鱼虱科

管口水蚤目的一科甲壳动物

海虱桡脚类下的鱼虱科生物。其下共有36个,当中最著名的疮痂鱼虱属鱼虱属就分别约有42和300个物种[2]它们是水生的 体外寄生虫,吃寄主的黏液、表皮组织及血液。其下的疮痂鱼虱属及鱼虱属会寄生在海鱼身上,尤其对饲养的鲑鱼造成很大的问题。

鱼虱科
雄性及雌性的鲑疮痂鱼虱
科学分类 编辑
界: 动物界 Animalia
门: 节肢动物门 Arthropoda
亚门: 甲壳亚门 Crustacea
纲: 桡足纲 Copepoda
目: 管口水蚤目 Siphonostomatoida
科: 鱼虱科 Caligidae
Burmeister, 1834[1]
[2]

见正文

异名
  • Eirgidae
  • Euryphoridae
  • Pargulidae

多样性

就海虱的认识主要是来自对饲养鲑鱼的感染。不过,就其生物学及与寄主间的关系所知甚少。

很多海虱都有特有的寄主,例如鲑疮痂鱼虱就特别喜欢寄生在鲑鱼当中,尤其是饲养的大西洋鲑。鲑疮痂鱼虱也可以寄生在其他鲑鱼,但程度各有不同。太平洋的鲑鱼,如银大麻哈鱼大麻哈鱼驼背大马哈鱼等对于鲑疮痂鱼虱会产生强烈的组织反应,在感染的首周就会将之排出。[3]太平洋的鲑疮痂鱼虱也可以发展到寄生到三刺鱼[4],不过大西洋的则没有出现这种情况。

海虱在浮游阶段如何扩散及寻找寄主仍然是迷。温度、光线及水流都是主因,其生存倚赖25‰以上的盐度[5][6][7][8]有指鲑疮痂鱼虱会向着光源向上迁徙,正好会遇到向下游的鲑鱼鱼秧。[9]潮间带浮游阶段的海虱可以被送到几十公里以外的地方。[7][10][11]

海虱会从溯河的鱼类身体上掉下及死去,例如回到淡水河流繁殖的鲑鱼。不过这些鲑鱼回到海中重新受感染的原因则不明。有指海虱可能是在河口生活的鱼类身体上存活下来,或是转往其他的寄主。至于海虱如何在野外鱼类之间传播也不详。成年的疮痂鱼虱属实验室环境下可以在不同鱼种间传播,但并不频繁;而鱼虱属则随时可以传播。

形态

鲑疮痂鱼虱的大小较鱼虱属大一倍。它们的身体有4个部分,包括头胸部、第四足体节、生殖部分及腹部[12]头胸部是一块阔板,当中包含了头三足的体节,可以像吸盘般将它们吸在寄主鱼类的身体上。所有的海虱都有像吸管的口器。第二触角及口附肢都进化到可以帮助将它们找著寄主。雄虱也会用第二触角来找住雌性进行交配[13]成年雌性比雄性大,生殖部分也非常大。鲑疮痂鱼虱会生两串共500-1000颗卵,成熟后卵会变深色。一只雌虱一生可以产6-11对卵串,估计寿命只有7个月。[7][9][14]

生命周期

海虱有浮游及寄生的阶段。所有阶段之间都会脱壳。[12][13][15][16]其中鲑疮痂鱼虱只要17-72日就可以由成长到成年。

海虱的卵会孵化出第一期无节幼体,继而脱壳成长至第二期无节幼体。两期的无节幼体阶段都不会进食,只靠卵黄供应能量,而且懂得游泳。到了成年阶段就会寻找寄主。水流、盐度、光线等都可以帮助它们寻找寄主。[7]海虱多会寄生在鱼类身体受水流影响较少的地方,特别是鳍及其他受保护的地方。[6][17]当幼体附在寄主身上后,就会觅食一段时间脱壳成第一期的附着幼体。海虱会继续发育成另外的3期附着幼体。所有四个附着幼体阶段都有一个前部丝体的结构,用来附着在寄主身上。不同的海虱有不同的繁殖时间及方法。亚成体及成体可以在鱼类的身体上移动,有时甚至可以在寄主之间转移。雌性成体的身体较大,也较为扁平。[18]

食性

无节幼体及幼体期都是不会进食的。当附在寄主身上时,它们就会发育到附着幼体阶段。附着幼体都有胃肠道,且会吃寄主的黏液及组织。亚成体及成体都是主动觅食者,有时甚至会吸食寄主的血液鲑疮痂鱼虱会分泌大量胰蛋白酶到寄主的黏液,帮助觅食及消化[3][19]其他的物质包括前列腺素E2,可以帮助觅食及避开寄主的免疫反应。[3][20]海虱是否疾病媒介则不明,但它们肯定带有细菌病毒[21]

疾病

病理学

 
大西洋鲑上的雌性鲑疮痂鱼虱

海虱会在附着的位置对寄主造成物质及酶促的破坏。它们在吃食的时候会造成像割开的伤口,严重程度会受到几个因素所影响,如寄主物种、年龄及健康。海虱感染会对寄主鱼造成慢性压力,包括破坏其表皮出现失血、电解质的改变及释放皮质醇。这样会降低寄主鱼的免疫反应,令它们易于染病。[22][23]

不同海虱的物种、发育阶段及数量也会造成不同程度的破坏。大西洋鲑的组织似乎对海虱没有任何反应。相反,银大麻哈鱼驼背大马哈鱼对于鲑疮痂鱼虱则有强烈的反应,包括表皮增生炎症。在感染后的第一周就会排斥海虱。[3]鲑疮痂鱼虱可以令饲养的大西洋鲑及野生的红大马哈鱼出现深深的损伤,尤其是在头部,严重的程度可以见到头颅骨

饲养与野生鱼类的关系

有指受到海虱感染的饲养鲑鱼会将病传播到周边野生的幼鱼,令它们的数量大减。[24]海虱,尤其是鲑疮痂鱼虱及多种鱼虱属都会令饲养及野生的鲑鱼致命。[25][26]海虱会迁徙及附在野生鲑鱼的皮肤上达几日之久。[27][28][29]大量受感染的鲑鱼可以令海虱的浓度攀升,游经此地的野生幼鲑鱼就会受到感染及死亡。[30][31]加拿大太平洋海岸,差不多有80%的驼背大马哈鱼死于海虱的感染。[24]

褐鳟的数量因感染海虱已于近年出现大幅下降。[32]

多项研究都指饲养鲑鱼会积聚海虱,并会破坏周边的野生鲑鱼群落。[31]其他研究却指饲养鲑鱼所感染的海虱,并不会对野生群落造成影响。[33]

下属分类

鱼虱科包括以下属:

参考文献

  1. ^ Caligidae. ITIS. 
  2. ^ 2.0 2.1 T. Chad Walter & Geoff Boxshall. Caligidae. World Copepoda database. World Register of Marine Species. 2009 [2010-01-12]. (原始内容存档于2019-10-17). 
  3. ^ 3.0 3.1 3.2 3.3 G. N. Wagner, M. D. Fast & S. C. Johnson. Physiology and immunology of Lepeophtheirus salmonis infections of salmonids. Trends in Parasitology. 2008, 24: 176–83. doi:10.1016/j.pt.2007.12.010. 
  4. ^ S. R. M. Jones, G. Prosperi-Porta, E. Kim, P. Callow & N. B. Hargreaves. The occurrence of Lepeophtheirus salmonis and Caligus clemensi (Copepoda: Caligidae) on threespine stickleback Gasterosteus aculeatus in coastal British Columbia. Journal of Parasitology. 2006, 92 (3): 473–80. PMID 16883988. doi:10.1645/GE-685R1.1. 
  5. ^ M. Costelloe, J. Costelloe, G. O’Donohoe, N. J. Coghlan, M. Oonk & Y. van der Heijden. Planktonic distribution of sea lice larvae Lepeophtheirus salmonis, in Killary harbor, west coast of Ireland (PDF). Journal of the Marine Biological Association of the United Kingdom. 1998, 78: 853–74. doi:10.1017/S0025315400044830. 
  6. ^ 6.0 6.1 R. L. Genna, W. Mordue, A. W. Pike & A. J. Mordue-Luntz. Light intensity, salinity, and host velocity influence presettlement intensity and distribution on hosts by copepodids of sea lice, Lepeophtheirus salmonis. Canadian Journal of Fisheries and Aquatic Sciences. 2005, 62: 2675–82. doi:10.1139/f05-163. 
  7. ^ 7.0 7.1 7.2 7.3 M. J. Costello. Ecology of sea lice parasitic on farmed and wild fish (PDF). Trends in Parasitology. 2006, 22: 475–83. doi:10.1016/j.pt.2006.08.006. 
  8. ^ Kenneth M. Brooks. The effects of water temperature, salinity, and currents on the survival and distribution of the infective copepodid stage of sea lice (Lepeophtheirus salmonis) originating on Atlantic salmon farms in the Broughton Archipelago of British Columbia, Canada. Reviews in Fisheries Science. 2005, 13: 177–204. doi:10.1080/10641260500207109. 
  9. ^ 9.0 9.1 P. A. Heuch, A. Parsons & K. Boxaspen. Diel vertical migration: a possible host finding mechanism in salmon lice (Lepeophtheirus salmonis) copepodid?. Canadian Journal of Fisheries and Aquatic Sciences. 1995, 52: 681–9. doi:10.1139/f95-069. 
  10. ^ M. A. McKibben & D. W. Hay. Distributions of planktonic sea lice larvae Lepeophtheirus salmonis in the inter-tidal zone in Loch Torrindon, western Scotland in relation to salmon farm production cycles. Aquaculture and Research. 2004, 35: 742–50. doi:10.1111/j.1365-2109.2004.01096.x. 
  11. ^ M. J. Costello. How sea lice from salmon farms may cause wild salmonid declines in Europe and North America and be a threat to fishes elsewhere. Proceedings of the Royal Society B. 2009, 276: 3385–94. doi:10.1098/rspb.2009.0771. 
  12. ^ 12.0 12.1 S. C. Johnson & L. J. Albright. The developmental stages of Lepeophtheirus salmonis (Krøyer, 1837) (Copepoda: Caligidae). Canadian Journal of Zoology. 1991, 69 (4): 929–50. doi:10.1139/z91-138. 
  13. ^ 13.0 13.1 M. Anstensrud. Moulting and mating in Lepeophtheirus pectoralis (Copepoda: Caligidae). Journal of the Marine Biological Association of the United Kingdom. 1990, 70: 269–81. doi:10.1017/S0025315400035396. 
  14. ^ A. Mustafa, G. A. Conboy & J. F. Burka. Life-span and reproductive capacity of sea lice, Lepeophtheirus salmonis, under laboratory conditions. Aquacul. Assoc. Canada Spec. Publ. 2001, 4: 113–4. 
  15. ^ Thomas A. Schram. Supplementary description of the developmental stages of Lepeophtheirus salmonis (Krøyer, 1837) (Copepoda: Caligidae). G. A. Boxshall & D. Defaye (编). Pathogens of Wild and Farmed Fish: Sea Lice. Chichester: Ellis Horwood. 1993: 30–50 [2010-12-23]. ISBN 0-13-015504-7. (原始内容存档于2012-11-12). 
  16. ^ A. W. Pike & S. L. Wadsworth. Sealice on salmonids: their biology and control. Advances in Parasitology. 1999, 44: 233–337. PMID 10563397. doi:10.1016/S0065-308X(08)60233-X. 
  17. ^ J. E. Bron, C. Sommerville, M. Jones & G. H. Rae. The settlement and attachment of early stages of the salmon louse, Lepeophtheirus salmonis (Copepoda: Caligidae) on the salmon host Salmo salar. Journal of Zoology. 1991, 224 (2): 201–12. doi:10.1111/j.1469-7998.1991.tb04799.x. 
  18. ^ C. D. Todd, A. M. Walker, J. E. Hoyle, S. J. Northcott, A. F. Walker & M. G. Ritchie. Infestations of wild adult Atlantic salmon (Salmo salar L.) by the ectoparasitic copepod sea louse Lepeophtheirus salmonis (Krøyer): prevalence, intensity and the spatial distribution of males and females on the host fish. Hydrobiologia. 2000, 429 (2–3): 181–96. doi:10.1023/A:1004031318505. 
  19. ^ Kara J. Firth, Stewart C. Johnson & Neil W. Ross. Characterization of proteases in the skin mucus of Atlantic salmon (Salmo salar) infected with the salmon louse (Lepeophtheirus salmonis) and in whole-body louse homogenates. Journal of Parasitology. 2000, 86 (6): 1199–205. PMID 11191891. doi:10.1645/0022-3395(2000)086[1199:COPITS]2.0.CO;2. 
  20. ^ M. D. Fast, N. W. Ross & S. C. Johnson. Prostaglandin E2 modulation of gene expression in an Atlantic salmon (Salmo salar) macrophage-like cell line (SHK-1). Developmental & Comparative Immunology. 2005, 29 (11): 951–63. PMID 15936074. doi:10.1016/j.dci.2005.03.007. 
  21. ^ A. Nylund, B. Bjørknes & C. Wallace. Lepeophtheirus salmonis – a possible vector in the spread of diseases on salmonids. Bull. Europ. Assoc. Fish Pathol. 1991, 11: 213–6. 
  22. ^ S. C. Johnson & L. J. Albright. Effects of cortisol implants on the susceptibility and the histopathology of the responses of naive coho salmo Oncorhynchus kisutch to experimental infection with Lepeophtheirus salmonis (Copepoda : Caligidae). Disease of Aquatic Organisms. 1992, 14: 195–205. doi:10.3354/dao014195. 
  23. ^ N. W. Ross, K. J. Firth, A. Wang, J. F. Burka & S. C. Johnson. Changes in hydrolytic enzyme activities of naive Atlantic salmon (Salmo salar) skin mucus due to infection with the salmon louse (Lepeophtheirus salmonis) and cortisol implantation. Disease of Aquatic Organisms. 2000, 41 (1): 43–51. PMID 10907138. doi:10.3354/dao041043. 
  24. ^ 24.0 24.1 M. Krkosek, J. S. Ford, A. Morton, S. Lele, R. A. Myers & M. A. Lewis. Declining wild salmon populations in relation to parasites from farm salmon. Science. 2007, 318 (5857): 1772. PMID 18079401. doi:10.1126/science.1148744. 
  25. ^ S. Bravo. Sea lice in Chilean salmon farms. Bull. Eur. Ass. Fish Pathol. 2003, 23: 197–200. 
  26. ^ Sea lice and salmon: elevating the dialogue on the farmed-wild salmon story (PDF). Watershed Watch Salmon Society. 2004 [2010-01-15]. (原始内容 (PDF)存档于2010-12-14). 
  27. ^ A. Morton, R. Routledge, C. Peet & A. Ladwig. Sea lice (Lepeophtheirus salmonis) infection rates on juvenile pink (Oncorhynchus gorbuscha) and chum (Oncorhynchus keta) salmon in the nearshore marine environment of British Columbia, Canada. Canadian Journal of Fisheries and Aquatic Sciences. 2004, 61 (2): 147–57. doi:10.1139/f04-016. 
  28. ^ Corey Ryan Peet. Interactions between sea lice (Lepeoptheirus salmonis and Caligus clemensii), juvenile salmon (Oncorhynchus keta and Oncorhynchus gorbuscha) and salmon farms in British Columbia (M.Sc.论文). University of Victoria. 2007. 
  29. ^ M. Krkošek, A. Gottesfeld, B. Proctor, D. Rolston, C. Carr-Harris & M. A. Lewis. Effects of host migration, diversity, and aquaculture on disease threats to wild fish populations. Proceedings of the Royal Society B. 2007, 274 (1629): 3141–9. PMC 2293942 . PMID 17939989. doi:10.1098/rspb.2007.1122. 
  30. ^ A. Morton, R. Routledge & M. Krkošek. Sea louse infestation in wild juvenile salmon and Pacific herring associated with fish farms off the east-central coast of Vancouver Island, British Columbia (PDF). North American Journal of Fisheries Management. 2008, 28: 523–32. doi:10.1577/M07-042.1. [永久失效链接]
  31. ^ 31.0 31.1 M. Krkošek, M. A. Lewis, A. Morton, L. N. Frazer & J. P. Volpe. Epizootics of wild fish induced by farm fish. Proceedings of the National Academy of Sciences. 2006, 103 (42): 15506–10. PMC 1591297 . PMID 17021017. doi:10.1073/pnas.0603525103. 
  32. ^ Charles Clover. The End of the Line: How Overfishing is Changing the World and What We Eat. London: Ebury Press. 2004. ISBN 0-09-189780-7. 
  33. ^ Peter Andreas Heuch, Pål Arne Bjørn, Bengt Finstad, Jens Christian Holst, Lars Asplin & Frank Nilsen. A review of the Norwegian ‘National Action Plan Against Salmon Lice on Salmonids’: The effect on wild salmonids. Aquaculture. 2005, 246 (1–4): 79–92. doi:10.1016/j.aquaculture.2004.12.027. 
  34. ^ Z. Kabata (1964) Copepoda parasitic on Australian fishes. I. Hermilius youngi sp. nov. (Caligidae), Annals and Magazine of Natural History, 7:82, 609-618, DOI: 10.1080/00222936408651504
  35. ^ Z. Kabata (1964) Copepoda parasitic on Australian fishes. II. Mappates alter sp. nov. (Caligidae), Annals and Magazine of Natural History, 7:82, 641-649, DOI: 10.1080/00222936408651509

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