三种海水经济鱼类早期发育生物学的研究
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摘要
海水经济鱼类的养殖在我国已经形成第四次海水养殖浪潮,经济效益显著,有力地推动了我国海水养殖的产业结构调整和可持续发展。然而在海水养殖发展过程中也存在着诸多问题,尤其是早期发育阶段的高死亡率,严重制约了我国海水养殖产业的稳定和健康发展。
海水鱼类养殖的关键为高质量,高存活率苗种的生产和培育,由于鱼类种类繁多,生物多样性丰富,对应实际的繁育技术,尤其是新品种的开发,必须要做出相应的调整。这就要求我们必须对每一种鱼类早期发育有所了解,并将形态和组织上的数据用于指导生产。
本文通过显微观察和组织学研究,主要描述和研究了我国北方三种重要的海水经济鱼类(条斑星鲽、杂交鲆、条石鲷)的早期发育生物学,并结合实际生产进一步阐明关键期的产生原因,机理以及采用相应的对策。具体结果如下:
1.条斑星鲽:作为冷温性鲆鲽鱼类,条斑星鲽早期发育过程的特征主要有:
①条斑星鲽受精卵无油球,卵子呈半浮性;不同步卵裂现象提前,发生在第三次卵裂;卵裂期裂球大小差异大。孵化过程较长,在水温8±0.3℃,盐度33的条件下,经9 d孵化。条斑星鲽胚胎发育的不同时期对温度的敏感性不同,其中原肠期对温度比较敏感。
②在8-10℃,盐度33的条件下,8-9 dph开口摄食。且开口时,其吻前端出现有一点状黑褐色素,构成了条斑星鲽仔鱼“开口期”的重要标志。卵黄囊于消失。在后期仔鱼末期,背鳍和臀鳍上形成特有的黑褐色条斑带。
③杯状细胞首先出现在咽腔后部和食道前段,胃腺和幽门盲囊出现于29 dph,变态期始于30dph。在条斑星鲽早期发育过程中,观察到其直肠粘膜层细胞质出现大量嗜伊红颗粒,为仔鱼肠道上皮吸收的蛋白质。
④首先淋巴化的免疫器官是头肾,然后是胸腺和脾脏,这与大部分硬骨鱼类不同。条斑星鲽除头肾和脾脏外,胸腺实质也形成MMCs。其中以脾脏形成MMCs最为丰富,形态多样。
2.杂交鲆:为同属的牙鲆和夏鲆间的远缘杂交种,其发育过程的特点为:
①在温度为15.4～16.0℃,杂交鲆胚胎从受精到孵化所需的时间为76 h左右,胚孔关闭前期,胚胎先出现视囊及克氏囊,而后形成体节。孵出前胚体在卵膜内环绕不到1周。
②孵化后消失。杂交鲆群体变态间隔长(34-60 dph),且变态高峰期出现的冠状幼鳍不明显(与母本牙鲆相比),数量为7-8根。
③组织学观察发现,其消化系统中胃腺出现较晚,且胃腺发育过程缓慢(与母本牙鲆相比)。甲状腺滤泡增生不明显,颜色较浅,数量较少。杂交鲆在早期发育过程中,并没有出现鳔原基。
3.条石鲷作为岩礁性的暖水性鱼类,早期发育过程也较为特殊,包括外形以及内部的器官结构。主要特点有:
①受精卵:受精卵卵黄上具有龟裂结构,为鱼卵的分类特征之一。
②初孵仔鱼:初孵仔鱼背鳍膜上的黑色素,从体背面向背鳍膜边缘移动,到3dph仔鱼基本消失,此为本种仔鱼发育所特有的特点。
③后期仔鱼和稚鱼:肠道肌肉层加厚明显,仔稚鱼胃肠排空率急剧上升,死亡率增加,通过改善常规的投饵方式部分解决了这个死亡高峰的问题。在幼鱼初期,牙齿融合为骨喙,为石鲷科鱼类的特征。
④胸腺上皮分泌细胞:类似的现象同样在虹鳟鱼中发现,但是虹鳟鱼胸腺上皮分泌细胞不如条石鲷的丰富,同样也不如条石鲷的排列整齐,而是零星分布在胸腺上皮与咽腔接触的表面。除了正常的造血器官—脾脏和头肾外,肝脏、胰腺和鳔等多种组织等也出现MMCs,此现象在硬骨鱼类不多见,一般发生在软骨鱼类。
Marine economical finfish aquaculture formed the fourth wave of Marine Aquaculture in China,acquired obvious economic value and social value,push Chinese marine aquaculture.However,during the development of marine aquaculture,there have some unresolved problems,especially the critical periods during the course of larval culture,still a obstacle hinder the stable and healthy development our Chinese marine aquaculture industries.
The key element of the marine finfish aquaculture is fingerling production with good quality and high survival rate.However,due to the abundant biodiversity of fish,we must modify our protocol in order to supply relevant species with the most suitable technology. At last,the data obtained from morphological and histological observation must be used in direction of the practical works.
The early life history of three important economical marine finfish in northern China (hybrid flounder,barfin flounder and Japanese parrotfish) were described the microscope observation and histological study,and further explained the emerged reasons,inner mechanism and relevant protocol of critical periods during larval early ontogeny.The concrete results are as below:
1.Barfin flounder:as a cold water flatfish,the early developmental characteristics is:
①The eggs were devoid of oil globule,and semi-pelagic feature;the asynchronous cleavage advanced,occurred on the third cleavage stage.The size heterogeneity between blastomeres was obvious.It had a long incubation time,under the temperature of 8±0.3℃,and salinity of 33,it took 9 d to hatch a larvae from the fertilized eggs.The different embryonic stage of barfin flounder had varied resistance to temperature and gastrula stage was sensitive to temperature.
②It also had a long period from hatching to feeding,under the temperature of 8-10 ℃,and salinity of 33,the larvae began exogenous feeding on circa 8-9 dph.A dot melanin on the beak was the main feature when the larvae fed.The yolk-sac disappeared on 14 dph.The special black-brown striped bars were seen at the posterior period of post-larvae stage.
③The goblet cells occurred on the posterior region of pharynx cavity and the anterior region of esophagus,and the gastric glands differentiated on 29 dph, metamorphosis began at 30 dph.During the early life stage,abundant acidophilic granules on the rectal epithelium indicated the protein-absorbing ability of the larvae intestine.
④The first lymphoid organ was head kidney,then thymus and spleen,which was different to the most marine teleosts.Besides head kidney and spleen,thymus also formed the MMCs.The MMCs in spleen was most abundant.
2.Hybrid flounder:as a hybrid of Japanese flounder and summer flounder-the same family,its early developmental characteristics is:
①Under the temperature of 15.4～16.0℃,the embryos of hybrid flounder took 76 h to hatch,before the blastopore closure stage,the embryo first formed the optic vesicle and kuffer's vesicle,then forming somites.Before hatching,the embryoic tail and nose never touched in egg membrane.
②The larvae began to feeding on 4 dph,and the yolk-sac & oil globule disappeared on 5 dph & 10 dph,respectively.The intervals of metamorphosis in hybrid flounder is longer(34-60 dph),and the emergence of the crown-like larval fins was not evident like its maternal flounder,however,the numbers of larval fins were 7-8.
③Through the histological observation,the gastric glands of digestive system appeared later,and the developmental duration of the gastric glands was also slower comparison with its maternal flounder.The increment of the thyroid follicles was not evident,the pale color and fewer numbers.During the early ontogeny of the hybrid flounder,no evident indicated the differentiation of anlage of swim bladder.
3.Japanese parrotfish:as a warm water species,lithophilic teleosts,the early developmental feature is:
①fertilized eggs:The yolk was segmented,and acts as a feature of fish egg classification.
②The newly-hatched larve:The melanin on the larval dorsal finfold in the newly-hatched larvae,moved from to the edge of the larval dorsal finfold,and disappeared on 3 dph.This phenomenon indicated the special ontogenetic characteristics of this species.
③post-larvae:The thickness of muscular tissue in intestine increased quickly during this period,and the gut evacuation rised abruptly,at the same time,the mortality rate rise up,and we resolved this problem by the means of changing the routine feeding pattern.During the juvenile stage,the jaw teeth of Japanese parrotfish fused into two whole teeth plates,which was a normal feature of Oplegnathidae.
④some secreting cells were found in the thymus during the larval development, which usually were found in some lower finfish,like salmonids.Through our studies,the numbers of secreting cells in rainbow trout was fewer than Japanese parrotfish,and its arrangement was also devoid of orderliness.MMCs was also normally found in the hemopoietic organs-spleen and head kidney,also found in the parenchyma of liver, pancreas and swim bladder,which usually occurred in elasmobranch fishes and few in teleosts.
海水鱼类养殖的关键为高质量,高存活率苗种的生产和培育,由于鱼类种类繁多,生物多样性丰富,对应实际的繁育技术,尤其是新品种的开发,必须要做出相应的调整。这就要求我们必须对每一种鱼类早期发育有所了解,并将形态和组织上的数据用于指导生产。
本文通过显微观察和组织学研究,主要描述和研究了我国北方三种重要的海水经济鱼类(条斑星鲽、杂交鲆、条石鲷)的早期发育生物学,并结合实际生产进一步阐明关键期的产生原因,机理以及采用相应的对策。具体结果如下:
1.条斑星鲽:作为冷温性鲆鲽鱼类,条斑星鲽早期发育过程的特征主要有:
①条斑星鲽受精卵无油球,卵子呈半浮性;不同步卵裂现象提前,发生在第三次卵裂;卵裂期裂球大小差异大。孵化过程较长,在水温8±0.3℃,盐度33的条件下,经9 d孵化。条斑星鲽胚胎发育的不同时期对温度的敏感性不同,其中原肠期对温度比较敏感。
②在8-10℃,盐度33的条件下,8-9 dph开口摄食。且开口时,其吻前端出现有一点状黑褐色素,构成了条斑星鲽仔鱼“开口期”的重要标志。卵黄囊于消失。在后期仔鱼末期,背鳍和臀鳍上形成特有的黑褐色条斑带。
③杯状细胞首先出现在咽腔后部和食道前段,胃腺和幽门盲囊出现于29 dph,变态期始于30dph。在条斑星鲽早期发育过程中,观察到其直肠粘膜层细胞质出现大量嗜伊红颗粒,为仔鱼肠道上皮吸收的蛋白质。
④首先淋巴化的免疫器官是头肾,然后是胸腺和脾脏,这与大部分硬骨鱼类不同。条斑星鲽除头肾和脾脏外,胸腺实质也形成MMCs。其中以脾脏形成MMCs最为丰富,形态多样。
2.杂交鲆:为同属的牙鲆和夏鲆间的远缘杂交种,其发育过程的特点为:
①在温度为15.4～16.0℃,杂交鲆胚胎从受精到孵化所需的时间为76 h左右,胚孔关闭前期,胚胎先出现视囊及克氏囊,而后形成体节。孵出前胚体在卵膜内环绕不到1周。
②孵化后消失。杂交鲆群体变态间隔长(34-60 dph),且变态高峰期出现的冠状幼鳍不明显(与母本牙鲆相比),数量为7-8根。
③组织学观察发现,其消化系统中胃腺出现较晚,且胃腺发育过程缓慢(与母本牙鲆相比)。甲状腺滤泡增生不明显,颜色较浅,数量较少。杂交鲆在早期发育过程中,并没有出现鳔原基。
3.条石鲷作为岩礁性的暖水性鱼类,早期发育过程也较为特殊,包括外形以及内部的器官结构。主要特点有:
①受精卵:受精卵卵黄上具有龟裂结构,为鱼卵的分类特征之一。
②初孵仔鱼:初孵仔鱼背鳍膜上的黑色素,从体背面向背鳍膜边缘移动,到3dph仔鱼基本消失,此为本种仔鱼发育所特有的特点。
③后期仔鱼和稚鱼:肠道肌肉层加厚明显,仔稚鱼胃肠排空率急剧上升,死亡率增加,通过改善常规的投饵方式部分解决了这个死亡高峰的问题。在幼鱼初期,牙齿融合为骨喙,为石鲷科鱼类的特征。
④胸腺上皮分泌细胞:类似的现象同样在虹鳟鱼中发现,但是虹鳟鱼胸腺上皮分泌细胞不如条石鲷的丰富,同样也不如条石鲷的排列整齐,而是零星分布在胸腺上皮与咽腔接触的表面。除了正常的造血器官—脾脏和头肾外,肝脏、胰腺和鳔等多种组织等也出现MMCs,此现象在硬骨鱼类不多见,一般发生在软骨鱼类。
Marine economical finfish aquaculture formed the fourth wave of Marine Aquaculture in China,acquired obvious economic value and social value,push Chinese marine aquaculture.However,during the development of marine aquaculture,there have some unresolved problems,especially the critical periods during the course of larval culture,still a obstacle hinder the stable and healthy development our Chinese marine aquaculture industries.
The key element of the marine finfish aquaculture is fingerling production with good quality and high survival rate.However,due to the abundant biodiversity of fish,we must modify our protocol in order to supply relevant species with the most suitable technology. At last,the data obtained from morphological and histological observation must be used in direction of the practical works.
The early life history of three important economical marine finfish in northern China (hybrid flounder,barfin flounder and Japanese parrotfish) were described the microscope observation and histological study,and further explained the emerged reasons,inner mechanism and relevant protocol of critical periods during larval early ontogeny.The concrete results are as below:
1.Barfin flounder:as a cold water flatfish,the early developmental characteristics is:
①The eggs were devoid of oil globule,and semi-pelagic feature;the asynchronous cleavage advanced,occurred on the third cleavage stage.The size heterogeneity between blastomeres was obvious.It had a long incubation time,under the temperature of 8±0.3℃,and salinity of 33,it took 9 d to hatch a larvae from the fertilized eggs.The different embryonic stage of barfin flounder had varied resistance to temperature and gastrula stage was sensitive to temperature.
②It also had a long period from hatching to feeding,under the temperature of 8-10 ℃,and salinity of 33,the larvae began exogenous feeding on circa 8-9 dph.A dot melanin on the beak was the main feature when the larvae fed.The yolk-sac disappeared on 14 dph.The special black-brown striped bars were seen at the posterior period of post-larvae stage.
③The goblet cells occurred on the posterior region of pharynx cavity and the anterior region of esophagus,and the gastric glands differentiated on 29 dph, metamorphosis began at 30 dph.During the early life stage,abundant acidophilic granules on the rectal epithelium indicated the protein-absorbing ability of the larvae intestine.
④The first lymphoid organ was head kidney,then thymus and spleen,which was different to the most marine teleosts.Besides head kidney and spleen,thymus also formed the MMCs.The MMCs in spleen was most abundant.
2.Hybrid flounder:as a hybrid of Japanese flounder and summer flounder-the same family,its early developmental characteristics is:
①Under the temperature of 15.4～16.0℃,the embryos of hybrid flounder took 76 h to hatch,before the blastopore closure stage,the embryo first formed the optic vesicle and kuffer's vesicle,then forming somites.Before hatching,the embryoic tail and nose never touched in egg membrane.
②The larvae began to feeding on 4 dph,and the yolk-sac & oil globule disappeared on 5 dph & 10 dph,respectively.The intervals of metamorphosis in hybrid flounder is longer(34-60 dph),and the emergence of the crown-like larval fins was not evident like its maternal flounder,however,the numbers of larval fins were 7-8.
③Through the histological observation,the gastric glands of digestive system appeared later,and the developmental duration of the gastric glands was also slower comparison with its maternal flounder.The increment of the thyroid follicles was not evident,the pale color and fewer numbers.During the early ontogeny of the hybrid flounder,no evident indicated the differentiation of anlage of swim bladder.
3.Japanese parrotfish:as a warm water species,lithophilic teleosts,the early developmental feature is:
①fertilized eggs:The yolk was segmented,and acts as a feature of fish egg classification.
②The newly-hatched larve:The melanin on the larval dorsal finfold in the newly-hatched larvae,moved from to the edge of the larval dorsal finfold,and disappeared on 3 dph.This phenomenon indicated the special ontogenetic characteristics of this species.
③post-larvae:The thickness of muscular tissue in intestine increased quickly during this period,and the gut evacuation rised abruptly,at the same time,the mortality rate rise up,and we resolved this problem by the means of changing the routine feeding pattern.During the juvenile stage,the jaw teeth of Japanese parrotfish fused into two whole teeth plates,which was a normal feature of Oplegnathidae.
④some secreting cells were found in the thymus during the larval development, which usually were found in some lower finfish,like salmonids.Through our studies,the numbers of secreting cells in rainbow trout was fewer than Japanese parrotfish,and its arrangement was also devoid of orderliness.MMCs was also normally found in the hemopoietic organs-spleen and head kidney,also found in the parenchyma of liver, pancreas and swim bladder,which usually occurred in elasmobranch fishes and few in teleosts.
引文
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