湖泊科学   2018, Vol. 30 Issue (2): 458-463.  DOI: 10.18307/2018.0217.
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研究论文

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姜涛, 刘洪波, 李孟孟, 杨健, 溯河洄游长江刀鲚(Coilia nasus)摄食虾类的调查. 湖泊科学, 2018, 30(2): 458-463. DOI: 10.18307/2018.0217.
[复制中文]
JIANG Tao, LIU Hongbo, LI Mengmeng, YANG Jian. Investigation on shrimp feeding of Coilia nasus during its anadromous migration along the Yangtze River. Journal of Lake Sciences, 2018, 30(2): 458-463. DOI: 10.18307/2018.0217.
[复制英文]

基金项目

中央级公益性科研院所基本科研业务费专项(2016PT01)、国家自然科学基金项目(31372533)和江苏省自然科学基金项目(BK20131089,BK20160204)联合资助

作者简介

姜涛(1984~), 男, 助理研究员; E-mail:jiangt@ffrc.cn

通信作者

杨健, E-mail:jiany@ffrc.cn

文章历史

2017-04-24 收稿
2017-07-07 收修改稿

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溯河洄游长江刀鲚(Coilia nasus)摄食虾类的调查
姜涛 1, 刘洪波 1, 李孟孟 2, 杨健 1,2     
(1: 中国水产科学研究院淡水渔业研究中心, 长江中下游渔业生态环境评价与资源养护重点实验室, 无锡 214081)
(2: 南京农业大学无锡渔业学院, 无锡 214081)
摘要:刀鲚(Coilia nasus)是长江的名贵经济鱼类.虽然传统上认为其溯河生殖洄游全过程不会摄食,但该问题一直尚未完全弄清.作为有效解明这一问题的第一步,本研究在前期长江流域干流和湖泊刀鲚资源调查的基础上,先利用耳石微化学技术筛选出长江河口区、江苏江段、安徽江段和鄱阳湖水域溯河洄游型的刀鲚个体,再对其胃、肠容物的大型游泳动物(鱼、虾类)进行调查分析.结果发现,在河口区、江苏和安徽江段所有刀鲚个体胃充塞度为0级,均未发现摄食有游泳动物;而鄱阳湖水域刀鲚个体的胃充塞度达4~5级,均发现有摄食淡水虾类的日本沼虾(Macrobrachium nipponense)和秀丽白虾(Exopalaemon modestus)的情况.其中29%和71%的刀鲚胃中分别含有1只和2只虾.所有刀鲚个体肠内均未发现有内容物.鉴于鄱阳湖已被确定为溯河洄游型刀鲚的产卵场之一,结果表明长江刀鲚在经河口,通过长江江苏和安徽江段干流到达鄱阳湖产卵场的过程中应该不会摄食大型游泳动物;而进入鄱阳湖产卵场后会开始摄食日本沼虾和秀丽白虾.这种两阶段的现象可能反映出了刀鲚的一种在洄游通道上节约能量,以利长距离溯河;而在产卵场补充能量,以利于性腺最终成熟的生存策略.
关键词刀鲚    溯河洄游    摄食    日本沼虾    秀丽白虾    长江    鄱阳湖    
Investigation on shrimp feeding of Coilia nasus during its anadromous migration along the Yangtze River
JIANG Tao 1, LIU Hongbo 1, LI Mengmeng 2, YANG Jian 1,2     
(1: Key Laboratory of Fishery Eco-environment Assessment and Resource Conservation in Middle and Lower Reaches of the Yangtze River, Freshwater Fisheries Research Center, Chinese Academy of Fishery, Wuxi 214081, P. R. China)
(2: Wuxi Fisheries College, Nanjing Agricultural University, Wuxi 214081, P. R. China)
Abstract: Estuarine tapertail anchovy(Coilia nasus) is a valuable commercial fish in the Yangtze River. It is traditionally believed that C. nasus do not feed during their anadromous migration, but this issue is not yet fully clarified. As a pilot study on feeding of C. nasus during the whole process of anadromous migration, the present study investigated the nekton(e. g. shrimp, fish) in stomach and intestinal contents of C. nasus fish (confirmed as anadromous individuals by otolith microchemical analysis) previously collected in the Yangtze River's estuary, Chongming Island of Shanghai City, sections in Jiangsu and Anhui Provinces, and connected Lake Poyang in Jiangxi Province. The results showed that C. nasus from the river estuary and sections of Jiangsu and Anhui Provinces did not feed nekton and their stomach fullness indices were at 0 degree. In contrast, all C. nasus from Lake Poyang obviously fed nekton and their stomach fullness indices were at 4 to 5 degree. Furthermore, the nektons in stomach contents of the fish were all freshwater shrimps (Macrobrachium nipponense and Exopalaemon modestus), and one and two prawns were found in the stomach contents of 29% and 71% of the C. nasus fish, respectively. However, no nekton residual could be found in the intestinal contents of C. nasus studied. Since the Lake Poyang has been confirmed as a spawning site of anadromous C. nasus, the present study revealed a possible two-step-feeding process for the fish in the Yangtze River, namely, C. nasus may not feed (at least nekton) when moving upstream through the estuary and migratory channel in the sections of Jiangsu and Anhui Provinces, while they may start to feed shrimps of M. nipponense and E. modestus when entering the spawning site in the Lake Poyang. These phenomena may reflect a survival strategy of Yangtze C. nasus, i. e., the fish keeps fasting to save energy for its long distance anadromous movement, but starts to feed to supply energy for full gonad maturation and finish breeding.
Keywords: Coilia nasus    anadromous migration    feeding    Macrobrachium nipponense    Exopalaemon modestus    Yangtze River    Lake Poyang    

刀鲚(Coilia nasus Temminck et Schlegel,1846)是我国的一种名贵鱼类,位居“长江三鲜”之首,历来也是我国长江流域重要的经济鱼类.由于近年来酷渔滥捕、水体污染和生境破坏等原因,虽然受到《长江刀鲚凤鲚专项管理暂行规定》、海洋伏季休渔政策等制度的保护,但刀鲚资源现状仍日渐堪忧.即便如此,人们对其依旧趋之若鹜,其售价更曾创造过单条(全长45.3 cm,体重325 g)5.9万元的天价.由于刀鲚是小型溯河洄游鱼类,活动范围覆盖海水、河口半咸水和淡水生境,即使在长江跟踪研究其全洄游过程亦十分困难,使得其很多生活习性(如溯河洄游过程中是否摄食等)尚未完全解明.这给有效地养护这种珍贵鱼类资源带来了极大的不便.历史上,长江刀鲚最远可以上溯至洞庭湖水域进行产卵繁殖[1],但目前其已很难上溯至该水域[2-4].笔者研究结果显示,其目前自河口上溯的最远位置可能为距长江口约1000 km的鄱阳湖上游信江水域[5].前人的研究认为刀鲚亲鱼在溯河繁殖洄游的全过程中并不进行摄食[6-8].然而,笔者在鄱阳湖星子—都昌一带所确认的长江刀鲚一个产卵场[9-10]水域中却发现洄游型刀鲚有摄食虾类的情况.鉴于这个发现与前人的观点相左,本研究中拟利用近期在刀鲚溯长江洄游路径沿线的河口上海崇明、江苏江段、安徽江段和江西鄱阳湖水域所获刀鲚成鱼,对其摄食游泳动物(如虾类、鱼类)的情况进行较为详细地调查.

需要注意的是,长江流域中除了分布有洄游型的刀鲚群体外,还分布有淡水定居型群体[1].传统的分析方法认为前者为长颌鲚,后者为短颌鲚.但在利用更为客观和准确的耳石微化学分析技术[11-12]进行相关研究后发现短颌鲚中存在有溯河洄游型的个体[13],长颌鲚中也存在有淡水定居型的个体[14].因此,为了更准确地达到研究目的,本研究将在上述各水域采集长江刀鲚并获取耳石的基础上,先利用耳石微化学技术筛选出溯河洄游型刀鲚个体,再分析其胃、肠容物中游泳动物的出现和残留状况.本研究期望一方面确证长江刀鲚在经河口通过长江江段洄游通道进入产卵场繁殖全过程中摄食以虾类为主游泳动物的情况,另一方面也为今后全面开展长江刀鲚溯河洄游过程中的摄食特征及能量利用和相关生存策略研究打下基础.

1 材料与方法

2008—2010和2013—2015年采自长江流域的上海崇明长江口,江苏的常熟、靖江、南京江段,安徽的和县、芜湖、无为、安庆江段以及江西鄱阳湖的湖口、星子、都昌、余干水域的刀鲚标本351尾,选取其中的179尾溯河洄游型长江刀鲚的个体(表 1)来调查其经过洄游路径沿线不同水域过程中的摄食虾类情况.鱼体性腺成熟度根据解剖后性腺的大小、颜色等特征进行目视判别[4, 15-16].

表 1 本研究所用长江不同水域溯河洄游型刀鲚标本的基本情况 Tab.1 Sampling details of anadromous Coilia nasus from different water areas along the Yangtze River

所有个体按惯例摘取左耳石为研究对象,利用耳石微化学方法分析其Sr和Ca元素含量特征,并以Sr/Ca×103值是否有大于3的阶段(对应于矢状面二维Sr元素分布图为绿色乃至黄色的高锶阶段,图 1)来确认其是否为溯河洄游型[9-10, 12](本研究只分析这类洄游型个体),之后解剖其胃和肠道,目视观察记录其充塞度.充塞度的级别依次分为: 0级:空胃; 1级:胃中有少量食物; 2级:食物占胃的1/2; 3级:食物较多,食物占胃的3/4; 4级:食物充塞全部胃; 5级:胃膨胀[17].随后剖开胃和肠道以分析其摄食情况.参考堵南山[18]和李隼等[19]等的特征描述来进行刀鲚胃肠中日本沼虾(Macrobrachium nipponense)、秀丽白虾(Exopalaemon modestus)等的鉴定和计数,如日本沼虾额剑上缘平直、秀丽白虾呈鸡冠状凸起且前段无齿,日本沼虾头胸甲有肝刺而秀丽白虾头胸甲有腮甲刺等.

图 1 典型溯河洄游型刀鲚耳石矢状面二维Sr元素的微化学图谱 Fig.1 A typical microchemical map of two-dimensional image of the Sr content in the otoliths of Yangtze anadromous Coilia nasus
2 结果与讨论

本研究所调查的所有洄游刀鲚经解剖后目视观察发现,长江河口区30尾个体、江苏江段41尾个体以及安徽江段59尾个体胃和肠道中均未发现有任何明显的游泳动物(虾类、鱼类)残留,其充塞度均为0级.而在鄱阳湖各水域采集的刀鲚个体100 %有明显的摄食迹象,胃容物中普遍有虾类(经鉴定为日本沼虾和秀丽白虾,图 2),其充塞度达4级(胃中有1只虾,性腺成熟度♂Ⅲ~♂Ⅳ、♀Ⅳ~♀Ⅴ期)到5级(胃中有2只虾,性腺成熟度♂Ⅳ和♂Ⅵ、♀Ⅲ~♀Ⅳ期),分别占刀鲚样本总数的29 %和71 %,而肠道充塞度均为0级,未发现有任何明显肠容物.从刀鲚摄食的虾种类看,胃内仅有秀丽白虾的有12尾(24.5 %),仅有日本沼虾的有27尾(55.1 %),同时有这2种虾的有10尾(20.4 %).由此可见刀鲚摄食虾类的行为具有某种选择性,蔡德霖等[20]和凡迎春等[21]的研究结果亦可得出这种结论.这可能显示虾除了可以提供较高的能量外,也可能与刀鲚较为特殊的颌骨结构(上颌骨较长,后端呈游离状且上下颌骨均分布有细齿)易于捕食这类饵料动物有关.鄱阳湖刀鲚胃中出现日本沼虾较多的情况也可能与该虾在鄱阳湖内为优势种,且在全湖呈均匀分布有关[22].与之相比,淀山湖内日本沼虾与秀丽白虾均为优势种,两者资源量相近[23],因此导致陆封型刀鲚胃中出现秀丽白虾与日本沼虾数量相近的现象[21].

图 2 鄱阳湖中溯河洄游型刀鲚胃容物中的秀丽白虾(a)和日本沼虾(b) Fig.2 Exopalaemon modestus (a) and Macrobrachium nipponense (b) in stomach contents of anadromous Coilia nasus collected from Lake Poyang

迄今的研究报道均认为长江刀鲚在上溯过程中会停止摄食[6-8].这与本研究长江口、江苏(近河口干流洄游通道)和安徽江段(下游干流洄游通道)刀鲚的胃、肠充塞度均为0级相吻合,同时也与其上溯过程中肥满度指标不断下降的结果相印证[24].然而,本研究在更为上游的鄱阳湖水域(产卵场)内所获刀鲚的胃中均发现有摄食迹象(充塞度甚至可达4~5级).其内清晰可见尚未消化的虾的情况更直观地反映了这些个体刚摄食完虾的行为.由此可见,长江刀鲚在上溯过程的终期(即进入产卵场后)应该会再行摄食(图 3),但胃饱满度高低与刀鲚性别与发育程度无明显关系,如胃饱满度4级的个体性腺成熟度可为♂Ⅲ~♂Ⅳ期、♀Ⅳ~♀Ⅴ期,胃饱满度5级的个体性腺成熟度可为♂Ⅳ和♂Ⅵ期、♀Ⅲ~♀Ⅳ期.

图 3 长江流域不同水域洄游型刀鲚的胃充塞度的动态变化(0级:空胃; 1级:胃中有少量食物; 2级:食物占胃的1/2; 3级:食物较多,食物占胃的3/4; 4级:食物充塞全部胃; 5级:胃膨胀[17]) Fig.3 The dynamics of stomach fullness of anadromous Coilia nasus from different water areas along the Yangtze River (0: empty; 1: traces of food; 2: half full; 3: 3/4 full; 4: full; 5: bursting[17])

对于长距离溯河生殖洄游的鱼类而言,其上溯过程中能量的合理利用十分重要[25].多数鱼类主要依靠其洄游前积累在脂肪中的能量进行生殖洄游[26-27].这部分能量除了很大一部分用以洄游至产卵场的运动消耗外,还有部分将用于性腺的最终成熟[28].长江刀鲚目前最远可上溯约1000 km至鄱阳湖上游的信江水域[5].如此长距离的洄游对于这种20~40 cm长的小型鱼类而言,能量的合理利用和分配较其他大型鱼类(如大麻哈鱼)显得更为重要[29].本研究结果显示,长江刀鲚具有特殊的繁殖对策,据其独特的摄食行为可将生殖洄游大致分为2个阶段: 1)不摄食上溯阶段,即当通过洄游通道(长江河口、江苏段以及安徽段)时亲鱼不进行摄食,此时其性腺成熟度较低,主要依靠其在上溯前(即越冬前期)所积累体内脂肪的消耗[30]来保证其长距离溯河; 2)摄食补充能量阶段,即当刀鲚在进入产卵场的鄱阳湖水域[10]后,可能会由于体内积累脂肪的能量基本消耗殆尽而开始摄食,以补充急需的能量并完成性腺的最后成熟和生殖活动.这样一方面可能缩短在洄游通道中所消耗的时间,规避亲鱼被捕食的风险; 另一方面也解决了小型鱼类长距离洄游时由于体型制约,大量积累的能量用于快速上溯,导致最终繁殖时可能无法满足性腺最终成熟和完成繁殖行为等所需能量消耗的问题.该水域的刀鲚捕食日本沼虾和秀丽白虾(图 2)可能是其快速补充能量消耗的一种策略.相似情况在前人黄河刀鲚的调查中也有发现[20].刀鲚进入东平湖(黄河刀鲚的传统产卵场)前不摄食比例占92.6 %以上,而入湖后摄食个体的比例占到81.7 %,且以虾类为主要摄食对象的更占到90.6 %,以此补充上溯时的能量消耗[20].

本研究刀鲚肠道内未见有明显的食物残渣.这一方面可能显示刀鲚在抵达鄱阳湖后刚开始摄食虾类,另一方面也可以佐证刀鲚在下游的河口区到安徽和江苏江段洄游过程中无明显摄食的现象.至于刀鲚是否在洄游过程中存在捕食其他饵料(如浮游生物等)的可能性,需要在下一步的研究中利用DNA条形码或者二代测序等技术进行深入地调查和确证.

综上所述,本研究证实长江刀鲚溯河生殖洄游过程中可能存在到达产卵场前不会明显摄食大型游泳动物,而到达产卵场后却会明显摄食这类动物(如虾类)2个阶段.而这种现象所包含的能量利用和生存策略很有可能是决定这种小型鱼类能够成功上溯约1000多km并完成繁殖活动的关键,亟需今后开展详细的调查和探索.

致谢: 本文中虾类的鉴定得到了中国水产科学研究院淡水渔业研究中心的傅洪拓研究员、龚永生副研究员和张敏莹副研究员的大力支持和帮助,在此一并致以诚挚的谢意.
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