FACE水稻茎蘖动态模型

借助农田开放式空气CO₂浓度增高（FACE）技术平台,以武香粳14为供试水稻品种,设置不同施N量处理,研究大气CO₂浓度为570 μmol·mol^-1（比对照高200 μmol·mol^-1）的FACE处理对水稻茎蘖动态的影响,并建立了相应的模拟模型：T_t=A₁(1+e^a1-b1t₎-A₂(1+e^a2-b2t)+C×[B₁(1+e^a3-b3t)-B₂(1+e^a4-b4t)]+D.模型以时间为驱动因子,描述了水稻茎蘖数随移栽天数的动态变化过程,对常规及CO₂浓度增加条件下水稻茎蘖的变化均有很好的拟合性.通过不同年份试验数据对模型的检验,预测根均方差（RMSE）最大为44.27个·m^-2,最小为13.96个·m^-2,且相关系数均达到了极显著水平.表明模型的预测程度较高,具有很好的适用性.     

热带季节雨林冠层树种绒毛番龙眼的光合生理生态特性

采用Li-6400便携式光合作用测定仪,对西双版纳热带季节雨林冠层树种绒毛番龙眼成树树冠上、中、下3层叶片进行了测定,分析西双版纳热带季节雨林冠层树木的光合作用.结果表明,绒毛番龙眼成树具有喜光的光合特性,光饱和点较高(1 000～1 500 μmol·m^-2·s^-1),而光补偿点较低(7.7～15.3 μmol·m^-2·s^-1),对光环境有较强的适应和调节能力,光合有效辐射是影响绒毛番龙眼光合日进程的关键因子;12月,叶片处于成熟期,生长良好,光合能力较强,树冠上层净光合速率（P_n）日变化为单峰型,最大净光合速率（A_max）约为8.9 μmol CO₂·m^-2·s^-1;4月处于新老树叶更替期,光合能力下降,树冠上层P_n日变化为双峰型,中午出现“午休”现象,树冠上层A_max约为4.3 μmol CO₂·m^-2·s^-1;7月上、中层叶片P_n为单峰型,下层出现“午休”.如人为使CO₂浓度在短期内迅速升高,则绒毛番龙眼的P_n会增加,而气孔导度和蒸腾速率降低;CO₂浓度从400 μmol·mol^-1升高到800 μmol·mol^-1时,干季水分利用效率（WUE）提高约50%～100％,雨季WUE较低.     

供氮和增温对倍增二氧化碳浓度下荫香叶片光合作用的影响

供给0～0.6 mg N的盆栽荫香(Cinnamomum burmannii)幼树分别生长在倍增CO ₂(+CO₂,731 μmol·mol^-1)和正常空气CO ₂浓度(CO ₂,365 μmol·mol^-1)的生长箱内,昼夜温度分别为25/23 ℃和32/25 ℃,自然光照下生长30 d.以生长在CO₂和25/23 ℃下的植株为对照研究增温和氮对+CO₂叶片光合作用的影响.结果表明,在+CO₂和25/23 ℃下无氮和氮处理植株的平均光合速率(Pnsat)较+CO₂和32/25 ℃下的叶片高5.1%,温度增高降低叶片Pnsat;而Pnsat随供氮而增高.在+CO₂条件下,生长在32/25 ℃下的叶片Rubisco最大羧化速率(Vcmax)和最大电子传递速率(Jmax)较25/23 ℃下的低(P＜0.05),温度增高降低+CO₂下叶片的Vcmax和Jmax在+CO2下叶片光合呼吸速率(Rp)较低,生长温度增高提升Rp.在CO₂下生长温度从25/23 ℃增至32/25 ℃,叶片的Rubisco含量(NR)和Rubisco活化中心浓度(M)降低,而供氮能增高NR和M.供氮能减缓温度增高对倍增CO₂下荫香叶片光合作用的限制.     

大气臭氧浓度升高对水稻叶片膜脂过氧化及保护酶活性的影响

采用开顶式气室（open top chambers,OTCs）装置,以水稻品种“3694繁”(Oryza sativa L.)为材料,研究3种处理：过滤大气（CF,O₃浓度约为20 nl·L^-1）、环境大气（NF,O₃浓度约为40 nl·L^-1）和高浓度O₃（EO,O₃浓度约为75 nl·L^-1）下叶片可溶性蛋白质含量、膜脂过氧化程度与主要保护酶活性的变化.结果表明：过滤大气与环境大气处理之间各个指标无显著差异.与CF处理相比,高浓度O₃处理条件下水稻叶片中可溶性蛋白含量显著下降,而过氧化氢（H₂O₂）和抗坏血酸（ASA）含量显著增加;超氧化物歧化酶（SOD）、过氧化氢酶（CAT）、过氧化物酶（POD）活性升高,最大升高幅度分别为93.7%、39.9%和312.4%;抗坏血酸过氧化物酶（APX）活性呈先升高后下降趋势; 不同O₃处理条件下叶片中丙二醛（MDA)含量无显著变化.表明抗氧化系统可有效抑制水稻叶片中的O₃胁迫引起的膜质过氧化作用,说明水稻品种“3694繁”对O₃胁迫有一定抗性.     

钾素水平对网纹甜瓜叶片光合特性及叶绿体亚显微结构的影响

在设施基质栽培条件下,研究了营养液中120、240及360 mg·L^-1 3个钾素水平对网纹甜瓜‘甜甜1号’叶片光合特性及叶绿体亚显微结构的影响.结果表明,营养液中钾素水平过低（120 mg·L^-1）或过高（360 mg·L^-1）均导致网纹甜瓜叶片净光合速率下降,使叶绿体片层结构混乱、变形和片层数减少,但对CO₂补偿点（70 μl·L^-1）、饱和CO₂（600 μl·L^-1）、光补偿点（50 μmol·m^-2·s^-1）无显著影响.适宜的钾素水平能显著提高叶片的饱和光强、羧化效率和表观量子效率,3个指标分别为1 200 μmol·m^-2·s^-1、0.1364和0.0237.在试验条件下,提高温室内基质栽培网纹甜瓜叶片光合效率的最适钾素水平为240 mg·L^-1.     

二氧化氮对樟树幼苗生长及光合作用的影响

采用开顶式人工熏气装置,对1年生樟树幼苗进行了为期2个月不同体积分数NO₂(0.1、0.5和4.0 μl·L^-1)熏气试验,研究其对幼苗生长及光合作用的影响.结果表明:0.5和0.1 μl·L^-1 NO₂处理促进了樟树幼苗生长,而4.0 μl·L^-1 NO₂处理则抑制其生长.各处理樟树幼苗叶片净光合速率(P_n)日变化呈不对称的双峰型曲线,存在光合“午休”现象;在光合日进程中,0.5 μl·L^-1 NO₂处理使叶片P_n提高,最大值达8.542 μmol CO₂·m^-2s^-1,4.0 μl·L^-1NO₂处理的大多数时段使P_n降低,而0.1 μl·L^-1 NO₂处理对P_n的影响则依时段而不同;0.5和4.0 μl·L^-1 NO₂处理提高了叶片气孔导度(G_s)和胞间CO₂浓度(C_i)的最大值和最小值,0.1 μl·L^-1 NO₂处理提高了C_i的最大值和最小值,降低了G_s的最大值和最小值.熏气处理中、后期,0.5μl·L^-1 NO₂处理叶片的日均净光合速率显著高于其他处理.在熏气处理前期,0.5和4.0 μl·L^-1 NO₂处理使叶片最大PSⅡ的光能转换效率(F_v/F_m)显著下降;在熏气处理后期,4.0 μl·L^-1 NO₂处理的叶片F_v/F_m仍显著低于对照.     

荔枝的光合特性

在低光强(0至0.15m mol m^-2s^-1)下,荔枝叶子的光合速率随光入射量子通量的增高而增大。在光强0.7m mol m^-2s^-1时光合速率为1.76μmol m^-2s^-1。光合作用的光补偿点约为0.02 m mol m^-2s^-1光量子。荔技叶子具有低的气孔对水分传导率。气孔对水分传导率和蒸腾速率在低光强时随入射量子通量增高而增大:而细胞间CO₂浓度随光强增高而下降。在光强高于0.2m mol m^-2 s^-1光量子时,细胞间CO₂浓度变化较少。在低光强时,叶子的水分利用效率（光合/蒸腾）随光强增高而增大。在光强高于0.2m mol m^-2 s^-1光量子时,水分利用效率明显降低。荔枝叶子的最适光合作用叶温为22-26℃。可能表明在华南夏季中午的高温限制荔枝的田间光合作用。外界CO₂浓度增高相应增高细胞间CO₂浓度。当细胞间CO₂浓度约低于230μ1.1^-1时,光合速率随细胞间CO₂浓度增高而增大。在更高的细胞间CO₂浓度对,光合速率变化则较少。荔枝叶子光合速率对叶子/空气水蒸汽压陡度的变化响应不敏感。气孔对水分传导率和细胞间CO₂浓度随叶子/空气水蒸汽压陡度增大略有降低。     

百菌清对土壤氧化亚氮和二氧化碳排放的影响

在25 ℃、60%WHC（最大持水量）的好氧条件下进行14 d的培养试验,研究杀菌剂百菌清在添加水平为0 mg·kg^-1（CK）、5.5 mg·kg^-1（田间施用量,FR）及110 mg·kg^-1(20FR)和220 mg·kg^-1（40FR）时对酸性、中性和碱性土壤中N₂O和CO₂排放的影响.结果表明：百菌清对N₂O和CO₂排放的影响取决于土壤类型和施用浓度.与对照相比,百菌清在20FR和40FR时显著抑制了酸性土壤N₂O的产生与排放;3种施用量均显著促进了中性土壤N₂O的排放,其中FR水平的促进效果最显著;高浓度（20FR和40FR）的百菌清在培养初期抑制了碱性土壤N₂O的排放,而在培养后期显著促进了N₂O的排放.田间用量的百菌清对土壤CO₂排放量没有明显影响;高浓度（20FR和40FR）时显著促进了酸性土壤CO₂的排放,显著抑制了中性和碱性土壤CO₂的排放.     

大气二氧化碳浓度变化对禾谷缢管蚜种群动态的影响

利用开顶式熏气室研究了大气CO₂浓度和土壤水分对禾谷缢管蚜Rhopalosiphum padi (L.)种群动态的影响,并分析了禾谷缢管蚜密度与被处理小麦叶片化学成分的关系。结果表明：（1）禾谷缢管蚜种群密度随CO₂浓度升高而持续增大并与土壤水分密切相关,各CO₂浓度下均以60%田间持水量时的密度最大;（2）CO₂和土壤水分对小麦叶片化学成分有明显的影响,麦叶水分、可溶性蛋白质、可溶性糖、淀粉含量随CO₂浓度和土壤水分含量上升而增加,纤维素含量随CO₂浓度上升而增加、随土壤水分含量上升而降低,单宁、丁布（DIMBOA）含量在CO₂浓度为550 μl/L时最高,但单宁含量随土壤水分上升而增加,丁布含量在60%田间持水量时最低;（3）禾谷缢管蚜密度与叶片水分、可溶性蛋白质、可溶性糖、淀粉含量呈正相关,与丁布、单宁含量呈负相关。结论：在未来的气候条件下,随着CO₂浓度升高禾谷缢管蚜种群可能会持续增长,这种增长在半干旱区更加突出。禾谷缢管蚜种群增长的原因之一是大气CO₂和土壤水分条件改变了植物的化学成分构成。     

在高CO2浓度下生长的小麦对棉铃虫生长发育和繁殖的影响

通过室内饲养实验研究了在高CO₂浓度（738.8±25.7μL/L）中生长的小麦对棉铃虫 Helicoverpa armigera (Hübner)生长发育,繁殖以及营养效应的影响。结果表明： （1）取食高CO₂浓度大气中生长的麦粒的棉铃虫对食料的取食量和粪便排泄量增加,与对照相比,取食量和粪便排泄量分别增加46.3%（P<0.05）和37.8%;（2）大气CO₂浓度增加影响了麦粒中的营养成分的含量,其中,可溶性蛋白、游离氨基酸、葡萄糖和总糖的含量及碳氮比(C∶N)都显著增加,果糖和粗蛋白的含量都显著降低;（3）大气 CO₂浓度升高所导致的麦粒营养成分的变化影响了棉铃虫幼虫的食物利用效率,与对照组相比,棉铃虫幼虫对食物的毛转化率和净转化率分别降低27.2%和25.4%,对食物的相对取食率则显著提高58.8%（P < 00.1）。据此推测,未来高CO₂浓度的大气环境会降低春小麦的营养价值,从而影响棉铃虫的生长发育,加重其对小麦的危害。     

Does nitrogen supply affect the response of wheat (Triticum aestivum cv. Hanno) to the combination of elevated CO(2) and O(3)?

Spring wheat (Triticum aestivum cv. Hanno) was grown at ambient (350 micromol mol(-1)) or elevated CO(2) (700 micromol mol(-1)) in charcoal/Purafil-filtered air (CFA <5 nmol mol(-1)) or ozone (CFA +75 nmol mol(-1) 7 h d(-1)) at three levels of N supply (1.5, 4 and 14 mM NO(-3)), to test the hypothesis that the combined impacts of elevated CO(2) and O(3) on plant growth and photosynthetic capacity are affected by nitrogen availability. Shifts in foliar N content reflected the level of N supplied, and the growth stimulation induced by elevated CO(2) was dependent on the level of N supply. At 60 d after transfer (DAT), elevated CO(2) was found to increase total biomass by 44%, 29%, 12% in plants supplied with 14, 4 and 1.5 mM NO(-3), respectively, and there was no evidence of photosynthetic acclimation to elevated CO(2) across N treatments; the maximum in vivo rate of Rubisco carboxylation (V(cmax)) was similar in plants raised at elevated and ambient CO(2). At 60 DAT, ozone exposure was found to suppress plant relative growth rate (RGR) and net photosynthesis (A) in plants supplied with 14 and 4 mM NO(-3). However, O(3) had no effect on the RGR of plants supplied with 1.5 mM NO(-3) and this effect was accompanied by a reduced impact of the pollutant on A. Elevated CO(2) counteracted the detrimental effects of O(3) (i.e. the same ozone concentration that depressed RGR and A at ambient CO(2) resulted in no significant effects when plants were raised at elevated CO(2)) at all levels of N supply and the effect was associated with a decline in O(3) uptake at the leaf level.     

红豆草与土壤氮含量对大气二氧化碳浓度升高的响应

在封闭的植物培养箱中,通过盆栽实验,研究了红豆草和土壤氮含量对CO₂浓度增加的响应.结果表明,与正常CO₂浓度(355～370 μmol·mol^-1)相比,CO₂浓度升高(700 μmol·mol^-1),植物生物量增加25.1%(P＜0.01),但植物体氮浓度降低25.3%(P＜0.001),植物全氮没有显著的变化.经3个月盆栽实验后,与原始土壤相比,两种CO₂浓度处理土壤全N、NO₃^--N和NH₄⁺-N都有所降低,而土壤微生物氮则显著增加,这可能与植物生长有关.不同CO₂浓度处理土壤NH₄⁺-N浓度基本一致,但在高CO₂浓度下,土壤NO₃^--N浓度显著降低,而微生物生物氮显著增加.对整个土壤-植物系统而言,盆栽实验后,整个系统全氮有少量增加,但变化不显著,特别是在高CO₂浓度条件下,土壤-植物系统全氮最大,这可能与培养材料红豆草为豆科植物,而且在高CO₂浓度下生物量增加,导致氮的固定量增加有关.     

Future CO2 concentrations, though not warmer temperatures, enhance wheat photosynthesis temperature responses

The temperature dependence of C3 photosynthesis is known to vary according to the growth environment. Atmospheric CO2 concentration and temperature are predicted to increase with climate change. To test whether long-term growth in elevated CO2 and temperature modifies photosynthesis temperature response, wheat (Triticum aestivum L.) was grown in ambient CO2 (370 micromol mol(-1)) and elevated CO2 (700 micromol mol(-1)) combined with ambient temperatures and 4 degrees C warmer ones, using temperature gradient chambers in the field. Flag leaf photosynthesis was measured at temperatures ranging from 20 to 35 degrees C and varying CO2 concentrations between ear emergence and anthesis. The maximum rate of carboxylation was determined in vitro in the first year of the experiment and from the photosynthesis-intercellular CO2 response in the second year. With measurement CO2 concentrations of 330 micromol mol(-1) or lower, growth temperature had no effect on flag leaf photosynthesis in plants grown in ambient CO2, while it increased photosynthesis in elevated growth CO2. However, warmer growth temperatures did not modify the response of photosynthesis to measurement temperatures from 20 to 35 degrees C. A central finding of this study was that the increase with temperature in photosynthesis and the photosynthesis temperature optimum were significantly higher in plants grown in elevated rather than ambient CO2. In association with this, growth in elevated CO2 increased the temperature response (activation energy) of the maximum rate of carboxylation. The results provide field evidence that growth under CO2 enrichment enhances the response of Rubisco activity to temperature in wheat.     

不同二氧化碳浓度条件下红松和长白赤松幼苗根际土壤微生物数量研究

以连续5年不同CO₂浓度(开顶箱700μmol·mol^-1、500μmol·mol^-1、对照箱和裸地)处理的长白赤松和红松幼苗为研究对象,在2003年7～9月分别对幼苗根际土壤细菌、真菌、放线菌数量进行比较研究.结果表明,高浓度CO₂处理对长白赤松幼苗根际土壤细菌数量起显著的(P≤0.001)促进作用,对根际真菌和放线菌数量的促进作用却不明显;对红松来说,除8月份700μmol·mol^-1CO₂处理和7月份500μmol·mol^-1CO₂处理之外,在各月份中受高浓度CO₂处理的根际土壤细菌数量均较对照箱和裸地显著增多(P≤0.001),而根际土壤真菌数量变化除9月份(P≤0.001)外均不明显,放线菌数量受高浓度CO₂的影响亦不明显.     

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.     

高浓度二氧化碳处理对红松幼苗土壤水解酶活性的影响

研究了700和500 μmol·mol^-1高浓度CO₂处理的红松幼苗0～10 cm土层土壤蛋白酶、脲酶、淀粉酶、转化酶和磷酸酶活性的变化.结果表明,土壤蛋白酶(除7月)、脲酶、淀粉酶(除7月)和磷酸酶(除9月)活性在高浓度CO₂条件下极显著增加,而转化酶(除9月)活性却极显著降低.不同高浓度CO₂对酶活性的影响程度不同,500 μmol·mol^-1浓度CO₂处理对蛋白酶和磷酸酶活性的影响较700 μmol·mol^-1处理明显,而700 μmol·mol^-1浓度CO₂处理对脲酶、淀粉酶和转化酶活性的影响较500 μmol·mol^-1显著.     

A whole-tree chamber system for examining tree-level physiological responses of field-grown trees to environmental variation and climate change

A whole-tree chamber (WTC) system was installed at Flakaliden in northern Sweden to examine the long-term physiological responses of field-grown 40-year-old Norway spruce trees [Picea abies (L.) Karst.] to climate change. The WTCs were designed as large cuvettes to allow the net tree-level CO(2) and water fluxes to be measured on a continuous basis. A total of 12 WTCs were used to impose combinations of atmospheric carbon dioxide concentration, [CO(2)], and air temperature treatments. The air inside the ambient and elevated [CO(2)] WTCs was maintained at 365 and 700 micromol mol(-1), respectively. The air temperature inside the ambient temperature WTCs tracked air temperature outside the WTCs. Elevated temperatures were altered on a monthly time-step and ranged between +2.8 and +5.6 degrees C above ambient temperature. The system allowed continuous, long-term measurement of whole-tree photosynthesis, night-time respiration and transpiration. The performance of the WTCs was assessed using winter and spring data sets. The ability of the WTC system to measure tree-level physiological responses is demonstrated. All WTCs displayed a high level of control over tracking of air temperatures. The set target of 365 micromol mol(-1) in the ambient [CO(2)] chambers was too low to be maintained during winter because of tree dormancy and the high natural increase in [CO(2)] over winter at high latitudes such as the Flakaliden site. Accurate control over [CO(2)] in the ambient [CO(2)] chambers was restored during the spring and the system maintained the elevated [CO(2)] target of 700 micromol mol(-1) for both measurement periods. Air water vapour deficit (VPD) was accurately tracked in ambient temperature WTCs. However, as water vapour pressure in all 12 WTCs was maintained at the level of non-chambered (reference) air, VPD of elevated temperature WTCs was increased.     

CO2浓度升高与氮添加对三江平原湿地小叶章生长的影响

利用开顶箱薰气室,设置正常大气CO₂浓度(350 μmol·mol^-1)、高CO₂浓度(700 μmol·mol^-1)2个CO₂水平和不施氮(0 g N·m^-2)、中氮(5 g N·m^-2)和高氮(15 g N·m^-2)3个氮素水平,研究CO₂浓度升高和氮肥施用对三江平原草甸小叶章生长的影响.结果表明：随着CO₂浓度升高,小叶章物候期提前,其中抽穗期提前1～2 d,成熟期提前3 d;不施氮、中氮和高氮水平下, CO₂浓度升高使小叶章的分蘖分别增加8.2%(P<0.05)、8.4%(P<0.05)和5.5%(P>0.05);在小叶章生长初期,CO₂浓度升高对其生物量的增加有促进作用,拔节期和抽穗期小叶章地上生物量分别增加12.4%和20.9%(P<0.05);生长后期则对小叶章地下生物量的促进作用增大,腊熟期和成熟期的地下生物量分别增加20.5%和20.9% (P<0.05).小叶章生物量对高浓度CO₂的响应与供氮水平有关,供氮充足条件下, 高浓度CO₂对生物量的促进效应更大.     

Effects of elevated CO2 on the capacity for photosynthesis of a single leaf and a whole plant, and on growth in a radish

The atmospheric concentration of CO2 will probably rise to about 700 micromol mol(-1) by the end of this century. The effects of elevated growth CO2 on photosynthesis are still not fully understood. Effects of elevated growth CO2 on the capacity for photosynthesis of a single leaf and a whole plant were investigated with the radish cultivar White Cherish. The plants were grown under ambient ( approximately 400 micromol mol(-1)) or elevated CO2 ( approximately 750 micromol mol(-1)). The rates of net photosynthesis per leaf area with a whole plant and a single leaf of plants of various ages (15-26 d after planting) were measured under ambient and elevated CO2. The rates of photosynthesis were increased by 20-28% by elevated CO2. There was no effect of elevated growth CO2 on the rate of photosynthesis, clearly indicating no downward acclimation of photosynthesis to elevated CO2. Elevated CO2 increased dry weight accumulation by >27%. The effect of elevated CO2 on other growth characteristics will also be shown.     

CO2浓度增加对长白山森林土壤甲烷氧化影响

大气CO2浓度升高可能对森林土壤的甲烷(CH4)氧化速率产生影响.本文采用开顶箱技术,对连续6年高浓度CO2(500 μmol·mol-1)处理的长白山森林典型树种蒙古栎树下土壤CH4氧化速率进行研究,并利用CH4氧化菌的16S rRNA特异性引物以及CH4单加氧酶功能基因引物分析了土壤中CH4氧化菌的群落结构与数量.结果表明:CO2浓度增高后,生长季土壤甲烷氧化量与对照和裸地相比分别降低了4％和22％;基于16S rRNA特异性引物的DGGE分析表明,CO2浓度增高导致两类甲烷氧化菌的多样性指数降低;CO2浓度增高对土壤中Ⅰ类甲烷氧化菌数量无显著影响,而使土壤中Ⅱ类甲烷氧化菌数量显著减少,功能基因pmoA拷贝数与对照和裸地相比分别降低了15％和46％.CO2浓度增高导致森林土壤甲烷氧化菌数量与活性降低,土壤含水量的增加可能是导致这一现象的主要原因.