长江中华绒螯蟹亲体和早期发育阶段对盐度的生理与行为响应
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摘要
中华绒螯蟹(Eriocheir sinensis)在我国俗称河蟹,隶属于甲壳纲(Crustacea),十足目(Decapoda),方蟹科(Crapsida),绒螯蟹属(Eriocheir),是甲壳类中少有的洄游性种类。在淡水河流、湖泊中完成生殖蜕壳后的中华绒螯蟹需向河口区域进行生殖洄游,在河口完成交配、抱卵、胚胎及幼体的发育。长江口是中华绒螯蟹最大、最优质的繁殖场。盐度是河口时空变化最为剧烈的因子之一,近年来长江口盐度变化加剧,盐水入侵严重,这可能对中华绒螯蟹的洄游、繁育造成重要的影响。
为全面评估河口盐度变动对中华绒螯蟹繁育的潜在影响,本研究以中华绒螯蟹亲体、后期胚胎及早期幼体为研究对象,从血淋巴渗透、代谢、免疫调节与肝胰腺消化酶调节以及行为方面研究了中华绒螯蟹亲体在洄游过程中对盐度的适应策略及耐盐阂值；以孵化率与初孵蚤状幼体的存活率为评价指标获得中华绒螯蟹胚胎及早期幼体发育的最适盐度范围,在此基础上分析了盐度胁迫后胚胎对能源物质的分解与利用特征、Na+、K+-ATPase的响应以及胰高血糖素(CHH)可能的神经内分泌调控作用；通过研究大眼幼体的盐度选择行为,不同盐度下的生态蜕壳及Na+/K+-ATPase活性对盐度的响应,探讨大眼幼体趋江洄游的行为与生理基础,初步阐明了不同生命阶段的中华绒螯蟹对盐度的生理与行为响应机制。本研究成果为全面评估河口盐度变动对中华绒螯蟹繁育的影响提供重要理论依据,对中华绒螯蟹野生资源的保护、利用和人工繁殖均具有重要意义,同时也丰富了甲壳类洄游生理学与渗透调节生理、行为研究的资料。主要研究结果如下：
1.中华绒螯蟹亲体在不同盐度下的行为反应
采用视频记录分析法比较研究了雌性中华绒螯蟹亲体适应了淡水、半咸水(盐度18)后以及在高盐(盐度18→30)、低盐(盐度18→0)胁迫下的行为反应。结果表明：在不同盐度下,蟹口器的活动性与其个体的运动活力呈现一致的反应模式,与淡水组相比,适应盐度18后蟹的运动活力和口器的活动性均增强(P<0.05)；低盐胁迫引起蟹运动活力和口器活动(P<0.05)增强,而高盐胁迫后蟹运动活力显著减弱(P<0.05),且口器的活动性也降低,且在低盐胁迫后蟹的运动活力和口器的活动性均显著强于淡水组(P<0.05)；第一触角回缩行为和封闭反应行为几乎仅出现在盐度18组以及高盐胁迫组(P<0.05),第一触角回缩至壳槽内的时间随着试验时间的延长而增加；第二触角抖动次数随着试验盐度的升高而逐渐降低,低盐胁迫后蟹的第二触角抖动次数显著高于其它三个试验组(P<0.05)；高、低盐胁迫均引起蟹的眼柄活动性增强,而适应盐度18后眼柄的活动性降低；盐度18组蟹清洁触角的频率显著高于淡水组(P<0.05),高、低盐胁迫均引起蟹的触角清洁频率显著降低(P<0.05)；腹部开合行为仅出现在低盐胁迫后。综合以上结果显示,从淡水适应半咸水后会刺激中华绒螯蟹亲体的运动活力及其它相关行为增强,蟹在低盐胁迫后出现腹部开合行为是其在低渗环境下通过暴露尾肠从而吸收水中离子的一种行为策略,蟹在高盐胁迫后表现出的封闭反应则有助于减少机体在高渗环境下对水分的吸收及盐分的被动扩散丢失。
2.中华绒螯蟹亲体的盐度选择行为
利用室内可控的六分室盐度选择装置结合视频监控系统,研究了洄游期的雌、雄中华绒螯蟹对盐度的选择行为。结果显示：雄蟹对盐度并未表现出显著的偏好行为,但其在盐度28分室中出现的次数最少,而在盐度35分室中出现最多,在21与35盐度分室中停留的时间略长于其它盐度分室；在盐度0-21范围内,雌蟹在各盐度分室中出现的次数随着盐度的升高而增加,在21盐度分室中达到最高,在28盐度分室中最低,雌蟹在35盐度分室中出现的次数接近盐度21分室的水平。在盐度7-28范围内,雌蟹在各盐度分室中停留的时间随着盐度的升高逐渐降低,而在35盐度分室中停留时间最长,且显著高于其在28盐度分室中停留时间(P<0.05)。本试验同时发现,在试验时间内,雄蟹的运动活力显著高于雌蟹,在提供可选择的盐度环境后,雌、雄蟹的运动活力均降低。结果提示,盐度在中华绒螯蟹生殖洄游寻找产卵场的过程中虽然发挥着一定的导向作用,但其它栖息地环境因子如水深、底质可能也是影响中华绒螯蟹在河口分布位置的重要因子。
3.盐度升高对中华绒螯蟹亲体渗透压、离子及离子调节酶的影响
通过逐步增盐法研究了中华绒螯蟹亲体盐度适应过程中血淋巴渗透压、离子含量以及后鳃Na+/K+-ATPase和碳酸酐酶(CA)活性的变化。结果显示：在0-21的盐度范围内,雌、雄中华绒螯蟹均显著调高其血淋巴渗透压(P<0.05),而在盐度达到35时略调低其血淋巴渗透压,雌、雄中华绒螯蟹的等渗点接近盐度28；随着环境水体盐度的增加和血淋巴渗透压的升高,中华绒螯蟹血淋巴Na+和Cl-含量升高,而Ca2+和K+含量呈现降低的趋势；盐度升高引起中华绒螯蟹后鳃Na+/K+-ATPase和CA活性降低,酶活性的降低是后鳃主动吸收Na+减少的直接生理反应；比较雌、雄中华绒螯蟹各检测指标可见,雌蟹血淋巴Ca2+和K+含量和鳃CA活性在一些盐度水平下显著高于雄蟹(P<0.05),而其它测定指标之间无显著的性别差异。研究结果表明,中华绒螯蟹亲体具有较强的高渗调节能力,但其低渗调节能力较弱。盐度升高后,中华绒螯蟹亲体主要通过调节血淋巴Na+和Cl-含量而调节血淋巴渗透浓度,Na+/K+-ATPase和CA在离子调节过程中发挥重要作用,雌、雄蟹的渗透调节能力不存在显著的差异。
4.盐度升高对中华绒螯蟹亲体血淋巴代谢、免疫指标及肝胰腺消化酶活性的影响
通过逐步增盐法,研究了从淡水逐步升高到盐度7、14、21、28、35后,雌、雄中华绒螯蟹血淋巴代谢指标(总蛋白蛋白、氧合血蓝蛋白、葡萄糖、甘油三酯、胆固醇、尿酸、尿素、肌酐)的含量、免疫酶(碱性磷酸酶、超氧化物歧化酶、酚氧化酶)和消化酶(淀粉酶、纤维素酶、胃蛋白酶、胰蛋白酶、脂肪酶)活性的变化。结果显示：随着盐度的增加,雌、雄中华绒螯蟹血淋巴总蛋白、氧合血蓝蛋白含量降低；雄蟹血淋巴甘油三酯、胆固醇、尿素、肌酐含量与雌蟹血淋巴甘油三酯、尿酸含量均随着盐度的升高先显著降低(P<0.05),在盐度升高到21时达到最低值,而后显著升高(P<0.05)；雄蟹血淋巴尿酸含量在0-21的盐度范围内呈现显著降低的趋势(P<0.05),盐度高于21时维持在稳定水平；雌蟹血淋巴尿素含量在盐度升高到21达到最低值(P>0.05),盐度高于21时显著升高(P<0.05)；盐度增加对雌蟹血淋巴肌酐、胆固醇含量无显著影响；雌、雄蟹血糖含量均在盐度升高到21时达到最高水平；雌蟹血淋巴多种代谢指标均高于雄蟹。
随着盐度的升高,雄蟹血淋巴酚氧化酶活力逐渐降低,雌蟹血淋巴酚氧化酶活力先降低,当盐度高于21后维持稳定水平；雌、雄蟹血淋巴碱性磷酸酶活性呈现先略升高而后逐渐降低的趋势；雌、雄蟹血淋巴超氧化物歧化酶活性先略降低,盐度高于21时被激活,且在盐度升高到35时酶活性显著高于盐度21时的水平(P<0.05)。除超氧化物歧化酶外,雌蟹血淋巴其它两种免疫酶活性均高于雄蟹。
雌、雄中华绒螯蟹肝胰腺淀粉酶活性最高,脂肪酶活性极低(P<0.05)；雌蟹肝胰腺淀粉酶、胃蛋白酶、胰蛋白酶活性均在盐度达到及高于28时显著降低(P<0.05),而纤维素酶活性在盐度升高到35时显著降低(P<0.05)；除脂肪酶外,雄蟹的肝胰腺消化酶活性均在盐度升高到14时显著升高(P<0.05),盐度升高到21时显著降低(P<0.05),之后维持稳定水平或进一步降低；除脂肪酶外,雄蟹肝胰腺消化酶活性在不同盐度水平(盐度21除外)下均显著高于雌蟹(P<0.05)。
综合上述研究结果显示：中华绒螯蟹亲体从淡水逐步适应海水的过程中多种血淋巴代谢指标均在盐度增加到21时达到最低点,这一盐度下机体可能用于渗透调节的能量最低,从而可以节约更多的能量用于繁殖。当水体盐度接近或超过28时中华绒螯蟹的免疫防御能力和消化能力均显著降低,最终可能对中华绒螯蟹的繁殖造成潜在的不利影响。在不同盐度下,雌蟹血淋巴代谢指标和免疫指标稍高于雄蟹,表明雌蟹的代谢水平和免疫防御能力相对较高。引起雄蟹肝胰腺消化酶活性降低的起始盐度值低于雌蟹,这从消化生化的角度表明雌蟹对盐度增加的适应能力强于雄蟹。
5.盐度对中华绒螯蟹胚胎及幼体发育的影响
研究了盐度对不同发育时期中华绒螯蟹离体胚胎发育、蚤状幼体存活的影响,结果显示：从眼点期开始盐度暴露其胚胎孵化率与幼体存活率均低于从原肠期和出膜前期开始暴露胚胎的孵化率,原肠期和出膜前期胚胎在盐度1-20范围内呈现较高的孵化率和较快的发育速度,且各盐度组间无显著差异(P>0.05)；盐度降低(盐度1、5)在一定程度上促进了胚胎的发育并提高了胚胎孵化率,且在眼点期表现出显著性(P<0.05)；盐度达到及高于25导致胚胎发育减缓、孵化率显著降低(P<0.05),其中原肠期和出膜前期胚胎孵化率在盐度达到35时几乎为零,而眼点期胚胎孵化率在盐度达到25时已低于10%；从原肠期和出膜前期开始盐度暴露其初孵蚤状幼体在盐度5-25的范围内具有较高的存活率,且各盐度组之间无显著差异,超出此盐度范围后初孵蚤状幼体的存活率急剧降低(P<0.05),盐度达到35时初孵蚤状幼体无法存活。眼点期开始盐度暴露后其初孵蚤状幼体存活率在15、20盐度下最高,在盐度10中略降低,盐度高于20或低于10均导致初孵蚤状幼体存活率显著降低(P<0.05),在盐度1、30、35中几乎无初孵蚤状幼体存活。
Ⅰ期蚤状幼体在1、40的盐度下暴露24h后全部死亡,盐度5组蚤状幼体死亡率从暴露后48h开始显著上升,到192h时接近100%,盐度25、30、35组蚤状幼体死亡率在盐度暴露后96h仍较低,之后开始显著升高,到192h时几乎全部死亡,盐度10、15、20组蚤状幼体在盐度暴露后120h死亡率仍较低,从144h开始逐渐升高,在192h死亡率达到60%,但仍显著低于其它各盐度组(P<0.05)。
综合上述研究结果表明,中华绒螯蟹原肠期和出膜前期胚胎发育及孵化的适宜盐度范围为5-20,眼点期胚胎发育的适宜盐度范围为10-20,Ⅰ期蚤状幼体发育的适宜盐度范围为10-20,超出上述盐度范围将显著影响中华绒螯蟹胚胎与早期幼体的发育,进而可能对其种群的繁衍造成潜在影响。
6.渗透胁迫对中华绒螯蟹胚胎生化成分含量与消化酶活性的影响
研究了渗透胁迫对不同发育期中华绒螯蟹胚胎中生化成分(蛋白、总脂、碳水化合物、水份)含量、胚胎体积(胚胎直径)以及消化酶(淀粉酶、胃蛋白酶、胰蛋白酶、脂肪酶)活性的影响。结果显示：随着胚胎的发育,对照组(盐度15)胚胎体积增加、水份含量升高(P<0.05)、蛋白含量降低(P<0.05)、胰蛋白酶与胃蛋白酶活性升高(P>0.05),脂类含量和脂肪酶活性均降低(P>0.05),碳水化合物含量和淀粉酶活性均在眼点期升高(P<0.05)而在出膜前又降低(P<0.05)；高、低渗胁迫均引起胚胎水份含量降低,其中高渗胁迫对三个发育时期胚胎水份含量的影响均具有显著性(P<0.05),而低渗胁迫的影响仅在眼点期表现出显著性(P<0.05)；低渗胁迫引起胚胎蛋白含量降低(P<0.05),而高渗胁迫导致胚胎蛋白含量升高(P<0.05)；高渗胁迫对胚胎脂类含量无显著影响(P>0.05),而低渗胁迫引起出膜前期胚胎脂类含量显著降低(P<0.05)；高、低渗胁迫对胚胎碳水化合物含量均无显著影响(P>0.05)；高、低渗胁迫均导致原肠期胚胎体积减小(P>0.05),而对眼点期胚胎体积无显著影响,高渗胁迫引起出膜前期胚胎体积减小(P>0.05)；除高、低渗胁迫均显著降低眼点期胚胎淀粉酶活性(P<0.05)而引起出膜前期胚胎胰蛋白酶活性略降低(P>0.05)外,低渗胁迫导致不同发育时期胚胎消化酶活性升高,而高渗胁迫导致酶活性降低；除脂肪酶外,渗透胁迫对原肠期胚胎消化酶活性的影响均具有显著性(P<0.05)。研究结果表明,主要能源物质在胚胎发育过程中被利用并重新合成,高渗胁迫抑制胚胎消化酶的活性,进而抑制胚胎对卵黄物质的分解、利用,导致胚胎中卵黄物质积累,从而影响胚胎的发育；低渗胁迫引起胚胎消化酶活性升高,加速胚胎对卵黄物质的分解与利用,在一定程度上促进胚胎的发育。
7.渗透胁迫对中华绒螯蟹胚胎CHH和Na+/K+-ATPase a-亚基nRNA表达及酶活性的影响
研究了渗透胁迫对中华绒螯蟹不同发育期胚胎Na+/K+-ATPase活性、a-亚基mRNA表达、CHH基因mRNA表达的影响。结果显示：对照组(盐度15)胚胎中Na+/K+-ATPase a-亚基mRNA表达量从原肠期到出膜前期升高了1.77倍(P<0.05),Na+/K+-ATPase活性升高了6.53倍(P<0.05)；与对照组相比,低渗胁迫诱导出膜前期胚胎Na+/K+-ATPase a-亚基表达显著增强(P<0.05),而对原肠期和眼点期胚胎无显著影响；低渗胁迫引起原肠期与眼点期胚胎Na+/K+-ATPase活性显著降低(P<0.05)；高渗胁迫诱导原肠期与眼点期胚胎Na+/K+-ATPase a-亚基的表达显著增强(P<0.05),同时引起原肠期、出膜前期Na+/K+-ATPase活性显著升高(P<0.05),而导致眼点期Na+/K+-ATPase活性显著降低(P<0.05)；从眼点期到出膜前期胚胎CHH mRNA表达显著增强,低渗胁迫导致CHH mRNA表达下调且对眼点期胚胎具有显著影响(P<0.05),高渗胁迫诱导CHH上调且对出膜前期胚胎具有显著影响(P<0.05)。研究结果显示,随着胚胎的发育,CHH表达增强、Na+/K+-ATPase mRNA转录、蛋白合成增加,CHH可能在胚胎应对渗透胁迫时发挥着重要的神经内分泌调控作用,对渗透胁迫后Na+/K+-ATPase的转录、蛋白合成起到调节作用。
8.中华绒螯蟹大眼幼体的盐度选择行为
利用七分室盐度选择装置研究了中华绒螯蟹大眼幼体对盐度的选择行为,结果显示：大眼幼体对低盐表现出明显的偏好行为(P<0.05),且这种行为在上午表现更为显著,上午(9：00-12：00)进行的两个重复组中3h内大眼幼体在淡水分室中的平均分布率最高,分别为60%-89%和69%-80%,分布率次高的为盐度5分室；下午(14：00-17：00)进行的两个重复组中大眼幼体在淡水分室中的分布率分别为29%-60%和31%-57%。上、下午的试验结果均显示,随着盐度的升高,大眼幼体在盐度分室中的分布率呈现降低的趋势。结果表明,中华绒螯蟹大眼幼体具有明显的偏好淡水、低盐的行为,且这种偏好程度可能受到光照强度的影响,中华绒螯蟹大眼幼体的低盐偏好行为与其在自然环境下的向岸洄游生态相吻合。本试验结果提示,大眼幼体趋淡水、低盐的行为可能是一种遗传行为而不受栖息环境的影响。
9.盐度对中华绒螯蟹大眼幼体蜕壳及Na+/K+-ATPase活性的影响
研究了不同盐度(0、5、10、15、20、25、30、35、40)对中华绒螯蟹蜕壳的影响,并测定了高、低渗胁迫后大眼幼体Na+/K+-ATPase活性的变化。结果显示：各盐度组大眼幼体在盐度暴露后第二天开始出现蜕壳现象,之后随着试验时间的延长,大眼幼体蜕壳率急剧升高(P<0.05),到试验的第5天所有盐度组大眼幼体几乎全部蜕壳为Ⅰ期仔蟹,且不同盐度组大眼幼体的蜕壳率无显著差异(P>0.05),但盐度达到40时大眼幼体在前期的蜕壳速度明显减缓；高、低盐胁迫均显著诱导大眼幼体Na+/K+-ATPase活性升高(P<0.05)。结果表明,大眼幼体已具有较强的高、低渗调节能力,可以耐受较广范围的盐度,并在此盐度环境下完成蜕壳,Na+/K+-ATPase在其高、低渗调节过程中均发挥重要作用。
Chinese mitten crab Eriocheir sinensis H. Milne Edwards,1853(Decapoda, Brachyura) is a diadromous freshwater crab. Adult crabs spend most of their lives growing in rivers and lakes but must migrate downstream towards the estuary or sea for reproduction after their puberty molt, where they reach maturity, complete mating and embryonic development as well as planktonic larvae release. Estuary features drastically temporal and spatial variation in salinity. Yangtze River estuary is the biggest and vital breeding ground of E. sinensis in China. Recently, saltwater intrudes the Yangtze River estuary more and more frequently, and it has become the focus of researches. Remarkable salinity change might exert a negatively potential effect on the reproduction and development of E. sinensis.
To evaluate the possible adverse implications of salinity change to E. sinensis comprehensively, we monitored the salinity preference behavior and the behavioral response of Chinese mitten crab E. sinensis broodstock in different salinities, and low salinity preference behavior of megalopas. We investigated the osmotic and ionic regulation, variation of ion-transporting enzymes activities, haemolymph metabolic and immune parameters and variation of digestive enzyme activities in both females and males, when crabs were exposed to freshwater and acclimated to step-wise salinities medium ranging from7to35ppt. We also studied the effect of salinity on E. sinensis embryo and larva development, and determined the developmental changes of energy source, digestive enzymes and Na+/K7-ATPase activity, and CHH and Na+/K+-ATPase a-subunit expression in the eggs of E. sinensis under salinity stress. Results obtained from this study will enhance our comprehension of the ontogeny of crustaceas'osmoregulation, especially during embryo stage and reproductive stage. Furthermore, this information is also great important to protect natural resources and to propagate of Chinese mitten crab E. sinensis artificially. These results will also provide theoretical basis for evaluating the potential effect of salinity change on E. sinensis.
1. Behavioral responses of female Chinese mitten crab Eriocheir sinensis broodstock to salinities
Behavioral response of female E. sinensis broodstock were monitored after acclimating crabs to freshwater (0ppt) and brackish water (18ppt) and hyper-osmotic (increased salinity from18to30ppt) or hypo-osmotic (decreased salinity from18to 0ppt) environment abruptly, by the method of video recording analysis. Locomotor activity and the frequency of mouthparts movements increased after acclimating crabs to brackish water (P<0.05) and hypo-osmotic environment. However, the two kinds of behaviors decreased after hyper-osmotic stress. Behaviors that the antennules retracted into the carapace for periods of time and the mouthparts closed were discovered only in the control group of18ppt and hyper-osmotic stress group (P<0.05). There was a decrease of the times of antennae flicking with the increase of environmental salinity. Behavior of the antennae flicking frequently showed up after hypo-osmotic stress (P<0.05). Behavior of eyestalk movement decreased after acclimating crabs to brackish water and increased after salinity stress. E. sinensis cleaned the antennae and antennules with the palps of the third maxillipeds. This behavior increased after gradual increase of environmental salinity, comparing with the response of crabs maintained in freshwater (P<0.05). Behavior of cleaning the antennae and antennules decreased after hypo-hyper osmotic stress (P<0.05). The abdomen extension behavior was observed only after hypo-osmotic stress. The results suggested that the increase of salinity induced the locomotor activity and other related behaviors of E. sinensis broodstock. Abdomen extension under hypo-osmotic stress could directly expose the hindgut to the water, and function as an additional means of ion uptake in lower salinity. Behavior of closure reaction under hyper-osmotic stress could help the reduction of water absorption and salt loss in high salinity environment.
2. Salinity preference behavior of Chinese mitten crab Eriocheir sinensis broodstock
Preference chambers are widely used to measure the responses of aquatic organisms to different environmental gradients, such as salinity, temperature and pollutants. A six-chambered device was designed to investigate the salinity preference of downstream-migrating Chinese mitten crab, Eriocheir sinensis. Occurrence numbers and retention time within6h are as indicators for evaluating the preference of E. sinensis to salinity. Males did not show preference for salinity. However, the occurrence number was the least at28ppt salinity chamber while was the most at35ppt salinity chamber. Males spend slightly more time in the chambers of21ppt and35ppt than others salinity chambers. Females occurred in the salinity chambers more and more frequently with the increase of salinity at a range0-21ppt. Occurrence number of females in chamber with21ppt was highest while lowest with28ppt. Females spends less and less time in salinity chambers with the increase of salinity at a range of0-28ppt. However, they retained in35ppt salinity chamber for the longest time. Males changed their positions more frequently than females (P<0.05). After offering different salinities in six chambers, locomotor activities of the crabs were weakening. Different salinity preference behaviors presented between females and males might be related to the asynchronism in gonadal development and the effect of salinity to their gonadal development. It is indicated that other factors such as water depth, substrate could be also vital to control their distribution in estuary.
3. Osmotic and ionic regulation and Na+/K+-ATPase, carbonic anhydrase activities in Chinese mitten crab Eriocheir sinensis broodstck exposed to different salinities
Chinese mitten crabs broodstock, are exposed to brackish water or seawater as an obligatory part of their reproductive migration. We investigated the changes in haemolymph osmolality, haemolymph Na+, Cl-, Ca2+and K+concentrations, as well as gill Na+/K+-ATPase (NKA) and carbonic anhydrase (CA) activity in both females and males, when in freshwater (0ppt,8mOsm (kg H2O)-1) and after step-wise acclimation to salinities ranging from7to35ppt (175-1044mOsm (kg H2O)-1). Both females and males strongly hyper-regulated their haemolymph osmolality over salinity range of0-21ppt, the isosmotic point approaching28ppt, and slightly hypo-regulated at35ppt. Haemolymph Na+and Cl-levels correlated positively with haemolymph osmolality. Ca2+and K+levels, however, were maintained at a relatively narrow range and showed a contrary trend in their response to salinity. This indicated that Na+and Cl-are the main contributors to increasing haemolymph osmolality. Reduced Na+/K+-ATPase and carbonic anhydrase activity in the posterior gills at elevated salinity suggest a direct physiological response to a decrease of Na+intake. No sexual differences existed for all parameters except for significantly higher haemolymph Ca2+and K+levels and CA activity in females than in males at some salinity levels. It is suggested that Eriocheir sinensis broodstock is a strong hyperosmoregulator but a weak hypo-osmoregulator. No sexual differences appeared to exist in osmotic and ionic regulation of Eriocheir sinensis broodstock exposed to different salinities.
4. Metabolic, immune and digestive response of Chinese mitten crab Eriocheir sinensis broodstock exposed to different salinities
To understand the metabolic, immune and digestive adjustments of Chinese mitten crab broodstock, Eriocheir sinensis, during their reproductive migration from freshwater to seawater, concentrations haemolymph metabolic variations (protein, glucose, triglycerides, cholesterol, urea, uric acid and creatinine) and activities of immune enzyme (alkaline phosphatase, superoxide dismutase and phenoloxidase) and digestive enzyme were determined after gradual acclimating of females and males from freshwater (0ppt) to different salinities (7,14,21,28,35ppt) medium. The results showed that increasing salinities caused a gradual decrease of hemolymph total protein and oxyhemocyanin contents of females and males. Some haemolymph metabolic variables reached a minimum at21ppt, and increased with both increase and decrease of the medium salinity (triglycerides, cholesterol, urea, creatinine in males and triglycerides, uric acid in females), or elevated either with the increase (urea in females) or decrease (uric acid in males) of medium salinities. Responses of haemolymph metabolic variations to salinity slightly differ between females and males. Hemolymph alkaline phosphatase activity increased slightly in7ppt; thereafter it dropped slowly until the end of experiment. Hemolymph phenoloxidase activity of females and males decreased gradually with the increase of environmental salinity. However, it maintained a stable level when the salinities were above21ppt in females. Hemolymph superoxide dismutase activity of males decreased with the increase of the medium salinity until21ppt and increased when the medium salinity above21ppt. All tested immune parameters except for SOD of females were significantly higher than those of males. Digestive enzyme activity except lipase of males were significantly higher than those of females at some salinities (P<0.05). Amylase, pepsin and trypsin activities of females reduced significantly when the salinity reached28ppt (P<0.05). Cellulase activity decreased when increasing the salinity to35(P<0.05). Remarkable decrease of males'digestive enzyme activities except lipase occurred at21and higher salinity, and an increase was shown in14ppt. The present results indicated that salinity of21ppt was the turning point of several biochemical parameters in E. sinensis broodstock. This salinity could be more'profitable'for E. sinensis broodstock from the energy-saving point of view. Immune and digestive enzyme activities of E. sinensis broodstock will be affected when the salinity was near to or above28ppt, which could exert a negative effect on reproduction. Metabolic and immune parameters in the haemolymph of females were slightly higher than those of males at some salinity levels, this suggestes higher metabolic level and greater immunity. The initial salinity that induced the decrease of digestive enzyme activity in males was lower than that in females; this result indicated that females were more tolerant elevated salinities than males in the digestive ability. This study might help understand some aspects of the metabolic, digestive and immune adjustments in euryhaline crabs related to osmoregulation.
5. Survival and development of Chinese mitten crab, Eriocheir sinensis embryo and Zoea in different salinities
Embryo and larval development of Chinese mitten crab. Eriocheir sinensis, is completed in the estuary, where they are often exposed to extreme salinity conditions. To test whether these salinity conditions have negative impacts on early development of embryo and zoea or not, we determined hatching rate, development time and survival rate when E. sinensis embryo for the gastrula, eyespot and pre-hatching stage were exposed to a salinity range of1-35ppt, and we also detected the survival rate of E. sinensis zoea I within192h later exposing them to different salinities. Reduced salinities (1and5ppt) accelerated embryonic development and increased hatching rate of eyespot and pre-hatching stage, but reduced salinities did not exert any effects on the gastrula stage development. Hatching rate significantly declined when the medium salinity reached at and above25ppt. No zoea I were hatched when eyespot stages embryos were exposed to30ppt and three stages embryos were exposed to35ppt. Higher survival rate of initial zoea I were presented when eyespot and pre-hatching stage embryos were exposed to salinity ranges of5-25ppt. However, survival rate of initial zoea I significantly declined when embryos were exposed to1and30ppt. All zoeas hardly survived at35ppt. Nearly40%zoeas I still survived until192h later during the range10-20ppt of salinity. However, all zoea I died24h later after exposure to salinity1and40ppt while died192h later at salinity range25-35and5ppt. The results showed that optimal salinity range was5-20ppt, when embryos began to be exposed to different salinities from gastrula stage and pre-hatching stage, while the salinity range became narrower (10-20ppt) for eyepot satge. The optimal salinity range was10-20ppt for zoea development. Out of above salinity ranges, will influence the development of embryo and larva, and then affect the wild population' reproduction.
6. Developmental changes of biochemical composition and digestive enzyme activity in the eggs of Chinese mitten crab, Eriocheir sinensis, under osmotic stress
Biochemical compositions (total carbohydrate, soluble protein, lipid and water content) contents, digestive enzymes (amylase, lipase, pepsin and trypsin) activity and embryo volume (egg diameter) have been determined in three developmental stages (gastrula, eyespot and pre-hatching stage) of E. sinensis eggs under hyper or hypo-osmotic stress. Increase of water contents (P<0.05), eggs diameter and pepsin and trypsin activities and decrease of soluble protein (P<0.05) and lipid contents and lipase activity (P<0.05) in eggs were noticed during the development of embryo. Total carbohydrate content and amylase activity increased at the eyespot stage (P<0.05), whereas they reduced during the prehatching stages (P<0.05). Hyper-osmotic (P<0.05) and hypo-osmotic stress reduced water content of eggs at each developmental stage. Although hyper-osmotic stress reduced the water content of eggs, significant effect presented only at eyespot stage (P<0.05). Soluble protein content of three developmental stages eggs decreased under hyper-osmotic stress while increased under hypo-osmotic stress (P<0.05). Total lipid content was unaffected by hyper-osmotic stress at any developmental stage, whereas it reduced significantly at pre-hatching stages under hypo-osmotic stress (P<0.05). Total carbohydrate contents did not show any significant variation after osmotic stress. Both hypo-osmotic and hyper-osmotic stress induced the decrease of egg diameters during the gastrula stage (P<0.05), however, no significant effect on eyespot stage. Egg diameter decreased before hatching, only under the hyper-osmotic stress (P<0.05). Among the four digestive enzymes assayed, amylase activities were the highest while lipase activities were the lowest. Hyper-osmotic stress reduced enzymes activities of the three developmental stage eggs, while hypo-osmotic stress enhanced the activities of digestive enzymes with the exception of reducing the amylase activities in eyespot stage (P<0.05) and trypsin activities in pre-hatching stage (P>0.05). Effect of salinity change on digestive enzymes is statically significant only during gastrula stage (P<0.05) except for lipase. It is suggested that salinity stress changed the activity of digestive enzymes which further affectes the utilizing of yolk substances, consequently, affects embryonic development. Dramatical variation of salinity in estuary ought to be having potential effect on embryonic development, especially induced by salt water encroachment.
7. Developmental changes of mRNA expression and activity of Na+/K+-ATPase, and expression of crustacean hyperglycemic hormone (CHH) gene in Chinese mitten crab Eriocheir sinensis eggs under osmotic stress
We examined ontogenetic and salinity-induced changes of Na+/K+-ATPase activity and mRNA expression of the a-subunit. We also detected the expression of crustacean hyperglycemic hormone (CHH) neuropeptides. which involved directly or indirectly in ionic and osmotic regulation, in the eyespot stage and pre-hatching stage. All activities and mRNA expression were determined24h later after directly exposeing embryos to1,15(control group), and35ppt. NKA activities were low in gastrula stage, and the enzyme activity increased6.53times in pre-hatching stage. Na+/K+-ATPase activity significantly decreased in gastrula and eyespot stages (P<0.05), and slightly reduced in pre-hatching stage at1ppt compared with15ppt, whereas the activity in the embryo of gastrula and pre-hatching stages was increased at salinity30ppt than at15and1ppt. However, increasing salinity reduced the enzyme activity (P<0.05) at eyespot stage, compared with the control group. The Na+/K+-ATPase a-subunit gene was expressed and elevated1.17times from gastrula stage to pre-hatching stage. There was no significant difference of Na+/K+-ATPase a-subunit gene expression between gastrula and eyespot stages, but expression increased remarkably before hatching (P<0.05). The expression of Na+/K+-ATPase a-subunit gene increased at30and1ppt compared with15ppt, but significant difference occurred only at eyepot stage under hyper salinity stress (P<0.05). Expression of CHH gene significantly increased with the development of embryo from eyespot stage to pre-hatching stage at all salinities (P<0.05), the expression remarkably elevated before hatching under hyper salinity stress while significantly decreased at eyespot stage under hypo salinity stress. It is suggested that although transcription and protein synthesis of Na+/K+-ATPase increased with the development of embryo, no consistent pattern between them under salinity stress. Neuroendocrine hormone CHH might be play important regulatory function in osmoregulation of embryo under osmotic stress.
8. Low salinity preference behavior of Chinese mitten crab Eriocheir sinensis megalopae
Up-estuary migration of crab larvae to adult habitats is thought to be started after molt to megalopae. However, the mechanisms used by crab megalopae for reinvasion of the estuary are unclear. We investigated the salinity preference behavior of E. sinensis megalopae in a seven choice-chambers device, to determine whether they showed preference behavior for the low salinity of the water or not. If low-salinity preference exists, this might be one of the bebavioral mechanisms for reinvasion estuary. Notable salinity gradients can be available in the seven choice-chambers device. Results revealed that most megalopa (60%-89%and69%-80%in two replicates, respectively) selected nominal0ppt as their final preference during9:00-12:00am. However, this behavioral weaken slightly (29%-60%and31%-57%megalopa prefer to nominal0ppt in two replicates, respectively) during14:00-17:00pm. Our findings reveal that E. sinensis megapolas obviously prefer low salinities, which associated with their strong osmoregulatory ability. Low-salinity preference behavior might be an important mechanism for megalopa's up-estuary migration. However, the extent of preference behavioral seems to be influenced by light. We speculated that the low-salinity preference behavior of megalopa is a hereditary behavioral while unaffected by environmental change. The importance of low-salinity preferences in relation to the megapola's up-estuary migration will be discussed in this study.
9. Effect of salinity on molt and Na+/K+-ATPase activity of E. sinensis megalopa
We determinted the percentage of megalopa molt to first juvenile instars under the condition of salinity range of1-40ppt, and detected the Na+/K+-ATPase activities of total megalopas after directly exposing them to1,15ppt (control group), and30ppt salinity. The result showed that nearly all the megalopas molted to the first juvenile instars at salinities range of1-40ppt on the fifth day of experiment, whereas molt delayed significantly at40ppt, compared with other salinities. Na+/K+-ATPase activity increased remarkably either at low salinity (1ppt) or at high salinity (30ppt) medium, compared with control group. The results suggest that Eriocheir sinensis megalopas possess moderate hyper-/hypo-regulating ability, and the osmoregulatory ability was also exhibited by the change of Na+/K+-ATPase activity under salinity stress. The results of this study provide physical evidence for comprehensing the up-estuary migration of crab larvae to adult habitats.
为全面评估河口盐度变动对中华绒螯蟹繁育的潜在影响,本研究以中华绒螯蟹亲体、后期胚胎及早期幼体为研究对象,从血淋巴渗透、代谢、免疫调节与肝胰腺消化酶调节以及行为方面研究了中华绒螯蟹亲体在洄游过程中对盐度的适应策略及耐盐阂值；以孵化率与初孵蚤状幼体的存活率为评价指标获得中华绒螯蟹胚胎及早期幼体发育的最适盐度范围,在此基础上分析了盐度胁迫后胚胎对能源物质的分解与利用特征、Na+、K+-ATPase的响应以及胰高血糖素(CHH)可能的神经内分泌调控作用；通过研究大眼幼体的盐度选择行为,不同盐度下的生态蜕壳及Na+/K+-ATPase活性对盐度的响应,探讨大眼幼体趋江洄游的行为与生理基础,初步阐明了不同生命阶段的中华绒螯蟹对盐度的生理与行为响应机制。本研究成果为全面评估河口盐度变动对中华绒螯蟹繁育的影响提供重要理论依据,对中华绒螯蟹野生资源的保护、利用和人工繁殖均具有重要意义,同时也丰富了甲壳类洄游生理学与渗透调节生理、行为研究的资料。主要研究结果如下：
1.中华绒螯蟹亲体在不同盐度下的行为反应
采用视频记录分析法比较研究了雌性中华绒螯蟹亲体适应了淡水、半咸水(盐度18)后以及在高盐(盐度18→30)、低盐(盐度18→0)胁迫下的行为反应。结果表明：在不同盐度下,蟹口器的活动性与其个体的运动活力呈现一致的反应模式,与淡水组相比,适应盐度18后蟹的运动活力和口器的活动性均增强(P<0.05)；低盐胁迫引起蟹运动活力和口器活动(P<0.05)增强,而高盐胁迫后蟹运动活力显著减弱(P<0.05),且口器的活动性也降低,且在低盐胁迫后蟹的运动活力和口器的活动性均显著强于淡水组(P<0.05)；第一触角回缩行为和封闭反应行为几乎仅出现在盐度18组以及高盐胁迫组(P<0.05),第一触角回缩至壳槽内的时间随着试验时间的延长而增加；第二触角抖动次数随着试验盐度的升高而逐渐降低,低盐胁迫后蟹的第二触角抖动次数显著高于其它三个试验组(P<0.05)；高、低盐胁迫均引起蟹的眼柄活动性增强,而适应盐度18后眼柄的活动性降低；盐度18组蟹清洁触角的频率显著高于淡水组(P<0.05),高、低盐胁迫均引起蟹的触角清洁频率显著降低(P<0.05)；腹部开合行为仅出现在低盐胁迫后。综合以上结果显示,从淡水适应半咸水后会刺激中华绒螯蟹亲体的运动活力及其它相关行为增强,蟹在低盐胁迫后出现腹部开合行为是其在低渗环境下通过暴露尾肠从而吸收水中离子的一种行为策略,蟹在高盐胁迫后表现出的封闭反应则有助于减少机体在高渗环境下对水分的吸收及盐分的被动扩散丢失。
2.中华绒螯蟹亲体的盐度选择行为
利用室内可控的六分室盐度选择装置结合视频监控系统,研究了洄游期的雌、雄中华绒螯蟹对盐度的选择行为。结果显示：雄蟹对盐度并未表现出显著的偏好行为,但其在盐度28分室中出现的次数最少,而在盐度35分室中出现最多,在21与35盐度分室中停留的时间略长于其它盐度分室；在盐度0-21范围内,雌蟹在各盐度分室中出现的次数随着盐度的升高而增加,在21盐度分室中达到最高,在28盐度分室中最低,雌蟹在35盐度分室中出现的次数接近盐度21分室的水平。在盐度7-28范围内,雌蟹在各盐度分室中停留的时间随着盐度的升高逐渐降低,而在35盐度分室中停留时间最长,且显著高于其在28盐度分室中停留时间(P<0.05)。本试验同时发现,在试验时间内,雄蟹的运动活力显著高于雌蟹,在提供可选择的盐度环境后,雌、雄蟹的运动活力均降低。结果提示,盐度在中华绒螯蟹生殖洄游寻找产卵场的过程中虽然发挥着一定的导向作用,但其它栖息地环境因子如水深、底质可能也是影响中华绒螯蟹在河口分布位置的重要因子。
3.盐度升高对中华绒螯蟹亲体渗透压、离子及离子调节酶的影响
通过逐步增盐法研究了中华绒螯蟹亲体盐度适应过程中血淋巴渗透压、离子含量以及后鳃Na+/K+-ATPase和碳酸酐酶(CA)活性的变化。结果显示：在0-21的盐度范围内,雌、雄中华绒螯蟹均显著调高其血淋巴渗透压(P<0.05),而在盐度达到35时略调低其血淋巴渗透压,雌、雄中华绒螯蟹的等渗点接近盐度28；随着环境水体盐度的增加和血淋巴渗透压的升高,中华绒螯蟹血淋巴Na+和Cl-含量升高,而Ca2+和K+含量呈现降低的趋势；盐度升高引起中华绒螯蟹后鳃Na+/K+-ATPase和CA活性降低,酶活性的降低是后鳃主动吸收Na+减少的直接生理反应；比较雌、雄中华绒螯蟹各检测指标可见,雌蟹血淋巴Ca2+和K+含量和鳃CA活性在一些盐度水平下显著高于雄蟹(P<0.05),而其它测定指标之间无显著的性别差异。研究结果表明,中华绒螯蟹亲体具有较强的高渗调节能力,但其低渗调节能力较弱。盐度升高后,中华绒螯蟹亲体主要通过调节血淋巴Na+和Cl-含量而调节血淋巴渗透浓度,Na+/K+-ATPase和CA在离子调节过程中发挥重要作用,雌、雄蟹的渗透调节能力不存在显著的差异。
4.盐度升高对中华绒螯蟹亲体血淋巴代谢、免疫指标及肝胰腺消化酶活性的影响
通过逐步增盐法,研究了从淡水逐步升高到盐度7、14、21、28、35后,雌、雄中华绒螯蟹血淋巴代谢指标(总蛋白蛋白、氧合血蓝蛋白、葡萄糖、甘油三酯、胆固醇、尿酸、尿素、肌酐)的含量、免疫酶(碱性磷酸酶、超氧化物歧化酶、酚氧化酶)和消化酶(淀粉酶、纤维素酶、胃蛋白酶、胰蛋白酶、脂肪酶)活性的变化。结果显示：随着盐度的增加,雌、雄中华绒螯蟹血淋巴总蛋白、氧合血蓝蛋白含量降低；雄蟹血淋巴甘油三酯、胆固醇、尿素、肌酐含量与雌蟹血淋巴甘油三酯、尿酸含量均随着盐度的升高先显著降低(P<0.05),在盐度升高到21时达到最低值,而后显著升高(P<0.05)；雄蟹血淋巴尿酸含量在0-21的盐度范围内呈现显著降低的趋势(P<0.05),盐度高于21时维持在稳定水平；雌蟹血淋巴尿素含量在盐度升高到21达到最低值(P>0.05),盐度高于21时显著升高(P<0.05)；盐度增加对雌蟹血淋巴肌酐、胆固醇含量无显著影响；雌、雄蟹血糖含量均在盐度升高到21时达到最高水平；雌蟹血淋巴多种代谢指标均高于雄蟹。
随着盐度的升高,雄蟹血淋巴酚氧化酶活力逐渐降低,雌蟹血淋巴酚氧化酶活力先降低,当盐度高于21后维持稳定水平；雌、雄蟹血淋巴碱性磷酸酶活性呈现先略升高而后逐渐降低的趋势；雌、雄蟹血淋巴超氧化物歧化酶活性先略降低,盐度高于21时被激活,且在盐度升高到35时酶活性显著高于盐度21时的水平(P<0.05)。除超氧化物歧化酶外,雌蟹血淋巴其它两种免疫酶活性均高于雄蟹。
雌、雄中华绒螯蟹肝胰腺淀粉酶活性最高,脂肪酶活性极低(P<0.05)；雌蟹肝胰腺淀粉酶、胃蛋白酶、胰蛋白酶活性均在盐度达到及高于28时显著降低(P<0.05),而纤维素酶活性在盐度升高到35时显著降低(P<0.05)；除脂肪酶外,雄蟹的肝胰腺消化酶活性均在盐度升高到14时显著升高(P<0.05),盐度升高到21时显著降低(P<0.05),之后维持稳定水平或进一步降低；除脂肪酶外,雄蟹肝胰腺消化酶活性在不同盐度水平(盐度21除外)下均显著高于雌蟹(P<0.05)。
综合上述研究结果显示：中华绒螯蟹亲体从淡水逐步适应海水的过程中多种血淋巴代谢指标均在盐度增加到21时达到最低点,这一盐度下机体可能用于渗透调节的能量最低,从而可以节约更多的能量用于繁殖。当水体盐度接近或超过28时中华绒螯蟹的免疫防御能力和消化能力均显著降低,最终可能对中华绒螯蟹的繁殖造成潜在的不利影响。在不同盐度下,雌蟹血淋巴代谢指标和免疫指标稍高于雄蟹,表明雌蟹的代谢水平和免疫防御能力相对较高。引起雄蟹肝胰腺消化酶活性降低的起始盐度值低于雌蟹,这从消化生化的角度表明雌蟹对盐度增加的适应能力强于雄蟹。
5.盐度对中华绒螯蟹胚胎及幼体发育的影响
研究了盐度对不同发育时期中华绒螯蟹离体胚胎发育、蚤状幼体存活的影响,结果显示：从眼点期开始盐度暴露其胚胎孵化率与幼体存活率均低于从原肠期和出膜前期开始暴露胚胎的孵化率,原肠期和出膜前期胚胎在盐度1-20范围内呈现较高的孵化率和较快的发育速度,且各盐度组间无显著差异(P>0.05)；盐度降低(盐度1、5)在一定程度上促进了胚胎的发育并提高了胚胎孵化率,且在眼点期表现出显著性(P<0.05)；盐度达到及高于25导致胚胎发育减缓、孵化率显著降低(P<0.05),其中原肠期和出膜前期胚胎孵化率在盐度达到35时几乎为零,而眼点期胚胎孵化率在盐度达到25时已低于10%；从原肠期和出膜前期开始盐度暴露其初孵蚤状幼体在盐度5-25的范围内具有较高的存活率,且各盐度组之间无显著差异,超出此盐度范围后初孵蚤状幼体的存活率急剧降低(P<0.05),盐度达到35时初孵蚤状幼体无法存活。眼点期开始盐度暴露后其初孵蚤状幼体存活率在15、20盐度下最高,在盐度10中略降低,盐度高于20或低于10均导致初孵蚤状幼体存活率显著降低(P<0.05),在盐度1、30、35中几乎无初孵蚤状幼体存活。
Ⅰ期蚤状幼体在1、40的盐度下暴露24h后全部死亡,盐度5组蚤状幼体死亡率从暴露后48h开始显著上升,到192h时接近100%,盐度25、30、35组蚤状幼体死亡率在盐度暴露后96h仍较低,之后开始显著升高,到192h时几乎全部死亡,盐度10、15、20组蚤状幼体在盐度暴露后120h死亡率仍较低,从144h开始逐渐升高,在192h死亡率达到60%,但仍显著低于其它各盐度组(P<0.05)。
综合上述研究结果表明,中华绒螯蟹原肠期和出膜前期胚胎发育及孵化的适宜盐度范围为5-20,眼点期胚胎发育的适宜盐度范围为10-20,Ⅰ期蚤状幼体发育的适宜盐度范围为10-20,超出上述盐度范围将显著影响中华绒螯蟹胚胎与早期幼体的发育,进而可能对其种群的繁衍造成潜在影响。
6.渗透胁迫对中华绒螯蟹胚胎生化成分含量与消化酶活性的影响
研究了渗透胁迫对不同发育期中华绒螯蟹胚胎中生化成分(蛋白、总脂、碳水化合物、水份)含量、胚胎体积(胚胎直径)以及消化酶(淀粉酶、胃蛋白酶、胰蛋白酶、脂肪酶)活性的影响。结果显示：随着胚胎的发育,对照组(盐度15)胚胎体积增加、水份含量升高(P<0.05)、蛋白含量降低(P<0.05)、胰蛋白酶与胃蛋白酶活性升高(P>0.05),脂类含量和脂肪酶活性均降低(P>0.05),碳水化合物含量和淀粉酶活性均在眼点期升高(P<0.05)而在出膜前又降低(P<0.05)；高、低渗胁迫均引起胚胎水份含量降低,其中高渗胁迫对三个发育时期胚胎水份含量的影响均具有显著性(P<0.05),而低渗胁迫的影响仅在眼点期表现出显著性(P<0.05)；低渗胁迫引起胚胎蛋白含量降低(P<0.05),而高渗胁迫导致胚胎蛋白含量升高(P<0.05)；高渗胁迫对胚胎脂类含量无显著影响(P>0.05),而低渗胁迫引起出膜前期胚胎脂类含量显著降低(P<0.05)；高、低渗胁迫对胚胎碳水化合物含量均无显著影响(P>0.05)；高、低渗胁迫均导致原肠期胚胎体积减小(P>0.05),而对眼点期胚胎体积无显著影响,高渗胁迫引起出膜前期胚胎体积减小(P>0.05)；除高、低渗胁迫均显著降低眼点期胚胎淀粉酶活性(P<0.05)而引起出膜前期胚胎胰蛋白酶活性略降低(P>0.05)外,低渗胁迫导致不同发育时期胚胎消化酶活性升高,而高渗胁迫导致酶活性降低；除脂肪酶外,渗透胁迫对原肠期胚胎消化酶活性的影响均具有显著性(P<0.05)。研究结果表明,主要能源物质在胚胎发育过程中被利用并重新合成,高渗胁迫抑制胚胎消化酶的活性,进而抑制胚胎对卵黄物质的分解、利用,导致胚胎中卵黄物质积累,从而影响胚胎的发育；低渗胁迫引起胚胎消化酶活性升高,加速胚胎对卵黄物质的分解与利用,在一定程度上促进胚胎的发育。
7.渗透胁迫对中华绒螯蟹胚胎CHH和Na+/K+-ATPase a-亚基nRNA表达及酶活性的影响
研究了渗透胁迫对中华绒螯蟹不同发育期胚胎Na+/K+-ATPase活性、a-亚基mRNA表达、CHH基因mRNA表达的影响。结果显示：对照组(盐度15)胚胎中Na+/K+-ATPase a-亚基mRNA表达量从原肠期到出膜前期升高了1.77倍(P<0.05),Na+/K+-ATPase活性升高了6.53倍(P<0.05)；与对照组相比,低渗胁迫诱导出膜前期胚胎Na+/K+-ATPase a-亚基表达显著增强(P<0.05),而对原肠期和眼点期胚胎无显著影响；低渗胁迫引起原肠期与眼点期胚胎Na+/K+-ATPase活性显著降低(P<0.05)；高渗胁迫诱导原肠期与眼点期胚胎Na+/K+-ATPase a-亚基的表达显著增强(P<0.05),同时引起原肠期、出膜前期Na+/K+-ATPase活性显著升高(P<0.05),而导致眼点期Na+/K+-ATPase活性显著降低(P<0.05)；从眼点期到出膜前期胚胎CHH mRNA表达显著增强,低渗胁迫导致CHH mRNA表达下调且对眼点期胚胎具有显著影响(P<0.05),高渗胁迫诱导CHH上调且对出膜前期胚胎具有显著影响(P<0.05)。研究结果显示,随着胚胎的发育,CHH表达增强、Na+/K+-ATPase mRNA转录、蛋白合成增加,CHH可能在胚胎应对渗透胁迫时发挥着重要的神经内分泌调控作用,对渗透胁迫后Na+/K+-ATPase的转录、蛋白合成起到调节作用。
8.中华绒螯蟹大眼幼体的盐度选择行为
利用七分室盐度选择装置研究了中华绒螯蟹大眼幼体对盐度的选择行为,结果显示：大眼幼体对低盐表现出明显的偏好行为(P<0.05),且这种行为在上午表现更为显著,上午(9：00-12：00)进行的两个重复组中3h内大眼幼体在淡水分室中的平均分布率最高,分别为60%-89%和69%-80%,分布率次高的为盐度5分室；下午(14：00-17：00)进行的两个重复组中大眼幼体在淡水分室中的分布率分别为29%-60%和31%-57%。上、下午的试验结果均显示,随着盐度的升高,大眼幼体在盐度分室中的分布率呈现降低的趋势。结果表明,中华绒螯蟹大眼幼体具有明显的偏好淡水、低盐的行为,且这种偏好程度可能受到光照强度的影响,中华绒螯蟹大眼幼体的低盐偏好行为与其在自然环境下的向岸洄游生态相吻合。本试验结果提示,大眼幼体趋淡水、低盐的行为可能是一种遗传行为而不受栖息环境的影响。
9.盐度对中华绒螯蟹大眼幼体蜕壳及Na+/K+-ATPase活性的影响
研究了不同盐度(0、5、10、15、20、25、30、35、40)对中华绒螯蟹蜕壳的影响,并测定了高、低渗胁迫后大眼幼体Na+/K+-ATPase活性的变化。结果显示：各盐度组大眼幼体在盐度暴露后第二天开始出现蜕壳现象,之后随着试验时间的延长,大眼幼体蜕壳率急剧升高(P<0.05),到试验的第5天所有盐度组大眼幼体几乎全部蜕壳为Ⅰ期仔蟹,且不同盐度组大眼幼体的蜕壳率无显著差异(P>0.05),但盐度达到40时大眼幼体在前期的蜕壳速度明显减缓；高、低盐胁迫均显著诱导大眼幼体Na+/K+-ATPase活性升高(P<0.05)。结果表明,大眼幼体已具有较强的高、低渗调节能力,可以耐受较广范围的盐度,并在此盐度环境下完成蜕壳,Na+/K+-ATPase在其高、低渗调节过程中均发挥重要作用。
Chinese mitten crab Eriocheir sinensis H. Milne Edwards,1853(Decapoda, Brachyura) is a diadromous freshwater crab. Adult crabs spend most of their lives growing in rivers and lakes but must migrate downstream towards the estuary or sea for reproduction after their puberty molt, where they reach maturity, complete mating and embryonic development as well as planktonic larvae release. Estuary features drastically temporal and spatial variation in salinity. Yangtze River estuary is the biggest and vital breeding ground of E. sinensis in China. Recently, saltwater intrudes the Yangtze River estuary more and more frequently, and it has become the focus of researches. Remarkable salinity change might exert a negatively potential effect on the reproduction and development of E. sinensis.
To evaluate the possible adverse implications of salinity change to E. sinensis comprehensively, we monitored the salinity preference behavior and the behavioral response of Chinese mitten crab E. sinensis broodstock in different salinities, and low salinity preference behavior of megalopas. We investigated the osmotic and ionic regulation, variation of ion-transporting enzymes activities, haemolymph metabolic and immune parameters and variation of digestive enzyme activities in both females and males, when crabs were exposed to freshwater and acclimated to step-wise salinities medium ranging from7to35ppt. We also studied the effect of salinity on E. sinensis embryo and larva development, and determined the developmental changes of energy source, digestive enzymes and Na+/K7-ATPase activity, and CHH and Na+/K+-ATPase a-subunit expression in the eggs of E. sinensis under salinity stress. Results obtained from this study will enhance our comprehension of the ontogeny of crustaceas'osmoregulation, especially during embryo stage and reproductive stage. Furthermore, this information is also great important to protect natural resources and to propagate of Chinese mitten crab E. sinensis artificially. These results will also provide theoretical basis for evaluating the potential effect of salinity change on E. sinensis.
1. Behavioral responses of female Chinese mitten crab Eriocheir sinensis broodstock to salinities
Behavioral response of female E. sinensis broodstock were monitored after acclimating crabs to freshwater (0ppt) and brackish water (18ppt) and hyper-osmotic (increased salinity from18to30ppt) or hypo-osmotic (decreased salinity from18to 0ppt) environment abruptly, by the method of video recording analysis. Locomotor activity and the frequency of mouthparts movements increased after acclimating crabs to brackish water (P<0.05) and hypo-osmotic environment. However, the two kinds of behaviors decreased after hyper-osmotic stress. Behaviors that the antennules retracted into the carapace for periods of time and the mouthparts closed were discovered only in the control group of18ppt and hyper-osmotic stress group (P<0.05). There was a decrease of the times of antennae flicking with the increase of environmental salinity. Behavior of the antennae flicking frequently showed up after hypo-osmotic stress (P<0.05). Behavior of eyestalk movement decreased after acclimating crabs to brackish water and increased after salinity stress. E. sinensis cleaned the antennae and antennules with the palps of the third maxillipeds. This behavior increased after gradual increase of environmental salinity, comparing with the response of crabs maintained in freshwater (P<0.05). Behavior of cleaning the antennae and antennules decreased after hypo-hyper osmotic stress (P<0.05). The abdomen extension behavior was observed only after hypo-osmotic stress. The results suggested that the increase of salinity induced the locomotor activity and other related behaviors of E. sinensis broodstock. Abdomen extension under hypo-osmotic stress could directly expose the hindgut to the water, and function as an additional means of ion uptake in lower salinity. Behavior of closure reaction under hyper-osmotic stress could help the reduction of water absorption and salt loss in high salinity environment.
2. Salinity preference behavior of Chinese mitten crab Eriocheir sinensis broodstock
Preference chambers are widely used to measure the responses of aquatic organisms to different environmental gradients, such as salinity, temperature and pollutants. A six-chambered device was designed to investigate the salinity preference of downstream-migrating Chinese mitten crab, Eriocheir sinensis. Occurrence numbers and retention time within6h are as indicators for evaluating the preference of E. sinensis to salinity. Males did not show preference for salinity. However, the occurrence number was the least at28ppt salinity chamber while was the most at35ppt salinity chamber. Males spend slightly more time in the chambers of21ppt and35ppt than others salinity chambers. Females occurred in the salinity chambers more and more frequently with the increase of salinity at a range0-21ppt. Occurrence number of females in chamber with21ppt was highest while lowest with28ppt. Females spends less and less time in salinity chambers with the increase of salinity at a range of0-28ppt. However, they retained in35ppt salinity chamber for the longest time. Males changed their positions more frequently than females (P<0.05). After offering different salinities in six chambers, locomotor activities of the crabs were weakening. Different salinity preference behaviors presented between females and males might be related to the asynchronism in gonadal development and the effect of salinity to their gonadal development. It is indicated that other factors such as water depth, substrate could be also vital to control their distribution in estuary.
3. Osmotic and ionic regulation and Na+/K+-ATPase, carbonic anhydrase activities in Chinese mitten crab Eriocheir sinensis broodstck exposed to different salinities
Chinese mitten crabs broodstock, are exposed to brackish water or seawater as an obligatory part of their reproductive migration. We investigated the changes in haemolymph osmolality, haemolymph Na+, Cl-, Ca2+and K+concentrations, as well as gill Na+/K+-ATPase (NKA) and carbonic anhydrase (CA) activity in both females and males, when in freshwater (0ppt,8mOsm (kg H2O)-1) and after step-wise acclimation to salinities ranging from7to35ppt (175-1044mOsm (kg H2O)-1). Both females and males strongly hyper-regulated their haemolymph osmolality over salinity range of0-21ppt, the isosmotic point approaching28ppt, and slightly hypo-regulated at35ppt. Haemolymph Na+and Cl-levels correlated positively with haemolymph osmolality. Ca2+and K+levels, however, were maintained at a relatively narrow range and showed a contrary trend in their response to salinity. This indicated that Na+and Cl-are the main contributors to increasing haemolymph osmolality. Reduced Na+/K+-ATPase and carbonic anhydrase activity in the posterior gills at elevated salinity suggest a direct physiological response to a decrease of Na+intake. No sexual differences existed for all parameters except for significantly higher haemolymph Ca2+and K+levels and CA activity in females than in males at some salinity levels. It is suggested that Eriocheir sinensis broodstock is a strong hyperosmoregulator but a weak hypo-osmoregulator. No sexual differences appeared to exist in osmotic and ionic regulation of Eriocheir sinensis broodstock exposed to different salinities.
4. Metabolic, immune and digestive response of Chinese mitten crab Eriocheir sinensis broodstock exposed to different salinities
To understand the metabolic, immune and digestive adjustments of Chinese mitten crab broodstock, Eriocheir sinensis, during their reproductive migration from freshwater to seawater, concentrations haemolymph metabolic variations (protein, glucose, triglycerides, cholesterol, urea, uric acid and creatinine) and activities of immune enzyme (alkaline phosphatase, superoxide dismutase and phenoloxidase) and digestive enzyme were determined after gradual acclimating of females and males from freshwater (0ppt) to different salinities (7,14,21,28,35ppt) medium. The results showed that increasing salinities caused a gradual decrease of hemolymph total protein and oxyhemocyanin contents of females and males. Some haemolymph metabolic variables reached a minimum at21ppt, and increased with both increase and decrease of the medium salinity (triglycerides, cholesterol, urea, creatinine in males and triglycerides, uric acid in females), or elevated either with the increase (urea in females) or decrease (uric acid in males) of medium salinities. Responses of haemolymph metabolic variations to salinity slightly differ between females and males. Hemolymph alkaline phosphatase activity increased slightly in7ppt; thereafter it dropped slowly until the end of experiment. Hemolymph phenoloxidase activity of females and males decreased gradually with the increase of environmental salinity. However, it maintained a stable level when the salinities were above21ppt in females. Hemolymph superoxide dismutase activity of males decreased with the increase of the medium salinity until21ppt and increased when the medium salinity above21ppt. All tested immune parameters except for SOD of females were significantly higher than those of males. Digestive enzyme activity except lipase of males were significantly higher than those of females at some salinities (P<0.05). Amylase, pepsin and trypsin activities of females reduced significantly when the salinity reached28ppt (P<0.05). Cellulase activity decreased when increasing the salinity to35(P<0.05). Remarkable decrease of males'digestive enzyme activities except lipase occurred at21and higher salinity, and an increase was shown in14ppt. The present results indicated that salinity of21ppt was the turning point of several biochemical parameters in E. sinensis broodstock. This salinity could be more'profitable'for E. sinensis broodstock from the energy-saving point of view. Immune and digestive enzyme activities of E. sinensis broodstock will be affected when the salinity was near to or above28ppt, which could exert a negative effect on reproduction. Metabolic and immune parameters in the haemolymph of females were slightly higher than those of males at some salinity levels, this suggestes higher metabolic level and greater immunity. The initial salinity that induced the decrease of digestive enzyme activity in males was lower than that in females; this result indicated that females were more tolerant elevated salinities than males in the digestive ability. This study might help understand some aspects of the metabolic, digestive and immune adjustments in euryhaline crabs related to osmoregulation.
5. Survival and development of Chinese mitten crab, Eriocheir sinensis embryo and Zoea in different salinities
Embryo and larval development of Chinese mitten crab. Eriocheir sinensis, is completed in the estuary, where they are often exposed to extreme salinity conditions. To test whether these salinity conditions have negative impacts on early development of embryo and zoea or not, we determined hatching rate, development time and survival rate when E. sinensis embryo for the gastrula, eyespot and pre-hatching stage were exposed to a salinity range of1-35ppt, and we also detected the survival rate of E. sinensis zoea I within192h later exposing them to different salinities. Reduced salinities (1and5ppt) accelerated embryonic development and increased hatching rate of eyespot and pre-hatching stage, but reduced salinities did not exert any effects on the gastrula stage development. Hatching rate significantly declined when the medium salinity reached at and above25ppt. No zoea I were hatched when eyespot stages embryos were exposed to30ppt and three stages embryos were exposed to35ppt. Higher survival rate of initial zoea I were presented when eyespot and pre-hatching stage embryos were exposed to salinity ranges of5-25ppt. However, survival rate of initial zoea I significantly declined when embryos were exposed to1and30ppt. All zoeas hardly survived at35ppt. Nearly40%zoeas I still survived until192h later during the range10-20ppt of salinity. However, all zoea I died24h later after exposure to salinity1and40ppt while died192h later at salinity range25-35and5ppt. The results showed that optimal salinity range was5-20ppt, when embryos began to be exposed to different salinities from gastrula stage and pre-hatching stage, while the salinity range became narrower (10-20ppt) for eyepot satge. The optimal salinity range was10-20ppt for zoea development. Out of above salinity ranges, will influence the development of embryo and larva, and then affect the wild population' reproduction.
6. Developmental changes of biochemical composition and digestive enzyme activity in the eggs of Chinese mitten crab, Eriocheir sinensis, under osmotic stress
Biochemical compositions (total carbohydrate, soluble protein, lipid and water content) contents, digestive enzymes (amylase, lipase, pepsin and trypsin) activity and embryo volume (egg diameter) have been determined in three developmental stages (gastrula, eyespot and pre-hatching stage) of E. sinensis eggs under hyper or hypo-osmotic stress. Increase of water contents (P<0.05), eggs diameter and pepsin and trypsin activities and decrease of soluble protein (P<0.05) and lipid contents and lipase activity (P<0.05) in eggs were noticed during the development of embryo. Total carbohydrate content and amylase activity increased at the eyespot stage (P<0.05), whereas they reduced during the prehatching stages (P<0.05). Hyper-osmotic (P<0.05) and hypo-osmotic stress reduced water content of eggs at each developmental stage. Although hyper-osmotic stress reduced the water content of eggs, significant effect presented only at eyespot stage (P<0.05). Soluble protein content of three developmental stages eggs decreased under hyper-osmotic stress while increased under hypo-osmotic stress (P<0.05). Total lipid content was unaffected by hyper-osmotic stress at any developmental stage, whereas it reduced significantly at pre-hatching stages under hypo-osmotic stress (P<0.05). Total carbohydrate contents did not show any significant variation after osmotic stress. Both hypo-osmotic and hyper-osmotic stress induced the decrease of egg diameters during the gastrula stage (P<0.05), however, no significant effect on eyespot stage. Egg diameter decreased before hatching, only under the hyper-osmotic stress (P<0.05). Among the four digestive enzymes assayed, amylase activities were the highest while lipase activities were the lowest. Hyper-osmotic stress reduced enzymes activities of the three developmental stage eggs, while hypo-osmotic stress enhanced the activities of digestive enzymes with the exception of reducing the amylase activities in eyespot stage (P<0.05) and trypsin activities in pre-hatching stage (P>0.05). Effect of salinity change on digestive enzymes is statically significant only during gastrula stage (P<0.05) except for lipase. It is suggested that salinity stress changed the activity of digestive enzymes which further affectes the utilizing of yolk substances, consequently, affects embryonic development. Dramatical variation of salinity in estuary ought to be having potential effect on embryonic development, especially induced by salt water encroachment.
7. Developmental changes of mRNA expression and activity of Na+/K+-ATPase, and expression of crustacean hyperglycemic hormone (CHH) gene in Chinese mitten crab Eriocheir sinensis eggs under osmotic stress
We examined ontogenetic and salinity-induced changes of Na+/K+-ATPase activity and mRNA expression of the a-subunit. We also detected the expression of crustacean hyperglycemic hormone (CHH) neuropeptides. which involved directly or indirectly in ionic and osmotic regulation, in the eyespot stage and pre-hatching stage. All activities and mRNA expression were determined24h later after directly exposeing embryos to1,15(control group), and35ppt. NKA activities were low in gastrula stage, and the enzyme activity increased6.53times in pre-hatching stage. Na+/K+-ATPase activity significantly decreased in gastrula and eyespot stages (P<0.05), and slightly reduced in pre-hatching stage at1ppt compared with15ppt, whereas the activity in the embryo of gastrula and pre-hatching stages was increased at salinity30ppt than at15and1ppt. However, increasing salinity reduced the enzyme activity (P<0.05) at eyespot stage, compared with the control group. The Na+/K+-ATPase a-subunit gene was expressed and elevated1.17times from gastrula stage to pre-hatching stage. There was no significant difference of Na+/K+-ATPase a-subunit gene expression between gastrula and eyespot stages, but expression increased remarkably before hatching (P<0.05). The expression of Na+/K+-ATPase a-subunit gene increased at30and1ppt compared with15ppt, but significant difference occurred only at eyepot stage under hyper salinity stress (P<0.05). Expression of CHH gene significantly increased with the development of embryo from eyespot stage to pre-hatching stage at all salinities (P<0.05), the expression remarkably elevated before hatching under hyper salinity stress while significantly decreased at eyespot stage under hypo salinity stress. It is suggested that although transcription and protein synthesis of Na+/K+-ATPase increased with the development of embryo, no consistent pattern between them under salinity stress. Neuroendocrine hormone CHH might be play important regulatory function in osmoregulation of embryo under osmotic stress.
8. Low salinity preference behavior of Chinese mitten crab Eriocheir sinensis megalopae
Up-estuary migration of crab larvae to adult habitats is thought to be started after molt to megalopae. However, the mechanisms used by crab megalopae for reinvasion of the estuary are unclear. We investigated the salinity preference behavior of E. sinensis megalopae in a seven choice-chambers device, to determine whether they showed preference behavior for the low salinity of the water or not. If low-salinity preference exists, this might be one of the bebavioral mechanisms for reinvasion estuary. Notable salinity gradients can be available in the seven choice-chambers device. Results revealed that most megalopa (60%-89%and69%-80%in two replicates, respectively) selected nominal0ppt as their final preference during9:00-12:00am. However, this behavioral weaken slightly (29%-60%and31%-57%megalopa prefer to nominal0ppt in two replicates, respectively) during14:00-17:00pm. Our findings reveal that E. sinensis megapolas obviously prefer low salinities, which associated with their strong osmoregulatory ability. Low-salinity preference behavior might be an important mechanism for megalopa's up-estuary migration. However, the extent of preference behavioral seems to be influenced by light. We speculated that the low-salinity preference behavior of megalopa is a hereditary behavioral while unaffected by environmental change. The importance of low-salinity preferences in relation to the megapola's up-estuary migration will be discussed in this study.
9. Effect of salinity on molt and Na+/K+-ATPase activity of E. sinensis megalopa
We determinted the percentage of megalopa molt to first juvenile instars under the condition of salinity range of1-40ppt, and detected the Na+/K+-ATPase activities of total megalopas after directly exposing them to1,15ppt (control group), and30ppt salinity. The result showed that nearly all the megalopas molted to the first juvenile instars at salinities range of1-40ppt on the fifth day of experiment, whereas molt delayed significantly at40ppt, compared with other salinities. Na+/K+-ATPase activity increased remarkably either at low salinity (1ppt) or at high salinity (30ppt) medium, compared with control group. The results suggest that Eriocheir sinensis megalopas possess moderate hyper-/hypo-regulating ability, and the osmoregulatory ability was also exhibited by the change of Na+/K+-ATPase activity under salinity stress. The results of this study provide physical evidence for comprehensing the up-estuary migration of crab larvae to adult habitats.
引文
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