条石鲷早期发育及相关酶活性的研究
摘要
条石鲷(Oplegnathus fasciatus),隶属鲈形目(Perciformes),石鲷科(Oplegnathidae),主要分布于我国黄海、东海、台湾沿海,日本北海道以南及韩国以南沿海海域。其肉质极为细致,口感好,是一种具有较高食用价值和观赏价值的优良海产鱼类。本文运用统计学、组织学和生理学的分析方法,详细地阐述了条石鲷早期发育阶段的生长模式、组织发育及相关酶活性的变化过程。所得
结果如下:
1.测定了条石鲷仔稚幼鱼(0—50日龄)全长和体重随日龄的生长变化,发现全长、体重随日龄的变化均符合Cubic函数关系式,其变化曲线呈S型。全长随日龄的变化可分为三个阶段,不同阶段的全长生长率明显不同,0-17日龄的全长生长率为0.243mm/d,17-35日龄的全长生长率为0.965mm/d,35-50日龄的全长生长率为0.823mm/d,3个阶段的全长生长率具有显著性差异(P<0.05)。体重随日龄的变化也分为三个明显的阶段,不同阶段的体重生长率明显不同,0-23日龄的体重生长率为8.958*10~(-4)g/d,23-32日龄的体重生长率为0.005 g/d,32-50日龄的体重生长率为0.019 g/d,3个阶段的体重生长率具有显著性差异(P<0.05)。
2.分析了条石鲷仔稚幼鱼头长、头高、体高、眼径、口裂、吻长、腹长、尾鳍长等外部形态参数与全长的变化关系,发现各功能器官的生长均呈现出异速生长的特点,但生长的拐点却各不相同。其结果表明,在条石鲷的早期发育过程中,有关运动、摄食、消化等的重要器官(眼、口、腹部、尾鳍等)具有优先发育的特征,且发育的时段各有差异,故在其苗种的培育过程中,应根据其生长、生态规律,为其创造最佳的环境条件,用以提高苗种的存活率。
3.通过组织学技术(HE染色法、Mallory三色法),观察了条石鲷在胚胎期卵黄多核体的形成、胚层的分化、神经胚的形成、体节的发生、克氏囊的出现及退化、主要器官的组织发生等过程。结果表明,条石鲷的胚胎发育过程有囊胚腔的出现;神经胚以次级神经胚的方式形成;未知嗜红颗粒的渗入导致了克氏囊的退化;眼、脑、心脏、消化管、肾脏等器官均已发生。并通过结合胚胎形态学的观察确定了条石鲷胚胎发育的准确时序,进而更详细地阐述了条石鲷的胚胎发育全过程。
4.通过组织学(HE染色法)技术,对条石鲷的主要功能器官(眼、鳃、心脏、鳔、口咽腔、食道、胃、肠、肝胰脏、肾脏、脾脏、胸腺)的组织发生、细胞分化、器官形成进行了观察,完整描述了条石鲷早期的整个组织发育过程;并辅以汞-溴酚蓝(mercury-bromophenol blue)法对仔鱼的蛋白样物质进行了定位和初步的定量,用以评估胰蛋白酶、胃蛋白酶、碱性磷酸酶、Na~+,K~+-ATP等酶原颗粒发生的时间和数量的变化,以期将条石鲷仔鱼组织器官的发育和相关酶活性的变化相联系;
5.以生物试剂盒测定了条石鲷早期发育过程中胃蛋白酶、碱性磷酸酶、钠钾三磷酸腺苷酶(Na~+,K~+-ATPase)、超氧化物歧化酶(SOD)的活性以及仔鱼可溶总蛋白量的变化(胰蛋白酶以BAPNA法测定),进而分析了以上几种特异性酶的相对活性变化。其中胰蛋白酶、碱性磷酸酶、Na~+,K~+-ATPase与SOD的活性从胚胎期开始就可以被检测到,并随着各自相关组织器官的发育而变化,而胃蛋白酶的活性在仔鱼胃的结构发育完善后(22日龄)才被检测到。胰蛋白酶、胃蛋白酶和碱性磷酸酶属于消化酶,其活性的变化与消化系统的组织发育密切相连;Na~+,K~+-ATPase主要参与离子的交换,其活性变化由泌氯细胞(鳃)决定;SOD的活性是非特异性免疫功能的指标,也反映了仔鱼免疫系统的发育状况。通过分析各特征性酶活性的变化,并结合各相应器官的组织发育,能更好地阐述了条石鲷早期发育的特点。
The rock bream, Oplegnathus fasciatus, a subtropical and carnivorous species, is an economically important marine fish in East Asia. The high commercial and ornamental value makes it a promising aquaculture species in the future. However, to some extent, the lack of information on ontogenetic development has restricted the breeding industry of this species. In this study, the allometric growth, histogenesis and enzymatic activity in rock bream were analyzed. The results are as follows:
1. At the general condition for fingerling-production, the total length and body weight of rock bream larvae were measured from hatching to 50 day after hatching. The increase of total length and body weight could be estimated with the Cubic function and took on the S-Curve. The curve could be divided into three phases and each phase possessed different growth rate.
2. The head length, head height, trunk height, eye diameter, mouth width, abdomen length, tail fin length and rostrum length of rock bream were measured and the relationship between them and the total length was analyzed. The result showed the allometric growth in rock bream larvae. By analyzing the inflexion points in growth curves, in combination with morphological development of the larvae, we found that some important organs (head, mouth, eye, digestive tract and fins) had developed prior to other ones. When rearing rock bream larvae, the best environmental condition should be established by making the important organs prior development.
3. The embryonic development of rock bream was studied from fertilization until hatching (HE & Mallory). The hatching occurred approximately at 25 h after fertilization at 23.5±0.5°C. The embryogenesis is divided into seven stages: Zygote period, Cleavage period, Morula period, Blastula period, Gastrula period, Segmentation period and Hatching period. The first cleavage furrow of rock bream fertilized eggs is vertically oriented, as is usual until horizontal cleavage occurs at the fifth cleavage. The blastocoel is observed between the blastoderm and I-YSL at blastula period. At 90%-epiboly stage, the earliest somitic furrow appears in the middle of embryo. The Kupffer’s vesicle consisting of ventrally I-YSL and dorsally columnar cells appears with the completion of epiboly. It degenerates gradually with the penetration of some eosinophilic granules and disappears completely at 20 h 30 min after fertilization. The digestive tract, a straight tubule, is differentiated into foregut, midgut and hindgut. The staging series provides a preliminary baseline reference for future studies on embryos of the rock bream.
4. The ontogeny of rock bream larvae was investigated by histological observation from hatching to 50 days after hatching (HE & mercury-bromophenol blue). The histogenesis of main organs (eyes, gill, brain, heart, digestive system, immune system and swim bladder) was described in detail. Overall, rock bream presented a similar pattern of histophysiological ontogeny of organs compared with other marine teleosts. The main interspecific variability encountered resides in the timing of organ development.
5. The activity of pepsin, alkaline phosphatase, Na~+ K~+-ATPase and Superoxide Dismutase (SOD) were assayed using test kits. Trypsin activity was measured using Nα-benzoyl-DL-arginine-p-nitroanilide (BAPNA) as substrate. Protein was determined by the Bradford method. The activities of the different enzymes assayed were expressed in relation to soluble protein in the extracts (specific activity). The analysis of enzymatic activity, in combination with the observation of organogenesis, can better illuminate the early development of rock bream.
结果如下:
1.测定了条石鲷仔稚幼鱼(0—50日龄)全长和体重随日龄的生长变化,发现全长、体重随日龄的变化均符合Cubic函数关系式,其变化曲线呈S型。全长随日龄的变化可分为三个阶段,不同阶段的全长生长率明显不同,0-17日龄的全长生长率为0.243mm/d,17-35日龄的全长生长率为0.965mm/d,35-50日龄的全长生长率为0.823mm/d,3个阶段的全长生长率具有显著性差异(P<0.05)。体重随日龄的变化也分为三个明显的阶段,不同阶段的体重生长率明显不同,0-23日龄的体重生长率为8.958*10~(-4)g/d,23-32日龄的体重生长率为0.005 g/d,32-50日龄的体重生长率为0.019 g/d,3个阶段的体重生长率具有显著性差异(P<0.05)。
2.分析了条石鲷仔稚幼鱼头长、头高、体高、眼径、口裂、吻长、腹长、尾鳍长等外部形态参数与全长的变化关系,发现各功能器官的生长均呈现出异速生长的特点,但生长的拐点却各不相同。其结果表明,在条石鲷的早期发育过程中,有关运动、摄食、消化等的重要器官(眼、口、腹部、尾鳍等)具有优先发育的特征,且发育的时段各有差异,故在其苗种的培育过程中,应根据其生长、生态规律,为其创造最佳的环境条件,用以提高苗种的存活率。
3.通过组织学技术(HE染色法、Mallory三色法),观察了条石鲷在胚胎期卵黄多核体的形成、胚层的分化、神经胚的形成、体节的发生、克氏囊的出现及退化、主要器官的组织发生等过程。结果表明,条石鲷的胚胎发育过程有囊胚腔的出现;神经胚以次级神经胚的方式形成;未知嗜红颗粒的渗入导致了克氏囊的退化;眼、脑、心脏、消化管、肾脏等器官均已发生。并通过结合胚胎形态学的观察确定了条石鲷胚胎发育的准确时序,进而更详细地阐述了条石鲷的胚胎发育全过程。
4.通过组织学(HE染色法)技术,对条石鲷的主要功能器官(眼、鳃、心脏、鳔、口咽腔、食道、胃、肠、肝胰脏、肾脏、脾脏、胸腺)的组织发生、细胞分化、器官形成进行了观察,完整描述了条石鲷早期的整个组织发育过程;并辅以汞-溴酚蓝(mercury-bromophenol blue)法对仔鱼的蛋白样物质进行了定位和初步的定量,用以评估胰蛋白酶、胃蛋白酶、碱性磷酸酶、Na~+,K~+-ATP等酶原颗粒发生的时间和数量的变化,以期将条石鲷仔鱼组织器官的发育和相关酶活性的变化相联系;
5.以生物试剂盒测定了条石鲷早期发育过程中胃蛋白酶、碱性磷酸酶、钠钾三磷酸腺苷酶(Na~+,K~+-ATPase)、超氧化物歧化酶(SOD)的活性以及仔鱼可溶总蛋白量的变化(胰蛋白酶以BAPNA法测定),进而分析了以上几种特异性酶的相对活性变化。其中胰蛋白酶、碱性磷酸酶、Na~+,K~+-ATPase与SOD的活性从胚胎期开始就可以被检测到,并随着各自相关组织器官的发育而变化,而胃蛋白酶的活性在仔鱼胃的结构发育完善后(22日龄)才被检测到。胰蛋白酶、胃蛋白酶和碱性磷酸酶属于消化酶,其活性的变化与消化系统的组织发育密切相连;Na~+,K~+-ATPase主要参与离子的交换,其活性变化由泌氯细胞(鳃)决定;SOD的活性是非特异性免疫功能的指标,也反映了仔鱼免疫系统的发育状况。通过分析各特征性酶活性的变化,并结合各相应器官的组织发育,能更好地阐述了条石鲷早期发育的特点。
The rock bream, Oplegnathus fasciatus, a subtropical and carnivorous species, is an economically important marine fish in East Asia. The high commercial and ornamental value makes it a promising aquaculture species in the future. However, to some extent, the lack of information on ontogenetic development has restricted the breeding industry of this species. In this study, the allometric growth, histogenesis and enzymatic activity in rock bream were analyzed. The results are as follows:
1. At the general condition for fingerling-production, the total length and body weight of rock bream larvae were measured from hatching to 50 day after hatching. The increase of total length and body weight could be estimated with the Cubic function and took on the S-Curve. The curve could be divided into three phases and each phase possessed different growth rate.
2. The head length, head height, trunk height, eye diameter, mouth width, abdomen length, tail fin length and rostrum length of rock bream were measured and the relationship between them and the total length was analyzed. The result showed the allometric growth in rock bream larvae. By analyzing the inflexion points in growth curves, in combination with morphological development of the larvae, we found that some important organs (head, mouth, eye, digestive tract and fins) had developed prior to other ones. When rearing rock bream larvae, the best environmental condition should be established by making the important organs prior development.
3. The embryonic development of rock bream was studied from fertilization until hatching (HE & Mallory). The hatching occurred approximately at 25 h after fertilization at 23.5±0.5°C. The embryogenesis is divided into seven stages: Zygote period, Cleavage period, Morula period, Blastula period, Gastrula period, Segmentation period and Hatching period. The first cleavage furrow of rock bream fertilized eggs is vertically oriented, as is usual until horizontal cleavage occurs at the fifth cleavage. The blastocoel is observed between the blastoderm and I-YSL at blastula period. At 90%-epiboly stage, the earliest somitic furrow appears in the middle of embryo. The Kupffer’s vesicle consisting of ventrally I-YSL and dorsally columnar cells appears with the completion of epiboly. It degenerates gradually with the penetration of some eosinophilic granules and disappears completely at 20 h 30 min after fertilization. The digestive tract, a straight tubule, is differentiated into foregut, midgut and hindgut. The staging series provides a preliminary baseline reference for future studies on embryos of the rock bream.
4. The ontogeny of rock bream larvae was investigated by histological observation from hatching to 50 days after hatching (HE & mercury-bromophenol blue). The histogenesis of main organs (eyes, gill, brain, heart, digestive system, immune system and swim bladder) was described in detail. Overall, rock bream presented a similar pattern of histophysiological ontogeny of organs compared with other marine teleosts. The main interspecific variability encountered resides in the timing of organ development.
5. The activity of pepsin, alkaline phosphatase, Na~+ K~+-ATPase and Superoxide Dismutase (SOD) were assayed using test kits. Trypsin activity was measured using Nα-benzoyl-DL-arginine-p-nitroanilide (BAPNA) as substrate. Protein was determined by the Bradford method. The activities of the different enzymes assayed were expressed in relation to soluble protein in the extracts (specific activity). The analysis of enzymatic activity, in combination with the observation of organogenesis, can better illuminate the early development of rock bream.
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