才女虫生物学特征及其寄生行为的基础研究
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摘要
才女虫属复合体(polydora-complex)是环节动物门(Annelida)、多毛纲(Polychaeta)、游走目(Erranta)、海稚虫科(Spionidae)中具变形第5刚节种类的统称。才女虫中的某些种类在含有碳酸钙的基质中钻凿管道,因此是贝类底播养殖业的重要病害之一。本论文以我国重要经济贝类虾夷扇贝和魁蚶寄生的才女虫为研究对象,系统研究了凿贝才女虫的形态结构特征、配子形成过程、幼虫发育过程和寄生行为,揭示了影响凿贝才女虫在虾夷扇贝上寄生数量的关键因素,同时研究了寄生在底播魁蚶上的后刺才女虫的胚胎和幼虫发育过程。详述如下:
     1.凿贝才女虫身体结构相对简单,主要由表皮、肌肉层、消化系统组成。体表有大量的腺细胞分泌粘液,用来腐蚀贝壳和润滑管道,疣足和肌肉发达,方便其挖掘贝壳。
     2.凿贝才女虫精子发生过程中顶体由两个前顶体颗粒融合而成;染色质收缩的过程没有微管参与;染色质在收缩过程中排列整齐;精子核后窝深陷;精子中片线粒体数量少。
     3.凿贝才女虫既可以进行有性繁殖又可以进行无性繁殖。在有性繁殖中卵在卵袋的保护下发育成3刚节浮游幼虫,然后进人水中经15-20d的浮游生活后转为底栖生活,并逐渐筑管隐居营管栖生活。在无性繁殖中身体细胞可以通过分化再生形成新的头部和尾部。
     4.凿贝才女虫有两种寄生方式,一种是筑造泥管寄生(TypeⅠ);另一种是以苔藓虫为媒介寄生(TypeⅡ)。其中TypeⅠ在所有的底质类型中都有分布,TypeⅡ只在沙石底的底质中分布。凿贝才女虫只在活体上寄生。
     5.虾夷扇贝上寄生的才女虫的附着数量随着底质中泥(直径<63μm)含量的增加而增加。凿贝才女虫寄生对2.5龄虾夷扇贝的壳高、干贝指数和干内脏指数没有显著影响。
     6.凿贝才女虫在扇贝贝壳的棱柱层筑造的管道内壁光滑,在扫描电镜下放大观察可以看到呈规则的沟壑状,与凿贝才女虫第五刚节变形的刚毛结构吻合,推测可能是由刚毛摩擦形成的。第五刚节变形刚毛由致密的钙质结构组成,非常坚硬,没有细胞结构,顶端非常尖锐。凿贝才女虫体表有大量的黏液腺,除了分泌粘液润滑身体外还分泌酸性黏液腐蚀贝壳,使贝壳变得相对松软。从以上分析可以推测凿贝才女虫凿贝筑管的过程为先分泌酸性黏液溶解腐蚀身体周围的贝壳,然后通过机械摩擦,用第五刚节变形的刚毛把松动的贝壳碎屑摩擦下来,从而形成光滑弯曲的管道。凿贝才女虫通过触须和口前叶收集沉积物颗粒,分泌粘液把这些颗粒粘附在一起筑造泥管。
     7.后刺才女虫与凿贝才女虫繁殖方式类似,都是把受精卵产在卵袋中,胚胎发育为三刚节幼虫后开始浮游生活,经过一段时间后开始下潜寻找合适的附着基进行附着变态。后刺才女虫幼虫可以捕捉并吞食贝类的D形幼虫。所以在贝类育苗中,才女虫不仅和贝类争夺饵料,还捕食贝类幼虫,给育苗业造成极大地危害。
     本研究开辟了我国贝类养殖业病害研究的新领域,在才女虫基础生物学、生态学、行为学、流行病学等方面取得的研究成果,为我国贝类才女虫病防控提供了重要的理论基础和应用知识。
Spionid worms of the genus Polydora inhabit the shells of many commerciallyimportant bivalves and are an important problem in molluscan aquaculture. TheJapanese scallop Patinopecten yessoensis and the bloody clam Scapharcabroughtoni have been very important commercial bivalves in China. These molluscsare often infested by Polydora ciliata and P. latispinosa, species belonging to theSpionidae, the most common parasitic polychaete group. A study was made of thebiological characters and the parasitic behavior of Polydora. The main results are asfollows.
     1. The body structure of P.ciliata is relatively simple insects mainly by skin,muscle layer, and the digestive system. A large number of gland cells secrete mucus,to pipeline shell and lubrication corrosion, muscular parapodium to facilitate itsexcavation shells.
     2. In P. ciliata, during spermiogenesis, the proacrosomal granules fuse to forma single, spherical acrosomal vesicle that migrates to the anterior pole of the cell.The chromatin condensation is regularly arranged as dense fibrils and it becomesmore condensed as the nucleus elongates. Microtubules do not surround the nucleusduring its elongation. At the time of nuclear condensation, mitochondria becomereduced in number but increase in size, causing deep indentations at the base of thenucleus. The mid-piece has a few mitochondria.
     3. Both sexual and asexual reproduction occurs in P.ciliata. The eggs are laid inegg-sacs attached to the wall of the parent’s burrow. The larvae are released at astage with three segments and lead a long planktonic life. Then each youngPolydora had collected debris and formed a tube. Asexual reproduction occursthrough architomy, in which a parent fragment regenerates a complete body.
     4. The first type was P. ciliata secreted mucus to form fragile mudtubes in theburrows with openings projected approximately2cm above the scallop shell surface.The other type was P. ciliata infested the scallop with the Bryozoa as the medium.
     5. The number of worms per infested scallop inhabiting sites with differentenvironmental conditions increased with increasing mud content (d<63μm) in thebottom sediments. The boring of P.ciliata showed no significant effect on the shellheight of Japanese scallops Patinopecten yessoensis.
     6. Scanning electron microscopy revealed many characteristic gully-likescratches on the surface of the burrows. The organs involved in rectilinear motion,such as the heavy spines of the fifth setiger, may be related to the horizontal scatcheson the surface of burrows. While tube-building P. ciliata lies with its head just visibleinside the rim of the tube. The palps emerge from the tube and are applied to thesubstratum with the ciliated groove downwards. The tips of the palps are forcedamong the bottom debris and sand grains are dislodged and carried posteriorly towardthe mouth. The ciliated groove ends short of the base of the palp and the mucus stringis forced out and met by the prostomium which bends towards it. The string is pickedup by the densely ciliated ventral epithelium of the prostomium and is carried towardsthe mouth.
     7. Both P. ciliata and Polydora latispinosalaid their eggs in egg-sacs. The larvaeare released at a stage with three segments and lead a long planktonic life. The larvaeof P. latispinosacan capture the D-shaped larvae of shellfish. So in molluscanaquaculture, the Polydora not only compete for food, but also prey shellfish larvae tocause great harm the nursery industry.
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