambient极显著增加117．0％和90．8％。根际呼吸速率在孕穗初期达到最大值;使根际呼吸在土壤呼吸中的比重由24．5％（LN）～26．7（HN）提高到39．8％（LN）～47．1％（HN）。CO2浓度升高与氮肥用量对根际呼吸产生交互效应。表明大气CO2浓度升高将加快土壤向大气的CO2排放,结果将有助于评价未来高CO2浓度背景下农田生态系统土壤碳的固定潜力。     

开放式空气CO2浓度增高对水稻冠层微气候的影响

利用位于江苏省无锡市安镇的我国唯一的农田开放式空气CO2 浓度增高 (FACE)系统平台 ,于2 0 0 1年 8月 2 6日至 10月 13日 (水稻抽穗至成熟期 )进行水稻作物冠层微气候连续观测 ,以研究FACE对水稻冠层微气候特征的影响 .结果表明 ,FACE降低了水稻叶片的气孔导度 ,FACE与对照水稻叶片气孔导度的差异上层叶片大于下层叶片 ,生长前期大于生长后期 .FACE使白天水稻冠层和叶片温度升高 ,这种差异生长前期大于生长后期 ;但FACE对夜间水稻冠层温度的影响不明显 .在水稻旺盛生长的抽穗开花期 ,晴天正午前后FACE水稻冠层温度比对照高 1.2℃ ;从开花至成熟期 ,FACE水稻冠层白天平均温度比对照高 0 .4 3℃ .FACE对冠层空气温度也有影响 ,白天水稻冠层空气温度FACE高于对照 ,这种差异随太阳辐射增强而增大且冠层中部大于冠层顶部 ;冠层中部空气温度FACE与对照的差异 (Tface-Tambient)日最大值在 0 .4 7～ 1.2℃之间 ,而冠层顶部的Tface-Tambient日最大值在 0 .37～ 0 .8℃之间 .夜间水稻冠层空气温度FACE与对照差别不大 ,变化在± 0 .3℃之内 .而FACE对水稻冠层空气湿度无显著影响 ,表明FACE使水稻叶片气孔导度降低 ,从而削弱了植株的蒸腾降温作用 ,导致水稻冠层温度和冠层空气温度升高 ,改变了整个水稻冠层的温度环     

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.     

Plant water relations at elevated CO2 -- implications for water-limited environments

Long-term exposure of plants to elevated [CO2] leads to a number of growth and physiological effects, many of which are interpreted in the context of ameliorating the negative impacts of drought. However, despite considerable study, a clear picture in terms of the influence of elevated [CO2] on plant water relations and the role that these effects play in determining the response of plants to elevated [CO2] under water-limited conditions has been slow to emerge. In this paper, four areas of research are examined that represent critical, yet uncertain, themes related to the response of plants to elevated [CO2] and drought. These include (1) fine-root proliferation and implications for whole-plant water uptake; (2) enhanced water-use efficiency and consequences for drought tolerance; (3) reductions in stomatal conductance and impacts on leaf water potential; and (4) solute accumulation, osmotic adjustment and dehydration tolerance of leaves. A survey of the literature indicates that the growth of plants at elevated [CO2] can lead to conditions whereby plants maintain higher (less negative) leaf water potentials. The mechanisms that contribute to this effect are not fully known, although CO2-induced reductions in stomatal conductance, increases in whole-plant hydraulic conductance and osmotic adjustment may be important. Less understood are the interactive effects of elevated [CO2] and drought on fine-root production and water-use efficiency, and the contribution of these processes to plant growth in water-limited environments. Increases in water-use efficiency and reductions in water use can contribute to enhanced soil water content under elevated [CO2]. Herbaceous crops and grasslands are most responsive in this regard. The conservation of soil water at elevated [CO2] in other systems has been less studied, but in terms of maintaining growth or carbon gain during drought, the benefits of CO2-induced improvements in soil water content appear relatively minor. Nonetheless, because even small effects of elevated [CO2] on plant and soil water relations can have important implications for ecosystems, we conclude that this area of research deserves continued investigation. Future studies that focus on cellular mechanisms of plant response to elevated [CO2] and drought are needed, as are whole-plant investigations that emphasize the integration of processes throughout the soil--plant--atmosphere continuum. We suggest that the hydraulic principles that govern water transport provide an integrating framework that would allow CO2-induced changes in stomatal conductance, leaf water potential, root growth and other processes to be uniquely evaluated within the context of whole-plant hydraulic conductance and water transport efficiency.     

开放式空气中CO_2浓度增高(FACE)对水稻生长和发育的影响

人类活动导致的大气和气候变化将极大地改变作物的生长环境,其中最大的一个变化就是大气二氧化碳(CO2)浓度的迅速上升:从工业革命前的平均270μmol/mol上升到目前的381μmol/mol,到2050年至少超过550μmol/mol。FACE(Free-air CO2 enrichment,开放式空气中CO2浓度增高)试验是目前评估未来高浓度CO2对作物生长和产量实际影响的最佳方法。水稻无疑是人类最重要的食物来源,迄今为止人类利用FACE技术开展水稻响应和适应的研究已有10a(19982008年)的历史。以生长发育为主线,首次系统综述了10a水稻FACE试验在该领域的研究成果,总结了FACE情形下高浓度CO2(模拟本世纪中叶大气CO2浓度)对主要供试水稻品种(小区面积大于4m2)光合作用、生育进程、地上部生长、地下部生长、物质分配、籽粒灌浆、产量构成以及倒伏性状等影响的研究进展,比较了FACE与非FACE研究之间以及中国和日本FACE研究(世界上唯一的两个大型水稻FACE研究)之间的异同点。根据研究进展以及当前的技术水平,文章最后提出了该领域的3个优先课题:(1)FACE情形下杂交稻生产力响应高于预期的生物学机制;(2)FACE情形下CO2与主要栽培措施的互作效应;(3)FACE情形下CO2与主要空气污染物臭氧的互作效应。这些响应的机理性解析将有助于从根本上减少人类预测未来粮食安全的不确定性,进而更加有效地制订出应对全球变化的适应策略。