鳜早期色素发育和色彩图案的形成
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摘要
色彩图案作为最显著的外部形态特征之一,在动物生存与交流中起着重要的作用。为了解鳜(Siniperca chautsi)早期发育过程中这一形态特征变化,采用CCD-Adapter解剖镜对鳜早期(胚胎期至出膜40日龄)体表色素细胞种类与分布、主要图案(条、带、斑)的形成过程进行了观察,同时对不同部位皮肤进行组织切片观察。结果显示,胚胎期,最早观察到黑色素细胞,位于卵黄囊和油球,出膜前,头部出现黄色素细胞;出膜后,黑色素细胞发育最为显著,红色素细胞出现在眼球后部和躯干前部;5日龄后,黄色素细胞发育增加,鱼体各部位均有分布,黑色素细胞继续发育,图案形成开始。鳜早期色彩图案形成过程:(1)躯干纵带:5日龄,背部出现少量黑色素,14日龄,背鳍基部黑色素与腹部黑色素相连;(2)头部过眼条带:10日龄,鳃盖后上方黑色素明显增多,12日龄,眼球后部经鳃盖后上缘至背部前端的条带形成,17日龄,上颌至眼球前部的条带形成;(3)头顶条带:6日龄,头顶正上方黑色斑点增多,18日龄,头部上方黑色斑块分别向前、后延伸,23日龄,头顶正上方黑色条带基本形成;(4)躯干斑块:8日龄,尾部底端出现一个较小的黑色斑块,15日龄,尾柄前部出现3个不规则黑色斑块,25日龄,躯干中后部5个近圆形黑色斑块形成。结果表明,鳜胚胎期至出膜40日龄,体表出现黑色素细胞、黄色素细胞和红色素细胞,体色以黑色为主,主要条带或斑块在仔鱼5日龄后按不同方式逐渐形成,不同皮肤部位的色素层组成与分布方式不同。
As one of the most prominent external morphological features,color patterns play an importantrole in animal survival and communication.In order to understand morphological changes in the early development of Siniperca chautsi,CCD-Adapter anatomical microscope was used to observe the formation,distribution and main patterns(stripe,band and spot) of the pigment cells in the early stage,from embryo stage up to 40 days after hatching.At the same time,the skin was observed after tissue section.The results show that melanophores are first observed in the yolk sac and oil balls at embryonic stage,xanthophores appear on the head at the pre-hatching stage.After hatching,melanophores develop most prominently and erythrophores appear behind the eye and in the front of trunk(Fig.1).After 5 days of age,xanthophores increase throughout the whole body and melanophores continue to develop and pattern formation begins(Fig.2).Early color pattern formation processes of S.chautsi include:1)Trunk band:some melanin on the back at 5 days old,the dorsal melanin and the abdominal melanin are connected at 14 days old(Fig.4 a﹣c).2) Eye strip:the melanin above the gill cover increases at 10 days old,the hind strip part forms from the upper edge of gill cover to the front of the back at 12 days old,the front strip part forms from the upper jaw to the front of the eye at 17 days old(Fig.4 d﹣f).3) Overhead strip:the black spots above the top of the head increase at 6 days old,these spots extend forward and backward at 18 days old.The black strip just above the top of the head forms at 23 days old(Fig.4 g﹣i).4) Trunk spots:a small black spot appears at the bottom of the tail at 8 days old,three irregular black patches appear in the front of the tail at 15 days old.Five round black spots formed in the back of the torso at 25 days old(Fig.4 j﹣l).The results show that melanophores,xanthophores and erythrophores appear at S.chautsi′s body surface in early stage,the whole body is dominated by black color,the main pigment strips and spots gradually form in different modes at the later larval stages,and pigment layer composition and distribution are different in different skin parts.
As one of the most prominent external morphological features,color patterns play an importantrole in animal survival and communication.In order to understand morphological changes in the early development of Siniperca chautsi,CCD-Adapter anatomical microscope was used to observe the formation,distribution and main patterns(stripe,band and spot) of the pigment cells in the early stage,from embryo stage up to 40 days after hatching.At the same time,the skin was observed after tissue section.The results show that melanophores are first observed in the yolk sac and oil balls at embryonic stage,xanthophores appear on the head at the pre-hatching stage.After hatching,melanophores develop most prominently and erythrophores appear behind the eye and in the front of trunk(Fig.1).After 5 days of age,xanthophores increase throughout the whole body and melanophores continue to develop and pattern formation begins(Fig.2).Early color pattern formation processes of S.chautsi include:1)Trunk band:some melanin on the back at 5 days old,the dorsal melanin and the abdominal melanin are connected at 14 days old(Fig.4 a﹣c).2) Eye strip:the melanin above the gill cover increases at 10 days old,the hind strip part forms from the upper edge of gill cover to the front of the back at 12 days old,the front strip part forms from the upper jaw to the front of the eye at 17 days old(Fig.4 d﹣f).3) Overhead strip:the black spots above the top of the head increase at 6 days old,these spots extend forward and backward at 18 days old.The black strip just above the top of the head forms at 23 days old(Fig.4 g﹣i).4) Trunk spots:a small black spot appears at the bottom of the tail at 8 days old,three irregular black patches appear in the front of the tail at 15 days old.Five round black spots formed in the back of the torso at 25 days old(Fig.4 j﹣l).The results show that melanophores,xanthophores and erythrophores appear at S.chautsi′s body surface in early stage,the whole body is dominated by black color,the main pigment strips and spots gradually form in different modes at the later larval stages,and pigment layer composition and distribution are different in different skin parts.
引文
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Ceinos R M,Raúl G,Kelsh R N,et al.2015.Pigment patterns in adult fish result from superimposition of two largely independent pigmentation mechanisms.Pigment Cell Melanoma Research,28(2):196-209.
Georg F H,Jana K,Hans-Martin M,et al.2013.Iridophores and their interactions with other chromatophores are required for stripe formation in zebrafish.Development,140(14):2997-3007.
Hirata M,Nakamura K,Kanemaru T,et al.2003.Pigment cell organization in the hypodermis of zebrafish.Developmental Dynamics,227(4):497-503.
Hirata M,Nakamura K,Kanemaru T,et al.2010.Pigment cell organization in the hypodermis of zebrafish.Developmental Dynamics,227(4):497-503.
Hirata M,Nakamura K,Kondo S.2005.Pigment cell distributions in different tissues of the zebrafish,with special reference to the striped pigment pattern.Developmental Dynamics,234(2):293-300.
Lopes S S,Yang X,Müller J,et al.2008.Leukocyte tyrosine kinase functions in pigment cell development.PLoS Genetics,4(3):e1000026.
Nakamasu A,Takahashi G,Kanbe A,et al.2009.Interactions between zebrafish pigment cells responsible for the generation of Turing patterns.Proceedings of the National Academy of Sciences of the United States of America,106(21):8429-8434.
Parichy D M.2006.Evolution of danio pigment pattern development.Heredity,97(3):200-210.
Ramachandran V S,Tyler C W,Gregory R L,et al.1996.Rapid adaptive camouflage in tropical flounders.Nature,379(6568):815-818.
Reiss P,Able K W,Nunes M S,et al.2012.Color pattern variation in Cichla temensis(Perciformes:Cichlidae):resolution based on morphological,molecular,and reproductive data.Neotropical Ichthyology,10(1):59-70.
Santos M E,Braasch I,Boileau N,et al.2014.The evolution of cichlid fish egg-spots is linked with a cis-regulatory change.Nature Communications,5(1):5149|DOI:10.1038/ncomms6149.
Sazima I C,Lucélia N M,et al.2006.Fallen leaves on the water-bed:diurnal camouflage of three night active fish species in an Amazonian streamlet.Neotropical Ichthyology,4(1):119-122.
Sugimoto M.2002.Morphological color changes in fish:regulation of pigment cell density and morphology.Microscopy Research&Technique,58(6):496-503.
蒋一珪.1959.梁子湖鳜鱼的生物学.水生生物集刊,3:375-384.
刘友亮,崔希群,陈敬存.1987.鳜鱼早期发育的生态形态学特征.水利渔业,(4):41-45.
苗田田,钟有奇,钱克林.等.2013.金鳜体色发生观察.水产科技情报,40(2):57-59.
赵丽丽,赵金良.2008.中国少鳞鳜不同地理群体遗传变异的AFLP分析.农业生物技术学报,16(5):907-908.
周才武,杨青,蔡德霖.1988.鳜亚科SINIPERCINAE鱼类的分类整理和地理分布.动物学研究,9(2):113-125.
Ceinos R M,Raúl G,Kelsh R N,et al.2015.Pigment patterns in adult fish result from superimposition of two largely independent pigmentation mechanisms.Pigment Cell Melanoma Research,28(2):196-209.
Georg F H,Jana K,Hans-Martin M,et al.2013.Iridophores and their interactions with other chromatophores are required for stripe formation in zebrafish.Development,140(14):2997-3007.
Hirata M,Nakamura K,Kanemaru T,et al.2003.Pigment cell organization in the hypodermis of zebrafish.Developmental Dynamics,227(4):497-503.
Hirata M,Nakamura K,Kanemaru T,et al.2010.Pigment cell organization in the hypodermis of zebrafish.Developmental Dynamics,227(4):497-503.
Hirata M,Nakamura K,Kondo S.2005.Pigment cell distributions in different tissues of the zebrafish,with special reference to the striped pigment pattern.Developmental Dynamics,234(2):293-300.
Lopes S S,Yang X,Müller J,et al.2008.Leukocyte tyrosine kinase functions in pigment cell development.PLoS Genetics,4(3):e1000026.
Nakamasu A,Takahashi G,Kanbe A,et al.2009.Interactions between zebrafish pigment cells responsible for the generation of Turing patterns.Proceedings of the National Academy of Sciences of the United States of America,106(21):8429-8434.
Parichy D M.2006.Evolution of danio pigment pattern development.Heredity,97(3):200-210.
Ramachandran V S,Tyler C W,Gregory R L,et al.1996.Rapid adaptive camouflage in tropical flounders.Nature,379(6568):815-818.
Reiss P,Able K W,Nunes M S,et al.2012.Color pattern variation in Cichla temensis(Perciformes:Cichlidae):resolution based on morphological,molecular,and reproductive data.Neotropical Ichthyology,10(1):59-70.
Santos M E,Braasch I,Boileau N,et al.2014.The evolution of cichlid fish egg-spots is linked with a cis-regulatory change.Nature Communications,5(1):5149|DOI:10.1038/ncomms6149.
Sazima I C,Lucélia N M,et al.2006.Fallen leaves on the water-bed:diurnal camouflage of three night active fish species in an Amazonian streamlet.Neotropical Ichthyology,4(1):119-122.
Sugimoto M.2002.Morphological color changes in fish:regulation of pigment cell density and morphology.Microscopy Research&Technique,58(6):496-503.
蒋一珪.1959.梁子湖鳜鱼的生物学.水生生物集刊,3:375-384.
刘友亮,崔希群,陈敬存.1987.鳜鱼早期发育的生态形态学特征.水利渔业,(4):41-45.
苗田田,钟有奇,钱克林.等.2013.金鳜体色发生观察.水产科技情报,40(2):57-59.
赵丽丽,赵金良.2008.中国少鳞鳜不同地理群体遗传变异的AFLP分析.农业生物技术学报,16(5):907-908.
周才武,杨青,蔡德霖.1988.鳜亚科SINIPERCINAE鱼类的分类整理和地理分布.动物学研究,9(2):113-125.
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