海洋无脊椎动物补体系统研究进展
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摘要
补体系统是先天免疫的重要组成部分,在免疫反应中发挥重要作用。补体的研究可以为海洋无脊椎动物提供病害防治的理论基础。本文介绍了海洋无脊椎动物补体系统中已报道的C1q、MBL、ficolin、MASP、B因子、C3和C6等成分,比较了海洋无脊椎动物补体成分的功能,描述了其补体激活途径和补体系统进化情况。目前,海洋无脊椎动物补体的研究发展很快,揭示可能存在与脊椎动物差异较大的补体系统,但是缺乏系统的基础研究而未能构建起理论模型。
The complement system is an important part of innate immunity and plays a vital role in immune responses. This study can provide theoretical basis for disease control of the economic farming of marine invertebrates. This article described the ingredients including C1 q,MBL,ficolin,MASP,B factor,C3,and C6 etc.,which had been found in the complement system of marine invertebrates. The inherent component of the complement system in marine invertebrates is a major study subject requiring deep research. Though our understanding of complement receptor,active fragment and binding protein is growing,it is still limited in mammalian vertebrates. The complement component of marine invertebrates is less than that in the vertebrates,there are different subtypes in terms of the amount in a single component,in most case,the number of subtypes is large. Different individuals vary greatly in the complement components and they have different expression in differet periods of developmental stage. We compared the function of complement component in marine invertebrate,finding that each complement component was presented in a variety of subtypes and there were differences between the functions of various subtypes. We also described some of the primary actors and pathways of the complement system in marine invertebrates. Lectin pathway and alternative pathway potentially existed in marine invertebrates,but the classic pathway and terminal pathway were not found yet. After comparison of the vertebrate and invertebrate complement pathway,we found that marine invertebrates may have their special features in terms of these pathways. The research on complement pathway in marine invertebrates is relatively limited. No systematic theoretical model has been developed to fully display the development of marine invertebrate complement pathway. This article also described the evolution of marine invertebrate complement system and compared with vertebrate the invertebrate features of evolution of complement component obviously. In the pathway there was not only the trace of evolution but also the existence of significant differences. At present,the state of research of complement system in invertebrates is primary. The development of marine invertebrates complement is fast,but short of basic research,failing to establish a large-scale system of theoretical models. With more finished full sequence of gene from different species,and the use of technology of height sequence in study,gradually,complement gene will be found,and the function and pathway will be improved in marine invertebrates. The exploratory study of complement system in marine invertebrate will offer a critical framework for better understanding of immune system and stress resistance in the growth of marine invertebrates.
The complement system is an important part of innate immunity and plays a vital role in immune responses. This study can provide theoretical basis for disease control of the economic farming of marine invertebrates. This article described the ingredients including C1 q,MBL,ficolin,MASP,B factor,C3,and C6 etc.,which had been found in the complement system of marine invertebrates. The inherent component of the complement system in marine invertebrates is a major study subject requiring deep research. Though our understanding of complement receptor,active fragment and binding protein is growing,it is still limited in mammalian vertebrates. The complement component of marine invertebrates is less than that in the vertebrates,there are different subtypes in terms of the amount in a single component,in most case,the number of subtypes is large. Different individuals vary greatly in the complement components and they have different expression in differet periods of developmental stage. We compared the function of complement component in marine invertebrate,finding that each complement component was presented in a variety of subtypes and there were differences between the functions of various subtypes. We also described some of the primary actors and pathways of the complement system in marine invertebrates. Lectin pathway and alternative pathway potentially existed in marine invertebrates,but the classic pathway and terminal pathway were not found yet. After comparison of the vertebrate and invertebrate complement pathway,we found that marine invertebrates may have their special features in terms of these pathways. The research on complement pathway in marine invertebrates is relatively limited. No systematic theoretical model has been developed to fully display the development of marine invertebrate complement pathway. This article also described the evolution of marine invertebrate complement system and compared with vertebrate the invertebrate features of evolution of complement component obviously. In the pathway there was not only the trace of evolution but also the existence of significant differences. At present,the state of research of complement system in invertebrates is primary. The development of marine invertebrates complement is fast,but short of basic research,failing to establish a large-scale system of theoretical models. With more finished full sequence of gene from different species,and the use of technology of height sequence in study,gradually,complement gene will be found,and the function and pathway will be improved in marine invertebrates. The exploratory study of complement system in marine invertebrate will offer a critical framework for better understanding of immune system and stress resistance in the growth of marine invertebrates.
引文
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[2]晏沐阳,宋宏彬,贾雷立,等.补体学[M].北京:科学出版,2009:1-7.YAN M Y,SONG H B,JIA L L,et al.Complementology[M].Beijing:Sience Press,2009:1-7.
[3]章跃陵,赵贤亮,严芳.无脊椎动物免疫分子的多态性[J].中国生物化学与分子生物学报,2010,26(3):209-214.ZHANG Y L,ZHAO X L,YAN F.Polymorphisms of immune molecules in invertebrates[J].Chinese Journal of Biochemistry and Molecular Biology,2010,26(3):209-214.
[4]SONG W C,SARRIAS M R,LAMBRIS J D.Complement and innate immunity[J].Immunopharmacology,2000,49(1):187-198.
[5]NONAKA M,KIMURA A.Genomic view of the evolution of the complement system[J].Immunogenetics,2006,58(9):701-713.
[6]ABBAS A K,LICHTMAN A H,PILLAI S.Cellular and Molecular Immunology[M].New York:Elsevier Medicine,2011.
[7]IWANAGA S,LEE B L.Recent advances in the innate immunity of invertebrate animals[J].BMB Reports,2005,38(2):128-150.
[8]HIBINO T,NONAKA M.A novel third complement component C3 gene of Ciona intestinalis expressed in the endoderm at the early developmental stages[J].Isj-Invertebrate Survival Journal,2013,10(1):29-37.
[9]AZUMI K,DE S R,DE T A,et al.Genomic analysis of immunity in a urochordate and the emergence of the vertebrate immune system:“Waiting for Godot”[J].Immunogenetics,2003,55(8):570-581.
[10]MARINO R,KIMURA Y,DE S R,et al.Complement in urochordates:Cloning and characterization of two C3-like genes in the ascidian Ciona intestinalis[J].Immunogenetics,2002,53(12):1055-1064.
[11]SODERGREN E,WEINSTOCK G M,DAVIDSON E H,et al.The genome of the sea urchin Strongylocentrotus purpuratus[J].Science,2006,314(5801):941-952.
[12]SIMAKOV O,MARLETAZ F,CHO S J,et al.Insights into bilaterian evolution from three spiralian genomes[J].Nature,2013,493(7433):526-531.
[13]LOWE T M,EDDY S R.tRNAscan-SE:A program for improved detection of transfer RNA genes in genomic sequence[J].Nucleic Acids Research,1997,25(5):955-964.
[14]WANG L L,WANG L L,ZHANG H,et al.A C1q domain containing protein from scallop Chlamys farreri serving as pattern recognition receptor with heat-aggregated Ig G binding activity[J].Plos One,2012,7(8):e43289.
[15]WANG L L,WANG L L,KONG P F,et al.A novel C1q DC protein acting as pattern recognition receptor in scallop Argopecten irradians[J].Fish&Shellfish Immunology,2012,33(2):427-435.
[16]GERDOL M,MANFRIN C,DE M G,et al.The C1q domain containing proteins of the Mediterranean mussel Mytilus galloprovincialis:A widespread and diverse family of immune-related molecules[J].Developmental&Comparative Immunology,2011,35(6):635-643.
[17]隋立军,刘卫东,李云峰,等.文蛤C1q基因的克隆与表达及其重组蛋白活性的研究[J].水产科学,2012,31(6):321-328.SUI L J,LIU W D,LI Y F,et al.Molecular cloning,mRNA expression and bioassay of recombinant protein of C1q gene from hard clam(Meretrix meretrix)[J].Fisheries Science,2012,31(6):321-328.
[18]XU T,XIE J S,LI J M,et al.Identification of expressed genes in c DNA library of hemocytes from the RLO-challenged oyster,Crassostrea ariakensis Gould with special functional implication of three complement-related fragments(Ca C1q1,Ca C1q2and Ca C3)[J].Fish&Shellfish Immunology,2012,32(6):1106-1116.
[19]SRIVASTAVA M,SIMAKOV O,CHAPMAN J,et al.The Amphimedon queenslandica genome and the evolution of animal complexity[J].Nature,2010,466(7307):720-726.
[20]CARLAND T M,GERWICK L.The C1q domain containing proteins:Where do they come from and what do they do?[J].Developmental&Comparative Immunology,2010,34(8):785-790.
[21]JIANG S,LI H,ZHANG D X,et al.A C1q domain containing protein from Crassostrea gigas serves as pattern recognition receptor and opsonin with high binding affinity to LPS[J].Fish&Shellfish Immunology,2015.
[22]KONG P F,ZHANG H,WANG L L,et al.Ai C1q DC-1,a novel g C1q-domain-containing protein from bay scallop Argopecten irradians with fungi agglutinating activity[J].Developmental&Comparative Immunology,2010,34(8):837-846.
[23]HUANG S F,YUAN S C,GUO L,et al.Genomic analysis of the immune gene repertoire of amphioxus reveals extraordinary innate complexity and diversity[J].Genome Research,2008,18(7):1112-1126.
[24]SKJOEDT M O,PALARASAH Y,RASMUSSEN K,et al.Two mannose-binding lectin homologues and an MBL-associated serine protease are expressed in the gut epithelia of the urochordate species Ciona intestinalis[J].Developmental&Comparative Immunology,2010,34(1):59-68.
[25]HUANG H Q,HUANG S F,YU Y C,et al.Functional characterization of a ficolin-mediated complement pathway in amphioxus[J].Journal of Biological Chemistry,2011,286(42):36739-36748.
[26]KENJO A,TAKAHASHI M,MATSUSHITA M,et al.Cloning and characterization of novel ficolins from the solitary ascidian,Halocynthia roretzi[J].Journal of Biological Chemistry,2001,276(23):19959-19965.
[27]ZHANG G F,FANG X D,GUO X M,et al.The oyster genome reveals stress adaptation and complexity of shell formation[J].Nature,2012,490(7418):49-54.
[28]SEKINE H,KENJO A,AZUMI K,et al.An ancient lectin-dependent complement system in an ascidian:Novel lectin isolated from the plasma of the solitary ascidian,Halocynthia roretzi[J].The Journal of Immunology,2001,167(8):4504-4510.
[29]BONURA A,VIZZINI A,SALERNO G,et al.Isolation and expression of a novel MBL-like collectin c DNA enhanced by LPS injection in the body wall of the ascidian Ciona intestinalis[J].Molecular Immunology,2009,46(11):2389-2394.
[30]FAN C X,ZHANG S C,LI L,et al.Fibrinogenrelated protein from amphioxus Branchiostoma belcheri is a multivalent pattern recognition receptor with a bacteriolytic activity[J].Molecular immunology,2008,45(12):3338-3346.
[31]JI X,AZUMI K,SASAKI M,et al.Ancient origin of the complement lectin pathway revealed by molecular cloning of mannan binding proteinassociated serine protease from a urochordate,the Japanese ascidian,Halocynthia roretzi[J].Proceedings of the National Academy of Sciences,1997,94(12):6340-6345.
[32]KIMURA A,SAKAGUCHI E,NONAKA M.Multicomponent complement system of Cnidaria:C3,Bf,and MASP genes expressed in the endodermal tissues of a sea anemone,Nematostella vectensis[J].Immunobiology,2009,214(3):165-178.
[33]HE Y A,TANG B,ZHANG S C,et al.Molecular and immunochemical demonstration of a novel member of Bf/C2 homolog in amphioxus Branchiostoma belcheri:Implications for involvement of hepatic cecum in acute phase response[J].Fish&Shellfish Immunology,2008,24(6):768-778.
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