绿鳍马面鲀胚后早期发育各阶段形态特征的研究.pdf

第 45 卷 第 2 期 渔 业 科 学 进 展 Vol.45,No.2 2 0 2 4 年4 月 PROGRESS IN FISHERY SCIENCES Apr.,2024 *中国水产科学研究院黄海水产研究所基本科研业务费(20603022022014)资助。朱金超，E-mail: 通信作者：陈四清，研究员，E-mail: 收稿日期:2022-12-08,收修改稿日期:2023-02-13 DOI:10.19663/j.issn2095-9869.202212080002 http:/ ZHU J C,BIAN L,PAN Y,LI F H,PAN L Y,ZHANG Z Y,WU D,CHANG Q,CHEN S Q.Morphological characteristics of postembryonic early development of the Thamnaconus septentrionalis.Progress in Fishery Sciences,2024,45(2):210219 绿鳍马面鲀胚后早期发育各阶段形态特征的研究*朱金超1,2 边 力2 潘 滢3 李凤辉2 潘鲁莹2 张子阳2 吴 丹2 常 青2 陈四清2(1.上海海洋大学水产与生命学院 上海 201306；2.中国水产科学研究院黄海水产研究所 农业农村部海洋渔业与可持续发展重点实验室 山东 青岛 266071；3.福建省农业科学院生物技术研究所 福建 福州 350003)摘要 本文研究了绿鳍马面鲀(Thamnaconus septentrionalis)胚后早期发育各阶段可观、可量和可数性状的变化。结果显示，在水温为(23.01.0)条件下，1 d 出现胸鳍；4 d 卵黄囊消失，鳔基和第 1 背鳍鳍基清晰可见；7 d 油球消失，出现腹鳍；8 d 第 1 背鳍鳍基和腹鳍出膜；9 d 披透明圆形小鳞；12 d 鳔和腹两侧后匙骨清晰可见；14 d 出现第 2 背鳍支鳍骨和臀鳍支鳍骨；16 d 披锥形小鳞，上、下颌出现锥形齿；19 d 分化出第 2 背鳍鳍条和臀鳍鳍条；20 d 尾柄下侧长出尾鳍鳍条；24 d 时，第 2 背鳍、臀鳍和尾鳍的鳍条形态发育完全；27 d 鳔大量充气。可量性状变化的曲线方程 R2 值均0.90，第 1 背鳍鳍高，430 d 呈指数递增；视囊 011 和 1120 d 为线性递增，011 d 递增程度0.90。第 1 背鳍鳍高在 430 d 呈指数递增；视囊 011 和 1120 d 为线性递增，011 d 递增程度1120 d，2130 d 为指数递增；口裂 413 和 2230 d 为线性递增，1322 d为线性递减；其余性状在拐点前为线性递增，拐点后为指数递增，拐点集中在 1923 d。各性状拐点的日龄和大小分别为：全长为20 d，(3.580.34)mm、体长为20 d，(3.320.18)mm、体高为20 d，(0.940.16)mm、耳石腔长为23 d，(0.3930.021)mm、耳石腔高为20 d，(0.3440.029)mm、矢耳石直径25 d，(0.0680.028)mm、视囊直径为11 d，(0.2820.059)mm；20 d，(0.714 0.042)mm、口裂为13 d，(0.5210.034)mm；22 d，(0.3250.028)mm、鳃盖长(19 d，0.4750.049)mm、鳃盖高为19 d，(0.412 0.024)mm、肝脏长为19 d，(0.5120.021)mm、肝脏高为16 d，(0.4250.029)mm、鳔长为21 d，(0.5150.042)mm、鳔高为19 d，(0.3860.034)mm。3 讨论 3.1 绿鳍马面鲀胚后早期发育可观和可数性状 本研究及已有的报道均表明(薛美岩,2011;张哲等,2021)，绿鳍马面鲀早期生长发育变化迅速，存在“变态”发育现象，与棘头梅童鱼、白条双锯鱼和北极茴鱼(刘俊果等,2018;吕爽等,2021;邢君霞等,2022)等仔、稚鱼的研究结果相似，仔、稚鱼和早期幼鱼三者之间形态存在明显差距。绿鳍马面鲀在 4 d吸收完卵黄囊，与蒙古鲌(Erythroculter mongolicus)和草海鲫鱼(Carassius auratus)(姜海峰等,2016;王金娜等,2023)卵黄囊消失时间相一致，而冷水性鱼类哲罗鲑(张永泉等,2008)卵黄囊消失时间则持续约 30 d，这一差异主要与其温度适应性和外界生物饵料有关。绿鳍马面鲀、蒙古鲌和鲫鱼苗种培育水温均为 23 左右，在自然条件下，水温为 23 时，外界生物饵料相对充足，及时有效开口可保证子代获得最大的存活率。油球于开口摄食后 34 d 吸收完全，与双棘刺尻鱼(Centropyge bispinosa)、拟 赤 梢 鱼(Pseudaspius leptocephalus)和北极茴鱼(杨建等,2021;李琰等,2022;邢君霞等,2022)等研究结果类似，原因可能是仔鱼刚进入外源性营养阶段，游动能力相对较弱，残留的油球可以提供一定的浮力，方便捕食，李琰等(2022)的研究也表明这一观点；另外，作为重要营养和能量物质，可降低仔鱼因开口无食而死亡的概率，以保证子代获得最大的存活率。本研究观察到的背部鳍膜伞形透明腺体组织，尚未见其他文献报道，猜测为早期辅助呼吸器官或鳔基充气所用。一方面鱼体刚破膜，鳃部处于分化阶段，观察 06 d 也未发现明显的鳃部结构；另一方面，此时鱼体变化迅速，代谢较旺盛，耗氧较大，鳃部供氧可能不足；同时，其下小管连接在腹腔顶部鳔基前端，鳔基出现时，其结构大小发生了明显变化。鳍条出现时序为胸鳍(1 d)、第 1 背鳍(4 d)、腹鳍(7 d)、第 2 背鳍(19 d)、臀鳍(19 d)和尾鳍(20 d)，与黄条、棘头梅童鱼和北极茴鱼(马宾等,2019;李文康等,2022;邢君霞等,2022)等出现时序一致；在观察中发现，第1 背鳍和腹鳍的变化与日龄相关性更大，第 2 背鳍和臀鳍的变化与全长相关性更大。原因可能是第 1 背鳍和腹鳍出现时间较早，个体间全长差异相对较小，同时，第 1 背鳍和腹鳍的出现除维持身体平衡外，也是仔鱼早期主要御敌手段。一方面鱼体在遇到危险时，二鳍均可竖起，“体高”会显著增加，吓阻捕食者；另一方面，二鳍鳍棘布满锥形倒刺易刺伤捕食者，从而降低捕食者对其仔、稚鱼的偏好，提高种群存活率，关健等(2011)的研究也表明这一点。第 2 背鳍和臀鳍出现时间点较晚，此时，鱼体已经披鳞，消化器官基本发育完善，游动能力较强，具备一定的御敌能力，第 2 背鳍和臀鳍主要作用是辅助平衡和游动；另在鱼体体长到达一定程度前，摄取的能量主要用于体长、体高的增加和体表鳞片的形成，以快速度过危险期。3.2 绿鳍马面鲀胚后早期发育可量性状 研究发现，绿鳍马面鲀胚后早期发育与多数鱼类一样(李军等,2017;徐永江等,2019;马宾等,2019)，表现为异速生长模式，也与薛美岩(2011)的研究相符；生长拐点集中在 1923 d，与张哲等(2021)的研究相一致。异速生长的主要原因：首先是早期发育营养源自卵黄囊和轮虫，外加仔鱼无法消化轮虫卵，营养较单一；其次是仔鱼生长初期，捕食能力差，消化系统不完善。后期投喂卤虫阶段，一方面饵料营养丰富，种类增多；另一方面鱼体消化系统较完善，摄食能力较强，杨建等(2021)研究拟赤梢鱼的生长发育也表明这一观点。有研究表明，仔、稚鱼这种异速生长特性与其生态习性相适应，有利于鱼体适应环境变化和提高存活率(Khemis et al,2013;Bignami et al,2013)。本研究中，430 d 内，第 1 背鳍鳍高呈指数增长，呈平缓趋势，预估后期幼鱼阶段呈线性增长，其与全长比 216 渔 业 科 学 进 展 第 45 卷 图 3 绿鳍马面鲀胚后早期发育生长曲线 Fig.3 Postembryonic early development and growth curve of T.septentrionalis 第 2 期 朱金超等:绿鳍马面鲀胚后早期发育各阶段形态特征的研究 217 在 27 d 达到最大值(0.583)，之后比值逐渐变小，这与鱼体后期快速生长和第 1 背鳍基本发育完全有直接关系。口裂呈先大后小再大的变化趋势，原因可能是前期口裂日益增大，一方面有助于提高捕食成功率，另一方面提高单位捕食次数的捕食量；至一定阶段，生长趋于稳定，上、下颌骨开口因口膜包裹日渐愈合，口裂因此变小，即表明鱼体由吞食性逐渐转为嘬食性；后期的增大则是随着鱼体的生长而变化。参 考 文 献 BIAN X D,WAN R J.Morphogenesis and taxonomic identification of larvae and juveniles in genus Myctophum rafinesque.Journal of Fisheries of China,2014,38(10):17311746 卞晓东,万瑞景.灯笼鱼属仔稚鱼的发育形态及其分类检索.水产学报,2014,38(10):17311746 BIGNAMI S,SPONAUGLE S,COWEN R K.Response to ocean acidification in larvae of a large tropical marine fish,Rachycentron 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YIN M C.Advances and studies on early life history of fish.Journal of Fisheries of China,1991(4):348358 殷名称.鱼类早期生活史研究与其进展.水产学报,1991(4):348 358 ZHANG Y Q,YIN J S,JIA Z H,et al.Embryonic and larval development in Hucho taimen.Journal of Dalian Ocean University,2008,23(6):425430 张永泉,尹家胜,贾钟贺,等.哲罗鱼胚胎和仔鱼发育的研究.大连水产学院学报,2008,23(6):425430 ZHANG Z,JIANG L L,WANG Z,et al.Growth,development,and feeding characteristics of Thamnaconus septentrionalis during early stages.Marine Sciences,2021,45(1):113 张哲,姜良龙,王臻,等.绿鳍马面鲀早期生长发育与摄食特性的研究.海洋科学,2021,45(1):113 (编辑 陈 严)Morphological Characteristics of Postembryonic Early Development of the Thamnaconus septentrionalis ZHU Jinchao1,2,BIAN Li2,PAN Ying3,LI Fenghui2,PAN Luying2,ZHANG Ziyang2,WU Dan2,CHANG Qing2,CHEN Siqing2(1.College of Fisheries and Life Sciences,Shanghai Ocean University,Shanghai 201306,China;2.Yellow Sea Fisheries Research Institute,Chinese Academy of Fishery Sciences,Key Laboratory of Marine Fisheries and Sustainable Development,Ministry of Agriculture and Rural Affairs,Qingdao 266071,China;3.Institute of Biotechnology,Fujian Academy of Agricultural Sciences,Fuzhou 350003,China)Abstract The morphological characteristics,ecological habits,and physiological structures of fish vary rapidly in the early growth and development stage.Understanding the changes in the postembryonic early development morphology of organs or tissues could provide basic information for the classification or identification of fish and the investigation or utilization of early resources.Morphological characteristic observation is one of the most commonly used methods.The existing studies on the early development of a certain trait in the late larval and juvenile stage are unclear.Most of the digital images are side views,resulting in a lack of dorsal views,ventral views,or local images,which is not conducive to the understanding of larval and juvenile.In this study,the variations in observable,measurable,and countable traits in postembryonic early development of Thamnaconus septentrionalis were explored using optical microscopy and stereomicroscopy.The observable traits included the developmental changes of yolk sac,scales,and notochord curvature.The measurable traits included the growth changes of total length,body length,body height,and oral fissure.The countable characters included the number of fins in the second dorsal fin,anal fin,caudal fin,and the number of pterygiophore.The results showed that at(23.01.0),the newly hatched larvae moved up and down in the water,with a phenomenon of convergent clustering under light irradiation;the pectoral fin appeared at 1 d,and the heart rate was approximately 50 beats/min;the optic capsule was completely black,the maxillar appeared,the oral fissure extended below the eyes,the yolk sac disappeared,and the first dorsal fin base appeared at 4 d;the girdle was single and colorless and began from the front edge of the belly,the oil globule disappeared,and the pelvic fin base appeared at 7 d;the first dorsal fin base and pelvic fin base broke membrane at 8 d;the first dorsal fin and pelvic fin were significantly longer than before,transparent rounded scales were observed at 9 d;swimming ability was improved,indicated by hovering back and forth gnawing Corresponding author:CHEN Siqing,E-mail: 第 2 期 朱金超等:绿鳍马面鲀胚后早期发育各阶段形态特征的研究 219 pool wall,transversally conical barbs appeared at the base of the first dorsal fin at 11 d;the swim bladder and postcleithrum on both sides of the belly were clearly visible at 12 d;the second dorsal fin and anal fin pterygiophore appeared at 14 d,the second dorsal fin pterygiophore spans seven sarcomeres and the anal fin pterygiophore spans six sarcomeres.There are 1012 melanin masses in the second dorsal fin and 56 in the anal fin,the conical teeth appeared on the upper and lower jaws at 16 d;the second dorsal fin and anal fin were differentiated at 19 d;the swim bladder fills about 25%of the abdominal cavity;notochord curves upward clearly and the stalk of caudal fin came out at 20 d;the fin strips of the second dorsal fin,anal fin,and caudal fin were well-developed at 24 d;the open mouth was round and changed from swallowing to sucking at 30 d,and the rainbow cells were colorful and the body surface near the gill was the most dense;the abdominal cavity has a strong metallic texture,the hard spine of the pelvic fin was degraded,close to the girdle at 39 d;45 black bands were arranged regularly on the body,and the lateral line was consistent with the vertebral column at 50 d.The R2 values of the curve equations of measurable traits were all greater than 0.90.The height of the first dorsal fin increased exponentially 430 d.The visual sac increased linearly from 011 d to 1120 d,and the increment of 011 d was less than that of 1120 d,while the increment of 2130 d was exponential.The oral fissure in 413 d and 2230 d increased linearly,while that in 1322 d decreased linearly.The total length,body length,and body height increased linearly before the inflection point and exponentially after the inflection point,which was concentrated in 1923 d.The number of fin pterygiophore of the second dorsal fin was 3136,while that in the anal fin was 3335.The number of pectoral fins and caudal fins were 1518 and 12,respectively.In conclusion,metamorphosis occurs in the postembryonic early development of T.septentrionalis,showing an allometric growth pattern,with a slow linear growth in the rotifer-feeding stage and a fast exponential growth in the artemia-feeding stage.The changes in the first dorsal fin and pelvic fin were correlated with age in days.The changes in the second dorsal and anal fins were correlated with the total length.This study enriches the early biological theory of T.septentrionalis and provides basic data for the investigation or utilization of its seedling breeding and resources.The partial enlarged detail is helpful for scholars to further understand the larval or juvenile fish and perform deep research.Key words Thamnaconus septentrionalis;Larvae;Juvenile;Early development;Morphological characteristics