海月水母（Aurelia sp.1）生活史关键过程研究
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摘要
近年来，全球海洋中水母的数量呈增加趋势，许多海域出现水母大规模暴发现象，并且频率和范围逐年扩大。我国所在的东亚海域是世界上大型水母暴发的“重灾区”，许多海域已经达到泛滥的程度。水母暴发对海洋渔业、旅游业、沿岸工业和人身安全等造成很大的威胁。
由于海月水母(Aurelia spp.)多分布在人类活动较集中的海湾、河口，因此对人类造成的影响较大，更容易造成危害。研究发现海月水母(Aurelia spp.)生活史策略复杂生态类型多样，而且存在亚种之间的区别。本文通过室内培养实验和野外调查工作，研究了中国近海水母暴发种海月水母(Aurelia sp.1)生活史关键过程及在胶州湾的生活史特征。分析特定阈值的环境因子对生活史过程及生殖策略的影响，以期揭示海月水母在中国近海大量发生的关键过程及暴发机制。同时可以将海月水母作为研究中国近海水母的模式生物，为开展其他钵水母的研究提供参考。
附着选择性实验发现，浮浪幼虫对人工材料(波纹板、网衣、PV管)具有偏好性，附着密度较大；水平放置的附着材料底部附着更多的水螅体。水螅体可以通过无性繁殖方式在新的栖息地形成水螅体种群；进行种群扩张过程中存在密度限制，较低密度时水螅体种群增长较快，高密度时水螅体种群增长较慢。日益增加的海洋废弃物(塑料、玻璃、木材)污染，以及海底构造物(海水养殖筏、海洋工程)为水螅体提供了更多的附着表面，利于栖息地的扩张，形成更大的水螅体种群。
环境因子影响实验结果表明，温度对横裂过程、碟状体产生影响显著；饵料对水螅体数量影响显著。水螅体从20℃降温至15℃和10℃可发生横裂产生碟状体，适宜横裂的温度范围内，相对高温和充足饵料有利于释放更多碟状体；适宜横裂的温度范围外，相对高温和充足饵料有利于水螅体数量的增加。适宜横裂的温度范围持续时间达到水螅体的响应时间阀值，才能够完成从底栖息阶段的水螅体到浮游阶段水母体的转化。在胶州湾温度周年变化范围内，经过低温处理的水螅体（5℃，2个月），升温后可发生横裂释放碟状体。在胶州湾，温度条件是控制水螅体横裂释放碟状体，从而形成海月水母暴发的关键因子。春季升温速率较慢时即适宜横裂温度范围持续时间较长，则延长横裂过程积累更多的碟状体，如果饵料充足则释放更多的碟状体，为形成海月水母暴发提供基础。不同温度下，水螅体对光照强度的响应不同。光照强度对水螅体数量，横裂过程及碟状体数量影响不显著。
温度和饵料种类对水螅体横裂与否有决定性作用。有饵料的环境下，将水螅体从20℃降温到15℃和10℃能够发生横裂释放碟状体，无饵料的环境下不能发生横裂。以卤虫为饵料的实验组，产生水螅体和碟状体的数量最大。环境中动物性饵料不足时，低温条件下浮游植物能够为水螅体提供饵料来源；高温条件下，高密度的浮游植物不利于水螅体存活。充足的动物性饵料和适宜的温度条件是水螅体发生横裂大量释放碟状体的基础条件。
对胶州湾2007-2011年夜光虫丰度的年际变化和海月水母丰度变化研究发现，发生海月水母暴发的2009年和2011年，海月水母出现的4~7月份夜光虫平均丰度分别为1112.3和712.5ind m~(-3)，非水母暴发年份4~7月份夜光虫平均丰度在2381.1到17565.4ind m~(-3)范围内。海月水母丰度大的年份，夜光虫丰度小。室内培养实验发现海月水母碟状体和水母体均能够摄食夜光虫。摄食率随伞径线性增加（0.5~8cm; R~2=0.3663, p<0.01）;随夜光虫细胞密度线性增加(10-2000cellL~(-1); R~2=0.955, p<0.01)。当夜光虫细胞密度为10ind/L时,碟状体对夜光虫的清除率为0.02h~(-1)；8cm的海月水母对夜光虫的清除率为0.36h~(-1)。因此，我们认为海月水母对夜光虫丰度存在潜在调控作用。
在胶州湾，每年5~6月大量出现海月水母(Aurelia sp.1)碟状体，碟状体大量出现时的温度范围在12~18℃，温度和海月水母碟状体出现时间关系密切。海月水母水母体七月份伞径达到最大值，八月份以后丰度及伞径均减小。碟状体在5℃和10℃下可以维持较高的存活率但是不能发育为水母体，在15和20℃时，可以迅速发育为水母体。结合野外调查和室内实验结果，初步构建了海月水母在胶州湾生活史模型。水螅体在胶州湾全年存在，可以通过出芽和足囊等方式进行无性繁殖，春季温度升高时水螅体发生横裂释放碟状体，五、六月温度上升到15℃以上时碟状体生长迅速发育成水母体，七月份以后水母体达到性成熟进行有性繁殖释放浮浪幼虫，九月末海月水母消亡。胶州湾码头、跨海大桥、养殖阀架等为水螅体附着提供了良好的基质，富营养化的高生产力环境提供了充足的饵料，为水母暴发提供了基础。
Jellyfish blooms occurred all over the world in recent years, especially the giantjellyfish blooms in the Far East seas including China coastal waters. Jellyfish bloomsdamage the structure and function of the marine ecosystem, causing many economicand social problems to the tourism because of jellyfish sting, to the coastal industriesby blocking the pipeline of its cooling system and to the fishery resources and fisheryactivities.
The jellyfish Aurelia spp. appears in estuaries and bays, causing more damage tohuman because of higher frequency of human activities. Life history of Aurelia spp. iscomplex with various ecological forms, recent research showed there are manysubspecies. In this research, we studied the key process of life cycle and the life cyclecharacter of Aurelia sp.1which blooms in coastal of China. Analyzed the effect ofenvironmental factors on the life cycle and reproduction strategy, intend to reveal thekey process and mechanism of the blooms of Aurelia sp.1in coastal of China. At thesame time, Aurelia sp.1can serve as a model organism of large jellyfish, providingconsult to the research of other scyphomedusa.
In this study, we examined planula of Aurelia sp.1settlement on living-oyster,oyster shell, scallop shell, floor plates, netting and PV-pipe, proliferation of polyps onthe surfaces of bamboo and cement block. The planula and polyps preferredman-made materials (floor plates, netting, and PV-pipe). The underside of floor plateswas strongly favorable for settlement. Polyps can form new polyps population through asexual reproduction on new habitat, the expanding process was limited bypolyps population density. When the density was relatively low, polyps populationincreased rapidly, and this increase was slower when density was relatively high.Increasing litter (plastics, glass, and wood) pollution and submarine constructions(mariculture and ocean engineering) provide more attachment surface and thusenlarge the areas of polyps distribution and form larger number of polyps populations.
Temperature and food play a vital role in determining asexual direction of jellyfish in its polyp stage. The combinations of four temperature grades (10,15,20and25℃) and three food levels (no food, feed once a week and feed everyday) were setup to exam Aurelia sp.1life cycle strategy. Temperature could determine thestrobilation and ephyrae release. Food determined the number of polyps. Temperaturechanging from20℃to15℃and10℃induced ephyrae release. In suitabletemperature range for strobilation, relatively higher temperature and sufficient foodenabled polyps to produce more ephyrae. At temperatures beyond range forstrobilation, sufficient food condition enabled polyp produce polyps. Highertemperature and lower food level induced higher polyp mortality. In the range ofannual temperature of Jiaozhou Bay, polyps go through low temperature process(5℃，2month) can strobilation and release ephyrae when temperature increased.Response of polyps to light density are different under different temperature, theeffect of light density on polyps number, strobilation process and number of ephyraeare not significant.
Environmental factors such as temperature and food type affect the rate ofasexual reproduction of jellyfish at the polyp stage. Combinations of threetemperatures (10,15, and20°C) and four food treatments (Prorocentrum donghaiense,Skeletonema costatum, Artemia sp, nauplii, and no food) were established to examinethe asexual reproduction strategy of Aurelia sp.1. Temperature and food typedetermined the strobilation of polyps. A change from20to15°C or10°C, induced polyps to release ephyrae when food was present, but without food, polyps did notstrobilate. Groups with Artemia sp. nauplii as prey produced more polyps and ephyraethrough buds, podocysts, and strobilation. Animal prey was better than plants as food.At20°C, the mortality rate of polyps exceeded50%in plant fed and unfed polyps.The number of polyps increased rapidly with Artemia nauplii as prey. We concludethat, when animal prey is limited, plants can serve as a nutrient source and satisfy theenergy requirements for polyps at lower temperatures. High densities ofphytoplankton can be lethal to polyps at high temperatures. Abundant animal prey andsuitable temperatures are essential conditions for polyps to strobilate and releaseephyrae, leading to jellyfish blooms.
Both Aurelia sp.1and Noctiluca scintillans are unwanted species that can bloom.Though low N.scintillans abundance had been recorded when Aurelia sp.1massivelyoccurred, no robust evidence exists on their interaction. In this study, inter-annualvariation of N.scintillans abundance was analyzed with field sampling data in theJiaozhouBay, and predation rate of Aurelia sp.1on N.scintillans was estimated withlaboratory incubation. During the existence period of Aurelia sp.1(April-July),average N.scintillans abundance was1112.3and712.5ind m~(-3)in bloom years of2009and2011, and varied between2381.1and17565.4ind m~(-3)in non-bloom years. Inlaboratory, Aurelia sp.1in both ephyrae and medusa stages ingested N.scintillans insignificant amount. The feeding rate linearly increased with both the bell diameter(0.5-8cm; R~2=0.3663, p<0.01) and prey concentrations (10-2000cell L~(-1); R~2=0.955,p<0.01). At the lowest concentration of10, the removal rate varied from0.02h~(-1)inephyra to0.36h~(-1)in8cm sized medusa. It was suggested that massively occurredAurelia sp.1can potentially depress N.scintillans abundance in natural environments.
In Jiaozhou Bay, ephyrae of Aurelia sp.1appeared from May to June, whentemperature was in the range of12to18℃in spring in Jiaozhou Bay. Survival rate ofephyrae is high in low temperature but can’t develop to medusa, growth rate of ephyrae is high when temperature beyond15℃. Polyps of Aurelia sp.1survivalduring the whole year, and can go through asexual reproduction through buds andpodocyst, ephyrae released when temperature increasing in spring, planula wereproduced through sexual reproduction of adult medusa of Aurelia sp.1during July andAugust, bell diameter of Aurelia sp.1decreased and medusa of Aurelia sp.1die out atthe end of September.
由于海月水母(Aurelia spp.)多分布在人类活动较集中的海湾、河口，因此对人类造成的影响较大，更容易造成危害。研究发现海月水母(Aurelia spp.)生活史策略复杂生态类型多样，而且存在亚种之间的区别。本文通过室内培养实验和野外调查工作，研究了中国近海水母暴发种海月水母(Aurelia sp.1)生活史关键过程及在胶州湾的生活史特征。分析特定阈值的环境因子对生活史过程及生殖策略的影响，以期揭示海月水母在中国近海大量发生的关键过程及暴发机制。同时可以将海月水母作为研究中国近海水母的模式生物，为开展其他钵水母的研究提供参考。
附着选择性实验发现，浮浪幼虫对人工材料(波纹板、网衣、PV管)具有偏好性，附着密度较大；水平放置的附着材料底部附着更多的水螅体。水螅体可以通过无性繁殖方式在新的栖息地形成水螅体种群；进行种群扩张过程中存在密度限制，较低密度时水螅体种群增长较快，高密度时水螅体种群增长较慢。日益增加的海洋废弃物(塑料、玻璃、木材)污染，以及海底构造物(海水养殖筏、海洋工程)为水螅体提供了更多的附着表面，利于栖息地的扩张，形成更大的水螅体种群。
环境因子影响实验结果表明，温度对横裂过程、碟状体产生影响显著；饵料对水螅体数量影响显著。水螅体从20℃降温至15℃和10℃可发生横裂产生碟状体，适宜横裂的温度范围内，相对高温和充足饵料有利于释放更多碟状体；适宜横裂的温度范围外，相对高温和充足饵料有利于水螅体数量的增加。适宜横裂的温度范围持续时间达到水螅体的响应时间阀值，才能够完成从底栖息阶段的水螅体到浮游阶段水母体的转化。在胶州湾温度周年变化范围内，经过低温处理的水螅体（5℃，2个月），升温后可发生横裂释放碟状体。在胶州湾，温度条件是控制水螅体横裂释放碟状体，从而形成海月水母暴发的关键因子。春季升温速率较慢时即适宜横裂温度范围持续时间较长，则延长横裂过程积累更多的碟状体，如果饵料充足则释放更多的碟状体，为形成海月水母暴发提供基础。不同温度下，水螅体对光照强度的响应不同。光照强度对水螅体数量，横裂过程及碟状体数量影响不显著。
温度和饵料种类对水螅体横裂与否有决定性作用。有饵料的环境下，将水螅体从20℃降温到15℃和10℃能够发生横裂释放碟状体，无饵料的环境下不能发生横裂。以卤虫为饵料的实验组，产生水螅体和碟状体的数量最大。环境中动物性饵料不足时，低温条件下浮游植物能够为水螅体提供饵料来源；高温条件下，高密度的浮游植物不利于水螅体存活。充足的动物性饵料和适宜的温度条件是水螅体发生横裂大量释放碟状体的基础条件。
对胶州湾2007-2011年夜光虫丰度的年际变化和海月水母丰度变化研究发现，发生海月水母暴发的2009年和2011年，海月水母出现的4~7月份夜光虫平均丰度分别为1112.3和712.5ind m~(-3)，非水母暴发年份4~7月份夜光虫平均丰度在2381.1到17565.4ind m~(-3)范围内。海月水母丰度大的年份，夜光虫丰度小。室内培养实验发现海月水母碟状体和水母体均能够摄食夜光虫。摄食率随伞径线性增加（0.5~8cm; R~2=0.3663, p<0.01）;随夜光虫细胞密度线性增加(10-2000cellL~(-1); R~2=0.955, p<0.01)。当夜光虫细胞密度为10ind/L时,碟状体对夜光虫的清除率为0.02h~(-1)；8cm的海月水母对夜光虫的清除率为0.36h~(-1)。因此，我们认为海月水母对夜光虫丰度存在潜在调控作用。
在胶州湾，每年5~6月大量出现海月水母(Aurelia sp.1)碟状体，碟状体大量出现时的温度范围在12~18℃，温度和海月水母碟状体出现时间关系密切。海月水母水母体七月份伞径达到最大值，八月份以后丰度及伞径均减小。碟状体在5℃和10℃下可以维持较高的存活率但是不能发育为水母体，在15和20℃时，可以迅速发育为水母体。结合野外调查和室内实验结果，初步构建了海月水母在胶州湾生活史模型。水螅体在胶州湾全年存在，可以通过出芽和足囊等方式进行无性繁殖，春季温度升高时水螅体发生横裂释放碟状体，五、六月温度上升到15℃以上时碟状体生长迅速发育成水母体，七月份以后水母体达到性成熟进行有性繁殖释放浮浪幼虫，九月末海月水母消亡。胶州湾码头、跨海大桥、养殖阀架等为水螅体附着提供了良好的基质，富营养化的高生产力环境提供了充足的饵料，为水母暴发提供了基础。
Jellyfish blooms occurred all over the world in recent years, especially the giantjellyfish blooms in the Far East seas including China coastal waters. Jellyfish bloomsdamage the structure and function of the marine ecosystem, causing many economicand social problems to the tourism because of jellyfish sting, to the coastal industriesby blocking the pipeline of its cooling system and to the fishery resources and fisheryactivities.
The jellyfish Aurelia spp. appears in estuaries and bays, causing more damage tohuman because of higher frequency of human activities. Life history of Aurelia spp. iscomplex with various ecological forms, recent research showed there are manysubspecies. In this research, we studied the key process of life cycle and the life cyclecharacter of Aurelia sp.1which blooms in coastal of China. Analyzed the effect ofenvironmental factors on the life cycle and reproduction strategy, intend to reveal thekey process and mechanism of the blooms of Aurelia sp.1in coastal of China. At thesame time, Aurelia sp.1can serve as a model organism of large jellyfish, providingconsult to the research of other scyphomedusa.
In this study, we examined planula of Aurelia sp.1settlement on living-oyster,oyster shell, scallop shell, floor plates, netting and PV-pipe, proliferation of polyps onthe surfaces of bamboo and cement block. The planula and polyps preferredman-made materials (floor plates, netting, and PV-pipe). The underside of floor plateswas strongly favorable for settlement. Polyps can form new polyps population through asexual reproduction on new habitat, the expanding process was limited bypolyps population density. When the density was relatively low, polyps populationincreased rapidly, and this increase was slower when density was relatively high.Increasing litter (plastics, glass, and wood) pollution and submarine constructions(mariculture and ocean engineering) provide more attachment surface and thusenlarge the areas of polyps distribution and form larger number of polyps populations.
Temperature and food play a vital role in determining asexual direction of jellyfish in its polyp stage. The combinations of four temperature grades (10,15,20and25℃) and three food levels (no food, feed once a week and feed everyday) were setup to exam Aurelia sp.1life cycle strategy. Temperature could determine thestrobilation and ephyrae release. Food determined the number of polyps. Temperaturechanging from20℃to15℃and10℃induced ephyrae release. In suitabletemperature range for strobilation, relatively higher temperature and sufficient foodenabled polyps to produce more ephyrae. At temperatures beyond range forstrobilation, sufficient food condition enabled polyp produce polyps. Highertemperature and lower food level induced higher polyp mortality. In the range ofannual temperature of Jiaozhou Bay, polyps go through low temperature process(5℃，2month) can strobilation and release ephyrae when temperature increased.Response of polyps to light density are different under different temperature, theeffect of light density on polyps number, strobilation process and number of ephyraeare not significant.
Environmental factors such as temperature and food type affect the rate ofasexual reproduction of jellyfish at the polyp stage. Combinations of threetemperatures (10,15, and20°C) and four food treatments (Prorocentrum donghaiense,Skeletonema costatum, Artemia sp, nauplii, and no food) were established to examinethe asexual reproduction strategy of Aurelia sp.1. Temperature and food typedetermined the strobilation of polyps. A change from20to15°C or10°C, induced polyps to release ephyrae when food was present, but without food, polyps did notstrobilate. Groups with Artemia sp. nauplii as prey produced more polyps and ephyraethrough buds, podocysts, and strobilation. Animal prey was better than plants as food.At20°C, the mortality rate of polyps exceeded50%in plant fed and unfed polyps.The number of polyps increased rapidly with Artemia nauplii as prey. We concludethat, when animal prey is limited, plants can serve as a nutrient source and satisfy theenergy requirements for polyps at lower temperatures. High densities ofphytoplankton can be lethal to polyps at high temperatures. Abundant animal prey andsuitable temperatures are essential conditions for polyps to strobilate and releaseephyrae, leading to jellyfish blooms.
Both Aurelia sp.1and Noctiluca scintillans are unwanted species that can bloom.Though low N.scintillans abundance had been recorded when Aurelia sp.1massivelyoccurred, no robust evidence exists on their interaction. In this study, inter-annualvariation of N.scintillans abundance was analyzed with field sampling data in theJiaozhouBay, and predation rate of Aurelia sp.1on N.scintillans was estimated withlaboratory incubation. During the existence period of Aurelia sp.1(April-July),average N.scintillans abundance was1112.3and712.5ind m~(-3)in bloom years of2009and2011, and varied between2381.1and17565.4ind m~(-3)in non-bloom years. Inlaboratory, Aurelia sp.1in both ephyrae and medusa stages ingested N.scintillans insignificant amount. The feeding rate linearly increased with both the bell diameter(0.5-8cm; R~2=0.3663, p<0.01) and prey concentrations (10-2000cell L~(-1); R~2=0.955,p<0.01). At the lowest concentration of10, the removal rate varied from0.02h~(-1)inephyra to0.36h~(-1)in8cm sized medusa. It was suggested that massively occurredAurelia sp.1can potentially depress N.scintillans abundance in natural environments.
In Jiaozhou Bay, ephyrae of Aurelia sp.1appeared from May to June, whentemperature was in the range of12to18℃in spring in Jiaozhou Bay. Survival rate ofephyrae is high in low temperature but can’t develop to medusa, growth rate of ephyrae is high when temperature beyond15℃. Polyps of Aurelia sp.1survivalduring the whole year, and can go through asexual reproduction through buds andpodocyst, ephyrae released when temperature increasing in spring, planula wereproduced through sexual reproduction of adult medusa of Aurelia sp.1during July andAugust, bell diameter of Aurelia sp.1decreased and medusa of Aurelia sp.1die out atthe end of September.
引文
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