西藏飞蝗各发育阶段的耐寒性

西藏飞蝗Locusta migratoria tibetensis Chen是青藏高原的重要农牧业害虫。对该虫各发育阶段的过冷却点和结冰点的测定表明,西藏飞蝗卵的过冷却点和结冰点为最低,分别为-22.02℃、-16.36℃,4龄蝻的过冷却点和结冰点最高,分别是-6.46℃和-5.05℃,西藏飞蝗在甘孜以卵越冬。     

西藏飞蝗（Locusta migratoria tibetensis Chen）耐寒性理化指标

西藏飞蝗是青藏高原上的一种重要害虫.近年来, 西藏飞蝗在西藏、青海玉树以及川西高原上危害十分严重.通过对该虫各发育阶段过冷却能力测定,研究了各发育阶段虫体内生理生化物质与过冷却能力之间的关系,揭示了该虫耐寒性规律.结果表明,西藏飞蝗蝗卵的过冷却点和结冰点为最低,分别为-22.02℃和-16.36℃.在1～4龄蝗蝻期,过冷却点随体内水分、糖原和脂肪含量随着虫体的发育而升高,其过冷却点分别为-9.28℃、-8.51℃、-7.18℃、-6.46℃;到5龄蝗蝻时,虫体脂肪和糖原含量达最高值,过冷却点下降为-8.07℃.讨论了虫体脂肪、糖原和甘油含量变化与其耐寒力之间的关系.     

越北腹露蝗卵的过冷却特性

利用热电偶法测定越北腹露蝗FruhstorferiolatonkinensisWill卵的过冷却点。结果表明,越北腹露蝗卵的过冷却点随季节变化而变动。卵的平均过冷却点从2004年9月初(刚产的卵)的-16.0℃显著下降到2005年1月的-24.7℃,随后卵的过冷却点显著增加到2005年3月的-10.7℃。滞育卵的平均过冷却点(-20.1℃)低于滞育后的平均过冷却点(-12.4℃)。滞育卵在5℃下适应30d后平均过冷却点显著减小到-25.9℃,而在30℃下适应30d后平均过冷却点显著增加到-8.2℃。     

光照与飞蝗卵耐寒性的关系

用热电偶法测定长光照(L：D=14：10)和短光照(L：D=10：14)条件下饲养的飞蝗所产卵的过冷却点;并对长、短光照组分别设置5个温度(0、-5、-10、-15和-20℃)处理,每一温度又设置5个时间(6h,1、3、5和10d)处理,然后检查其28℃的孵化数,以此计算低温存活率和半致死温度。长、短光照组卵的过冷却点没有差异;两种光照条件下的低温存活率随着卵处理温度的降低和时间的延长而下降,在-5和-10℃时短光照组的低温存活率显著高于长光照组;卵的半致死温度随低温处理时间的延长而升高,短光照组的半致死温度明显低于长光照组。接受短光照的飞蝗母本能产出耐寒性较高的卵,暗示秋天所产的卵能更成功地越冬。     

意大利蝗越冬卵抗寒性初步研究

过冷却能力是昆虫越冬的主要策略之一。本研究测定了新疆乌鲁木齐2015年9月至次年5月意大利蝗卵过冷却点及土壤温度等指标的变化。结果显示,9月至次年3月, 80%以上的蝗卵过冷却点低于-24.0℃;4月和5月,蝗卵过冷却点显著上升至-22.7±0.7℃和-21.0±0.7℃,过冷却点在-23.9℃～-20.0℃的蝗卵比例分别达77.1%和52.1%。越冬过程中蝗卵含水量在土壤温度降至零下时短暂下降后迅速回升,并在越冬后的4月和5月剧烈上升。土壤温度(地下5 cm)较高且变化幅度不大,1月开始降至零下,2月达到最低。较高且稳定的土壤温度可能是意大利蝗卵维持较低过冷却点的原因,意大利蝗卵较强的抗寒能力是新疆草原蝗灾持续暴发的潜在原因。研究结果对开展新疆意大利蝗监测预警有很大意义。     

飞蝗不同地理种群抗寒性研究

新疆和硕、哈密和天津北大港飞蝗种群的过冷却点随发育阶段而升高,在胚胎发育的各期都具有较低的过冷却点。飞蝗卵的过冷却点在这3个地理种群间无明显差异。雄性蝻和成虫双雌性的过冷却能力强,和硕飞蝗的成虫过冷点比北大港的飞蝗成虫要低。低温胁迫可诱导蝗卵以糖元为朱料合成抗冻保护剂,和硕飞蝗种群的卵主要合成甘油和山梨醇,北大港飞蝗种群的卵主要合成海藻糖,而哈密飞蝗种群的卵却可以合成甘油、山梨醇和海藻糖等抗冻保护剂。糖元磷酸化酶的总量随发育阶段而下降,但热休克和冷休克并不改变其总量,冷休克在胚胎发育的各阶段可以使其活性升高10%-40%,北大港飞蝗种群和哈密飞蝗种群的蝗卵经热休克2h处理后,糖元磷酸化酶的变化与在冷休克下的情况相似,而和硕飞蝗处群胚胎发育的Ⅱ期,热休克诱导糖元磷酸化酶活性的升高程度较低。     

桔小实蝇自然种群蛹和越冬成虫的耐寒性

本文通过过冷却点和低温耐寒能力的测定,研究了我国桔小实蝇Boctrocera dorsalis (Hendel)不同季节种群和地理种群蛹的耐寒性规律。结果表明：桔小实蝇自然种群蛹的低温存活力有明显的季节规律,冬前种群的耐寒性显著强于夏季种群。如在0℃低温下暴露2天,夏季种群基本全部死亡,而冬前种群存活率仍为61.4%。2005-2006年不同月份桔小实蝇1日龄蛹过冷却点差异明显,以2月份均值最高,为-9.26℃,8月份最低,为-15.20℃;且2月份桔小实蝇蛹个体过冷却点值出现明显的分化现象,分别在-15℃左右和-6℃左右出现两个明显的聚集区。所研究的5个桔小实蝇地理区域种群没有出现明显的耐寒性分化。结合本文结果,文中还就桔小实蝇自然种群耐寒性影响因素进行了讨论。     

椰心叶甲的耐寒力测定

测定椰心叶甲Brontispalongissima(Gestro)的过冷却点和结冰点。各虫态中过冷却点值以卵和蛹最低,分别为-9.8℃和-9.0℃,成虫较高,雌、雄成虫分别为-5.9℃和-5.5℃;幼虫的过冷却点最高,平均为-4.5℃。结冰点由高到低依次为2龄幼虫、5龄幼虫、4龄幼虫、1龄幼虫、3龄幼虫、雄成虫、雌成虫、蛹和卵,分别为-0.3,-0.8,-1.1,-1.3,-1.3,-2.4,-2.7,-5.4和-5.5℃。综合分析认为,蛹、卵的耐寒能力最强。根据广东地区常年平均气温分析了在自然条件下该虫各虫态越冬情况,但由于生物学特性的限制,越冬虫态主要是成虫、蛹。     

稻水象甲越冬成虫的耐寒力测定

测定了浙江乐清 (北纬 2 8 1 4°,东经 1 2 0 94°)稻水象甲LissorhoptrusoryzophilusKuschel越冬成虫的耐寒力 ,从 1 1月下旬至 3月下旬平均过冷却点温度为 -1 5 3～ -2 1 2℃ ,结冰点温度为 -1 4 0～ -1 8 6℃。结合发生地气温条件 ,讨论了该地冬春季低温对该虫越冬群存活的影响。     

温度及土壤含水量对淡剑纹灰翅夜蛾死亡率的影响

淡剑纹灰翅夜蛾Spodopteradepravata(Bulter)在 2 2 ,2 5 ,2 8,3 1 ,3 4,3 6,3 8℃恒温条件下卵孵化率分别为 96. 0 0 ,91 .0 0 ,83. 90 ,84. 81 ,83 . 60 ,80 %和 0。高龄幼虫在低温环境下 ,一般随温度的降低、时间的延长死亡率增加。过冷却点为 (-6. 81± 1 . 63 )℃ ,体液冰点为 (-3 .5 1± 1 . 5 6)℃。 1 0 %土壤含水量时该虫羽化率最高 ,40 %及其以上羽化率为 0。     

光周期对中华通草蛉自然越冬成虫及实验种群幼虫耐寒能力的影响

【目的】滞育诱导期进行短光照处理可影响昆虫耐寒性。为明确光周期对中华通草蛉Chrysoperla sinica (Tjeder)耐寒性的影响, 针对中华通草蛉滞育解除过程及非滞育虫态的耐寒性进行了一系列研究。【方法】测定了中华通草蛉自然越冬成虫的过冷却点（supercooling point, SCP）以及长光周期（15L∶9D）和短光周期（9L∶15D）条件下自然越冬成虫在滞育解除过程中在-12℃下的死亡率, 并测定了室内长、 短两种光周期下实验种群2龄和3龄幼虫的过冷却点（SCP）、 结冰点（freezing point, FP）以及-7℃下的死亡率。【结果】中华通草蛉12月份的自然越冬成虫SCP集中在-10~-14℃之间。SCP低于-12℃的个体占43.70%, 且-12℃处理1 d死亡率为62.00%。-12℃处理1 d条件下的长、 短光周期处理自然越冬成虫, 除处理0 d外, 长光周期处理死亡率均高于短光周期处理的, 且在处理15 （P=0.012）, 20 （P=0.01）和25 d （P=0.001）差异显著。中华通草蛉试验种群相同龄期幼虫在短光周期下的SCP和FP均高于长光周期下, 但差异不显著(P>0.05); 但在-7℃下, 2龄幼虫短光周期下的低温死亡率为67.00%±4.04%, 显著低于长光周期下的低温死亡率（78.00%±1.33%）（P=0.011）, 3龄幼虫短光周期条件下低温死亡率为24.33%±1.33%, 显著低于长光周期下的低温死亡率（53.00%±3.46%）（P=0.002）。【结论】中华通草蛉为结冰敏感型, 诱导滞育的短光照处理可提高其幼虫期及滞育解除过程中成虫的耐寒能力。     

HISTORICAL EVIDENCE FOR POPULATION DYNAMICS OF TIBETAN MIGRATORY LOCUST AND THE FORECAST OF ITS OUTBREAK*

Abstract The Tibetan migratory locust (Locusta migratoria tibetensis Chen) is the highest altitude distributed subspecies among the 10 subspecies of migratory locusts. It was discovered and described as new subspecies in 1963. It is mostly distributed above an elevation of 3 000 m, with the highest up to 4 600 m, on the “Roof of the World,” viz. the Qinghai-Xizang Plateau in southwestern China. Recent study on the historic literature revealed ancient records of locust plagues caused by Locusta migratoria tibetensis Chen in many regions of Xizang (Tibet). These disasters took place during 124 years stretching from 1828 to 1952. Forty-five places were infested by locust swarms, and from 1846 to 1857 locust disasters occurred sucessively in 12 years, and affected 18 places of Xizang. At the severe disaster regions crops were damaged by locusts in such a degree that there was no harvest at all. At the same time, locust plagues due to another subspecies also occurred in the plains between the Yellow River and Huaihe and Haihe Rivers in East China. The disasters in Xizang were caused by the Tibetan migratory locust, while in Huang-Huai-Hai Plain disasters were due to the Oriental migratory locust Locusta migratoria manilensis (Meyen). This study not only provides the evidence that the Tibetan migratory locust has been existing as a separate subspecies with a long history, but also reveals the relevant years of locust plagues, their regional distribution and intermittent rules of locust plagues. It also offers a scientific basis for forecasting Tibetan migratory locust disasters and related monitoring strategies; as well as understanding the close relationship between the outbreak of the migratory locust and drought.     

西藏飞蝗蝗蝻群集迁移特性及其群集效应

西藏飞蝗Locusta migratoria tibetnsis Chen暴发成灾的重要原因之一是蝗蝻具有群集迁移危害习性。为阐明西藏飞蝗灾变的行为机制,为西藏飞蝗的监测预警和防治提供科学依据,利用视频跟踪技术测定了自然环境中西藏飞蝗蝗蝻群集迁移的运动速度、方向,建立自推进粒子模型模拟蝗蝻群集迁移行为,分析群集迁移效应。结果表明,①不同自然环境中的西藏飞蝗蝗蝻在群集迁移过程中,群体内个体的运动表现出定向集体运动,群集迁移速度为0.1256 m/s,0.2 m以内的个体蝗蝻方向趋向一致。沙滩、翻耕农田和草地蝗蝻群运动一致性参数均较高,分别为0.8502、0.7870和0.6987。②西藏飞蝗蝗蝻群由分散运动转变为群集迁移存在临界密度,密度较低时群体内个体分散运动,当蝗蝻密度达到12-15头/m²时,蝗蝻群体由分散运动转变为高度一致的群集迁移运动。③蝗蝻群通过群集迁移可以显著增加迁移距离,随机运动蝗蝻1 d扩展只有70-80 m,而群集迁移1 d最大距离可达2.5 km。蝗蝻群集迁移可以提高发现特别是远距离食物等资源的概率,使群体中更多的个体受益。④尽管未发现室外蝗蝻群存在先验个体,但模拟发现在群集迁移群体中,只需要少数先验个体（3%-5%）即可引导整个蝗蝻群运动。     

温度对意大利蝗呼吸代谢的影响

【目的】意大利蝗Calliptamus italicus L.是新疆荒漠、半荒漠草原的主要危害种类,前期研究表明其发生与新疆气候变暖显著相关,本研究进一步探讨气候变暖条件下意大利蝗的呼吸代谢适应机制。【方法】应用多通道昆虫呼吸仪测定了15, 20, 25, 30和35℃不同温度条件下意大利蝗成虫的呼吸率、代谢率、CO₂释放率、Q₁₀(每升高10℃呼吸率的变化幅度)及呼吸商并分析其变化特征。【结果】15℃时意大利蝗成虫的呼吸率、代谢率、CO₂释放率均显著低于其他温度（P<0.01）, 35℃时3项指标值显著高于其他温度（P<0.01）,表明低于15℃和高于35℃的温度条件都对意大利蝗的呼吸代谢产生明显影响;在20~30℃之间,意大利蝗的呼吸率、代谢率及CO2释放率变化幅度小且平稳,差异均不显著（P>0.01）,表明该温度范围是意大利蝗生长发育的适宜温度条件。不同温度下意大利蝗呼吸率的Q₁₀值显示, 20~30℃温度范围内的 Q₁₀值最小,为1.03, 15~25℃的Q10值为1.43, 25~35℃的Q10值最大,为2.42,说明意大利蝗的呼吸代谢活动对温度变化敏感,并表明20~30℃是意大利蝗生长发育的适宜范围。各温度条件下意大利蝗呼吸商的差异不显著（P>0.01）,平均为0.9450,判断意大利蝗呼吸代谢消耗的底物主要为糖类物质。【结论】意大利蝗生长发育的适宜温度范围是20~30℃,预示着在气候持续变暖背景下,意大利蝗仍将是新疆草原最重要的生物灾害之一。     

Effects of summer frost exposures on the cold tolerance strategy of a sub-Antarctic beetle

The sub-Antarctic beetle Hydromedion sparsutum (Coleoptera, Perimylopidae) is common locally on the island of South Georgia where sub-zero temperatures can be experienced in any month of the year. Larvae were known to be weakly freeze tolerant in summer with a mean supercooling point (SCP) around -4 degrees C and a lower lethal temperature of -10 degrees C (15min exposure). This study investigated the effects of successive freezing exposures on the SCP and subsequent survival of summer acclimatised larvae. The mean SCP of field fresh larvae was -4.2+/-0.2 degrees C with a range from -1.0 to -6.1 degrees C. When larvae were cooled to -6.5 degrees C on 10 occasions at intervals of 30min and one and four days, survival was 44, 70 and 68%, respectively. The 'end of experiment' SCP of larvae surviving 10 exposures at -6.5 degrees C showed distinct changes and patterns from the original field population depending on the interval between exposure. In the 30min interval group, most larvae froze between -6 and -8 degrees C, a depression of up to 6 degrees C from the original sample; all larvae were dead when cooling was continued below the SCP to -12 degrees C. In the one and four day interval groups, most larvae froze above -6 degrees C, showing no change as a result of the 10 exposures at -6.5 degrees C. As with the 30min interval group, some larvae froze below -6 degrees C, but with a wider range, and again, all were dead when cooled to -12 degrees C. However, in the one and four day interval groups, some larvae remained unfrozen when cooled to -12 degrees C, a depression of their individual SCP of at least 6 degrees C, and were alive 24h after cooling. In a further experiment, larvae were cooled to their individual SCP temperature at daily intervals on 10 occasions to ensure that every larva froze every day. Most larvae which showed a depression of their SCP of 2-4 degrees C from their day one value became moribund or died after six or seven freezing events. Survival was highest in larvae with SCPs of -2 to -3 degrees C on day one and which froze at this level on all 10 occasions. The results indicate that in larvae in which the SCP is lowered following sub-zero exposure, the depression of the SCP is greatest in individuals that do not actually freeze. Further, the data suggest that after successive frost exposures in early winter the larval population may become segregated into two sub-populations with different overwintering strategies. One group consists of larvae that freeze consistently in the temperature range from -1 to -3 degrees C and can survive multiple freeze-thaw cycles. A second group with lower initial SCPs (around -6 degrees C), or which fall to this level or lower (down to -12 degrees C) after freezing on one or more occasions, are less likely to freeze through extended supercooling, but more likely to die if freezing occurs.     

Composition and emission dynamics of migratory locust volatiles in response to changes in developmental stages and population density

Chemical communication plays an important role in density‐dependent phase change in locusts. However, the volatile components and emission patterns of the migratory locust, Locusta migratoria, are largely unknown. In this study, we identified the chemical compositions and emission dynamics of locust volatiles from the body and feces and associated them with developmental stages, sexes and phase changes. The migratory locust shares a number of volatile components with the desert locust (Schistocerca gregaria), but the emission dynamics of the two locust species are significantly different. The body odors of the gregarious nymphs in the migratory locust consisted of phenylacetonitrile (PAN), benzaldehyde, guaiacol, phenol, aliphatic acids and 2,3‐butanediol, and PAN was the dominant volatile. Volatiles from the fecal pellets of the nymphs primarily consist of guaiacol and phenol. Principal component analysis (PCA) showed significant differences in the volatile profiles between gregarious and solitary locusts. PAN and 4‐vinylanisole concentrations were significantly higher in gregarious individuals than in solitary locusts. Gregarious mature males released significantly higher amounts of PAN and 4‐vinylanisole during adulthood than mature females and immature adults of both sexes. Furthermore, PAN and 4‐vinylanisole were completely lost in gregarious nymphs during the solitarization process, but were obtained by solitary nymphs during gregarization. The amounts of benzaldehyde, guaiacol and phenol only unidirectionally decreased from solitary to crowded treatment. Aliphatic aldehydes (C7 to C10), which were previously reported as locust volatiles, are now identified as environmental contaminants. Therefore, our results illustrate the precise odor profiles of migratory locusts during developmental stages, sexes and phase change. However, the function and role of PAN and other aromatic compounds during phase transition need further investigation.     

改治结合,根除蝗害的关键因子是“水”!

改治结合,根除蝗害的关键因子是“水”。其主要原因为:(1)水灾是为东亚飞蝗创造发生地和转移到临近地区危害的关键因子;(2)东亚飞蝗产卵的适宜的土壤含水量为10%～20%(沙土为10%～12%,壤土为15%～18%,粘土为18%～20%);(3)先涝后旱,飞蝗成片;(4)黄(河)患是东亚飞蝗蝗害发生的源泉。南水北调东线工程的实施,有利于滨湖(库)蝗区东亚飞蝗的持续控制。     

中国嵩草属植物地理分布模式和适应的气候特征

为了明确嵩草属(Kobresia)植物分布与气候要素的关系, 收集了嵩草属植物地理分布资料和气象台站气候数据, 应用ArcGIS软件及SPSS软件中的聚类分析方法, 分析了嵩草属植物地理分布模式和适应的气候特征。结果显示: 嵩草属植物分布在青藏高原、西北、华北和东北部分地区, 广泛分布13种, 间断分布10种, 分布海拔为1 400-5 000 m, 经度和纬度范围分别为81-112° E和23-46° N。嵩草属植物适应的气候要素平均值范围: 年生物学温度为4-19 ℃, 年平均气温为0-20 ℃, 年平均最高气温为7-28 ℃, 年平均最低气温为-6-16 ℃, 极端最高气温为25-40 ℃, 极端最低气温为-37.0-0.0 ℃, 1月和7月平均气温分别为-14-13 ℃和11-24 ℃, 1月和7月最高气温分别为-7-23 ℃和18-30 ℃, 1月和7月最低气温分别为-22-7 ℃和5-20 ℃, 春夏秋冬季气温分别为-4-19 ℃、9-23 ℃、6-21 ℃和-11-15 ℃, 温暖指数为23-159 ℃, 寒冷指数为-36-0 ℃, 年降水量为154-1 500 mm, 春夏秋冬降水量分别为19-135 mm、53-662 mm、48-545 mm和5-92 mm, Holdridge潜在蒸散量为261-1 100 mm, Thornthwaite潜在蒸发量为399-895 mm, 干燥度为167-786, 湿润指数为179-816, 4-10月日照时数为990-2 100 h。在热量要素平均值较低和中等、降水量与干燥湿润度平均值中等或辐射时数平均值较高范围下分布种数较多。嵩草属植物适应的气候要素极值, 年平均气温最小最大值范围为-6-21 ℃, 年平均最低气温最小值最高气温最大值范围为-12-28 ℃, 极端最低气温最小值最高气温最大值范围为-48-42 ℃, 最冷最热月气温范围为-32-33 ℃, 冬夏季最低最高气温范围为-20-25 ℃, 降水量最小最大值范围为15-1 800 mm, 干燥度最小最大值范围为7-890, 日照时数最小最大值范围为701-2 300 h。在热量要素极值较低、降水量及干燥度极值中等或日照时数极值较大范围下分布种数较多。说明嵩草属植物主要适应于低温亚湿润型和中温湿润型气候。     

Olfactory discrimination of two enantiomers of 4-methyl-hexanoic acid by the migratory locust and the honeybee

Summary Single olfactory receptor cells of the migratory locust and of honeybee drones were stimulated with the two enantiomeric forms of 4-methylhexanoic acid. In the majority of cells, equal stimulus quantities of either the (+)- or the (–)-isomer elicited a greater number of nerve impulses. Behavioral discrimination of the two enantiomers was demonstrated in drones. Here, the proboscis extension response served as indicator of the discrimination following the conditioning to one or the other steric antipode. These findings are explained by a minimum of two variants of one structurally highly stabilized type of acceptor present in different numbers on the membranes of individual cells of this type.     

寄主对桔小实蝇耐寒性的影响

为了研究寄主营养对桔小实蝇耐寒性的作用,测定了以15种果蔬饲养的桔小实蝇1日龄蛹的过冷却点(supercooling points,SCP); 再选取南瓜、西红柿、柑桔、番石榴和杨桃等5种果蔬,测定了桔小实蝇3龄老熟幼虫、1日龄蛹、3日龄蛹、5日龄蛹、7日龄蛹和雌雄成虫的过冷却点,并观察了1日龄蛹的低温存活力。结果表明：（1）15种果蔬饲养所得的桔小实蝇1日龄蛹SCP均值在-11.03℃～-13.17℃,不同寄主发育的桔小实蝇SCP值存在显著性差异,其中以取食蒲桃的最高,为-11.03℃,取食苦瓜的最低,为-13.17℃。（2）5种果蔬饲养所得的桔小实蝇各虫态的SCP均值存在极显著差异(F_(4,863)＝35.6,P<0.01); 同一寄主上的桔小实蝇不同虫态之间SCP均值也达到极显著性差异(F_(6,863)＝392.9,P<0.01); 且寄主和发育龄期之间存在着极显著的交互作用(F_(24,863)＝9.4,P<0.01)。（3）桔小实蝇各发育阶段,SCP值表现一定变化： 老熟幼虫发育至1日龄蛹,SCP值变化不大; 蛹发育至3、5和7天过冷却能力明显增强,降至-20℃左右,但他们之间没有明显区别; 羽化后3～5天的成虫SCP值又升高至-10℃左右。老熟幼虫、1日龄蛹和2～3日龄成虫与3日龄、5日龄和7日龄蛹的SCP值之间有显著性差异。（4）将5种果蔬饲养所得的桔小实蝇1日龄蛹置于6℃和-3℃下进行较长时间(1～8天)和较短时间(1～8 h)的低温处理,发现番石榴、杨桃和南瓜发育的蛹经低温处理后的校正羽化率较西红柿和柑桔发育的蛹高; 同样在0℃、3℃、6℃和9℃处理(2天)的实验中,得出相似的结果。因此,本实验结果表明桔小实蝇幼虫由于生活寄主的不同使得其下一代蛹的耐寒性产生了差异,引起其差异的原因值得进一步研究。