【《对虾免疫机制综述》5900字】

对虾免疫机制综述目录TOC\o"1-3"\h\u26099对虾免疫机制综述 161841.1免疫识别 1207501.2体液免疫 2163821.3细胞免疫 6作为无脊椎动物，对虾通常缺乏淋巴细胞或基于抗体的适应性免疫，仅依靠先天免疫系统来抵御入侵的病原体。尽管如此，对虾已发展出非常复杂的针对病原体的先天免疫系统ADDINEN.CITE<EndNote><Cite><Author>Wang</Author><Year>2013</Year><RecNum>132</RecNum><DisplayText>[100]</DisplayText><record><rec-number>132</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613553056">132</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Wang,X.W.</author><author>Wang,J.X.</author></authors></contributors><auth-address>TheKeyLaboratoryofPlantCellEngineeringandGermplasmInnovationofMinistryofEducation,SchoolofLifeSciences,ShandongUniversity,Jinan,Shandong250100,China.</auth-address><titles><title>Patternrecognitionreceptorsactingininnateimmunesystemofshrimpagainstpathogeninfections</title><secondary-title>FishShellfishImmunol</secondary-title><alt-title>Fish&shellfishimmunology</alt-title></titles><periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></periodical><alt-periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></alt-periodical><pages>981-9</pages><volume>34</volume><number>4</number><edition>2012/09/11</edition><keywords><keyword>AminoAcidSequence</keyword><keyword>Animals</keyword><keyword>Aquaculture</keyword><keyword>Host-PathogenInteractions/immunology</keyword><keyword>Immunity,Innate/immunology</keyword><keyword>MolecularSequenceData</keyword><keyword>Penaeidae/*immunology</keyword><keyword>Receptors,PatternRecognition/*immunology</keyword><keyword>SignalTransduction/immunology</keyword></keywords><dates><year>2013</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>1050-4648</isbn><accession-num>22960101</accession-num><urls></urls><electronic-resource-num>10.1016/j.fsi.2012.08.008</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[100]。先天免疫是一种古老且保守的保护机制，它在后生动物的进化早期出现，分为体液免疫和细胞免疫ADDINEN.CITEADDINEN.CITE.DATA[101,102]。体液免疫和细胞免疫相互作用、相互依存、协同作用以保护机体并清除异物和病原体ADDINEN.CITE<EndNote><Cite><Author>Amparyup</Author><Year>2013</Year><RecNum>126</RecNum><DisplayText>[103]</DisplayText><record><rec-number>126</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613546354">126</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Amparyup,P.</author><author>Charoensapsri,W.</author><author>Tassanakajon,A.</author></authors></contributors><auth-address>CenterofExcellenceforMolecularBiologyandGenomicsofShrimp,DepartmentofBiochemistry,FacultyofScience,ChulalongkornUniversity,Bangkok10330,Thailand.piti.amp@biotec.or.th</auth-address><titles><title>Prophenoloxidasesystemanditsroleinshrimpimmuneresponsesagainstmajorpathogens</title><secondary-title>FishShellfishImmunol</secondary-title><alt-title>Fish&shellfishimmunology</alt-title></titles><periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></periodical><alt-periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></alt-periodical><pages>990-1001</pages><volume>34</volume><number>4</number><edition>2012/09/11</edition><keywords><keyword>Animals</keyword><keyword>Aquaculture</keyword><keyword>CatecholOxidase/*immunology</keyword><keyword>EnzymePrecursors/*immunology</keyword><keyword>Host-PathogenInteractions</keyword><keyword>Immunity,Innate/immunology</keyword><keyword>Melanins/immunology</keyword><keyword>Penaeidae/enzymology/*immunology</keyword><keyword>Receptors,PatternRecognition/immunology</keyword><keyword>SerineEndopeptidases/immunology</keyword></keywords><dates><year>2013</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>1050-4648</isbn><accession-num>22960099</accession-num><urls></urls><electronic-resource-num>10.1016/j.fsi.2012.08.019</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[103]。对虾的免疫反应主要发生在血淋巴中，血淋巴包含三种主要类型的血细胞，分别为透明、颗粒和半颗粒细胞ADDINEN.CITE<EndNote><Cite><Author>Martin</Author><Year>1985</Year><RecNum>129</RecNum><DisplayText>[104]</DisplayText><record><rec-number>129</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613547130">129</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Martin,G.G.</author><author>Graves,B.L.</author></authors></contributors><auth-address>DepartmentofBiology,OccidentalCollege,LosAngeles,California90041.</auth-address><titles><title>Finestructureandclassificationofshrimphemocytes</title><secondary-title>JMorphol</secondary-title><alt-title>Journalofmorphology</alt-title></titles><periodical><full-title>JMorphol</full-title><abbr-1>Journalofmorphology</abbr-1></periodical><alt-periodical><full-title>JMorphol</full-title><abbr-1>Journalofmorphology</abbr-1></alt-periodical><pages>339-348</pages><volume>185</volume><number>3</number><edition>1985/09/01</edition><dates><year>1985</year><pub-dates><date>Sep</date></pub-dates></dates><isbn>0022-2887</isbn><accession-num>29976016</accession-num><urls></urls><electronic-resource-num>10.1002/jmor.1051850306</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[104]。1.1免疫识别先天免疫系统根据入侵微生物保守的分子模式，即病原体相关分子模式（pathogen-associatedmolecularpatterns，PAMPs）来区分自我和非自我ADDINEN.CITEADDINEN.CITE.DATA[91]。当PAMPs被宿主细胞表面的模式识别受体（patternrecognitionreceptors，PRRs）识别并结合后，会触发一系列免疫反应，从而导致先天免疫系统的激活ADDINEN.CITEADDINEN.CITE.DATA[105]。PAMPs通常是微生物表面的多糖和糖蛋白，如革兰氏阳性菌的脂多糖（lipopolysaccharide，LPS），革兰氏阴性菌的肽聚糖（peptidoglycan，PGN）和磷壁酸（lipotechoicacid，LTA），以及真菌细胞的葡聚糖（glucans）。此外，PAMPs也可以是多核苷酸，如非甲基化CpGDNA，病毒的单链和双链RNA。不同的PAMPs由特定的PRRs识别。迄今为止，已在虾中鉴定出11种PRRs，它们在虾免疫反应中具有不同功能ADDINEN.CITE<EndNote><Cite><Author>Wang</Author><Year>2013</Year><RecNum>132</RecNum><DisplayText>[100]</DisplayText><record><rec-number>132</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613553056">132</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Wang,X.W.</author><author>Wang,J.X.</author></authors></contributors><auth-address>TheKeyLaboratoryofPlantCellEngineeringandGermplasmInnovationofMinistryofEducation,SchoolofLifeSciences,ShandongUniversity,Jinan,Shandong250100,China.</auth-address><titles><title>Patternrecognitionreceptorsactingininnateimmunesystemofshrimpagainstpathogeninfections</title><secondary-title>FishShellfishImmunol</secondary-title><alt-title>Fish&shellfishimmunology</alt-title></titles><periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></periodical><alt-periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></alt-periodical><pages>981-9</pages><volume>34</volume><number>4</number><edition>2012/09/11</edition><keywords><keyword>AminoAcidSequence</keyword><keyword>Animals</keyword><keyword>Aquaculture</keyword><keyword>Host-PathogenInteractions/immunology</keyword><keyword>Immunity,Innate/immunology</keyword><keyword>MolecularSequenceData</keyword><keyword>Penaeidae/*immunology</keyword><keyword>Receptors,PatternRecognition/*immunology</keyword><keyword>SignalTransduction/immunology</keyword></keywords><dates><year>2013</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>1050-4648</isbn><accession-num>22960101</accession-num><urls></urls><electronic-resource-num>10.1016/j.fsi.2012.08.008</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[100]。其中，Toll样受体（Tolllikereceptor，TLR）是第一个详细研究也是目前研究最为深入的PRRs家族。TLR是一类经典的跨膜糖蛋白，其胞外结构域包含多种富含亮氨酸重复（LRR）的基序，而胞质区具有与白细胞介素1受体（IL-1R）同源的TIR结构域ADDINEN.CITE<EndNote><Cite><Author>Yang</Author><Year>2007</Year><RecNum>133</RecNum><DisplayText>[106]</DisplayText><record><rec-number>133</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613567241">133</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Yang,Li-Shi</author><author>Yin,Zhi-Xin</author><author>Liao,Ji-Xiang</author><author>Huang,Xian-De</author><author>Guo,Chang-Jun</author><author>Weng,Shao-Ping</author><author>Chan,Siu-Ming</author><author>Yu,Xiao-Qiang</author><author>He,Jian-Guo%JMolecularimmunology</author></authors></contributors><titles><title>ATollreceptorinshrimp</title></titles><pages>1999-2008</pages><volume>44</volume><number>8</number><dates><year>2007</year></dates><isbn>0161-5890</isbn><urls></urls></record></Cite></EndNote>[106]。目前，在日本囊对虾（Marsupenaeusjaponicus）、中国对虾（Fenneropenaeuschinensis）、斑节对虾（Penaeusmonodon）、凡纳滨对虾中均报道了TLR，并已证明其参与细菌和病毒的防御ADDINEN.CITEADDINEN.CITE.DATA[107-109]。另一种研究较多的PRRs——凝集素（Lectin）包含碳水化合物识别结构域（CRD），它能够特异性结合细胞表面的多种碳水化合物，促进细胞吞噬和诱导细菌凝集。目前，在虾中已鉴定出7种凝集素，即C型、L型、P型、M型，纤维蛋白原样结构域凝集素，半乳糖凝集素和钙连蛋白/钙网蛋白，其中研究较多的是C型凝集素，主要在肝胰腺或血细胞中合成ADDINEN.CITE<EndNote><Cite><Author>Tassanakajon</Author><Year>2013</Year><RecNum>137</RecNum><DisplayText>[110]</DisplayText><record><rec-number>137</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613572245">137</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Tassanakajon,A.</author><author>Somboonwiwat,K.</author><author>Supungul,P.</author><author>Tang,S.</author></authors></contributors><auth-address>CenterofExcellenceforMolecularBiologyandGenomicsofShrimp,DepartmentofBiochemistry,FacultyofScience,ChulalongkornUniversity,254PhayathaiRoad,Bangkok10330,Thailand.anchalee.k@chula.ac.th</auth-address><titles><title>Discoveryofimmunemoleculesandtheircrucialfunctionsinshrimpimmunity</title><secondary-title>FishShellfishImmunol</secondary-title><alt-title>Fish&shellfishimmunology</alt-title></titles><periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></periodical><alt-periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></alt-periodical><pages>954-67</pages><volume>34</volume><number>4</number><edition>2012/10/13</edition><keywords><keyword>Animals</keyword><keyword>Hemocytes/immunology</keyword><keyword>Hemolymph/immunology</keyword><keyword>Immunity,Innate/genetics/immunology</keyword><keyword>Penaeidae/genetics/*immunology</keyword></keywords><dates><year>2013</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>1050-4648</isbn><accession-num>23059654</accession-num><urls></urls><electronic-resource-num>10.1016/j.fsi.2012.09.021</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[110]。然而，大多数对虾PRRs的研究仅集中在生物活性和生化特性上，PRRs的功能研究，尤其是体内参与免疫反应的研究仍然不足ADDINEN.CITE<EndNote><Cite><Author>Wang</Author><Year>2013</Year><RecNum>132</RecNum><DisplayText>[100]</DisplayText><record><rec-number>132</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613553056">132</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Wang,X.W.</author><author>Wang,J.X.</author></authors></contributors><auth-address>TheKeyLaboratoryofPlantCellEngineeringandGermplasmInnovationofMinistryofEducation,SchoolofLifeSciences,ShandongUniversity,Jinan,Shandong250100,China.</auth-address><titles><title>Patternrecognitionreceptorsactingininnateimmunesystemofshrimpagainstpathogeninfections</title><secondary-title>FishShellfishImmunol</secondary-title><alt-title>Fish&shellfishimmunology</alt-title></titles><periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></periodical><alt-periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></alt-periodical><pages>981-9</pages><volume>34</volume><number>4</number><edition>2012/09/11</edition><keywords><keyword>AminoAcidSequence</keyword><keyword>Animals</keyword><keyword>Aquaculture</keyword><keyword>Host-PathogenInteractions/immunology</keyword><keyword>Immunity,Innate/immunology</keyword><keyword>MolecularSequenceData</keyword><keyword>Penaeidae/*immunology</keyword><keyword>Receptors,PatternRecognition/*immunology</keyword><keyword>SignalTransduction/immunology</keyword></keywords><dates><year>2013</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>1050-4648</isbn><accession-num>22960101</accession-num><urls></urls><electronic-resource-num>10.1016/j.fsi.2012.08.008</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[100]。1.2体液免疫体液免疫反应包括凝血反应，多种抗菌肽（antimicrobialpeptide，AMP）的合成，酚氧化酶原（prophenoloxidase，proPO）系统以及应激反应蛋白和分子的释放ADDINEN.CITEADDINEN.CITE.DATA[111]。1.2.1凝血反应对虾具有开放的循环系统，其外骨骼是抵御病原体以及维持血淋巴和组织完整性的重要屏障，当它受到破坏时，为防止失血和微生物的入侵，损伤部位的快速凝血是重要的免疫机制ADDINEN.CITE<EndNote><Cite><Author>Tassanakajon</Author><Year>2013</Year><RecNum>137</RecNum><DisplayText>[110]</DisplayText><record><rec-number>137</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613572245">137</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Tassanakajon,A.</author><author>Somboonwiwat,K.</author><author>Supungul,P.</author><author>Tang,S.</author></authors></contributors><auth-address>CenterofExcellenceforMolecularBiologyandGenomicsofShrimp,DepartmentofBiochemistry,FacultyofScience,ChulalongkornUniversity,254PhayathaiRoad,Bangkok10330,Thailand.anchalee.k@chula.ac.th</auth-address><titles><title>Discoveryofimmunemoleculesandtheircrucialfunctionsinshrimpimmunity</title><secondary-title>FishShellfishImmunol</secondary-title><alt-title>Fish&shellfishimmunology</alt-title></titles><periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></periodical><alt-periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></alt-periodical><pages>954-67</pages><volume>34</volume><number>4</number><edition>2012/10/13</edition><keywords><keyword>Animals</keyword><keyword>Hemocytes/immunology</keyword><keyword>Hemolymph/immunology</keyword><keyword>Immunity,Innate/genetics/immunology</keyword><keyword>Penaeidae/genetics/*immunology</keyword></keywords><dates><year>2013</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>1050-4648</isbn><accession-num>23059654</accession-num><urls></urls><electronic-resource-num>10.1016/j.fsi.2012.09.021</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[110]。甲壳类动物的凝血反应通常伴随着损伤部位的血细胞聚集而发生，在新的角质层形成之前形成一个暂时的细胞屏障。迄今为止，在无脊椎动物中报道了两种凝血机制。一种在甲壳动物和昆虫中，由血细胞转谷氨酰胺酶（transglutaminase，TGase）促使凝结蛋白（clottingprotein，CP）聚合而形成的凝块或凝胶组成；另一种存在于中华鲎（Tachypleustridentatus）体内，由微生物细胞壁成分（如LPS和β-1,3-葡聚糖）激活的蛋白水解级联反应组成ADDINEN.CITEADDINEN.CITE.DATA[112]。已有研究证明对虾凝血反应包括CP和TGaseADDINEN.CITE<EndNote><Cite><Author>Maningas</Author><Year>2013</Year><RecNum>139</RecNum><DisplayText>[113]</DisplayText><record><rec-number>139</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613574496">139</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Maningas,M.B.</author><author>Kondo,H.</author><author>Hirono,I.</author></authors></contributors><auth-address>GraduateSchoolofMarineScienceandTechnology,TokyoUniversityofMarineScienceandTechnology,Konan4-5-7,Tokyo108-8477,Japan.</auth-address><titles><title>Molecularmechanismsoftheshrimpclottingsystem</title><secondary-title>FishShellfishImmunol</secondary-title><alt-title>Fish&shellfishimmunology</alt-title></titles><periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></periodical><alt-periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></alt-periodical><pages>968-72</pages><volume>34</volume><number>4</number><edition>2012/10/10</edition><keywords><keyword>Animals</keyword><keyword>AntimicrobialCationicPeptides/immunology</keyword><keyword>Aquaculture</keyword><keyword>BloodCoagulation/immunology</keyword><keyword>Immunity,Innate/immunology</keyword><keyword>Penaeidae/*immunology</keyword><keyword>SerineEndopeptidases/*immunology</keyword><keyword>Transglutaminases/immunology</keyword></keywords><dates><year>2013</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>1050-4648</isbn><accession-num>23044383</accession-num><urls></urls><electronic-resource-num>10.1016/j.fsi.2012.09.018</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[113]。其中，CP已在凡纳滨对虾、斑节对虾和日本囊对虾中得到鉴定，分子量均约为380~420kDa且N末端高度同源ADDINEN.CITEADDINEN.CITE.DATA[114,115]。对虾CP为同源二聚体糖蛋白，由两个相同的亚基组成，通过二硫键共价连接ADDINEN.CITEADDINEN.CITE.DATA[112]。TGase是一种钙依赖性酶，能够在CP间形成共价的γ-谷氨酰-ε-赖氨酸键ADDINEN.CITEADDINEN.CITE.DATA[116,117]（图5）。当对虾组织受伤或感染后，TGase从血细胞或其他组织中释放出来，以钙依赖的方式催化血浆蛋白CP的交联和聚集。有研究表明，血浆蛋白主要成分血蓝蛋白可能通过调节虾中的TGase来积极调节血淋巴凝结ADDINEN.CITEADDINEN.CITE.DATA[118]。此外，虾的凝血反应还与AMP的激活有关ADDINEN.CITEADDINEN.CITE.DATA[119]。图5对虾凝血反应ADDINEN.CITE<EndNote><Cite><Author>Sritunyalucksana</Author><Year>2000</Year><RecNum>141</RecNum><DisplayText>[120]</DisplayText><record><rec-number>141</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613630490">141</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Sritunyalucksana,Kallaya</author><author>Soderhall,Kenneth%JAquaculture</author></authors></contributors><titles><title>TheProPOandClottingSysteminCrustaceans</title></titles><pages>53-69</pages><volume>191</volume><number>1-3</number><dates><year>2000</year></dates><urls></urls></record></Cite></EndNote>[120]Fig5Clottingreactioninshrimp1.2.2AMPAMP广泛存在于无脊椎动物中，是先天免疫系统的重要效应分子，并作为抵御入侵微生物的第一道防线ADDINEN.CITE<EndNote><Cite><Author>Hancock</Author><Year>2000</Year><RecNum>161</RecNum><DisplayText>[121]</DisplayText><record><rec-number>161</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613744723">161</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Hancock,R.E.</author><author>Diamond,G.</author></authors></contributors><auth-address>DeptofMicrobiologyandImmunology,UniversityofBritishColumbia,300-6174UniversityBlvd,Vancouver,CanadaV6T1Z3.bob@cmdr.ubc.ca</auth-address><titles><title>Theroleofcationicantimicrobialpeptidesininnatehostdefences</title><secondary-title>TrendsMicrobiol</secondary-title><alt-title>Trendsinmicrobiology</alt-title></titles><periodical><full-title>TrendsMicrobiol</full-title><abbr-1>Trendsinmicrobiology</abbr-1></periodical><alt-periodical><full-title>TrendsMicrobiol</full-title><abbr-1>Trendsinmicrobiology</abbr-1></alt-periodical><pages>402-10</pages><volume>8</volume><number>9</number><edition>2000/09/16</edition><keywords><keyword>AminoAcidSequence</keyword><keyword>Anti-InfectiveAgents/*pharmacology</keyword><keyword>BaseSequence</keyword><keyword>Cations/pharmacology</keyword><keyword>*Immunity,Innate</keyword><keyword>Models,Biological</keyword><keyword>MolecularSequenceData</keyword><keyword>Peptides/genetics/*pharmacology</keyword></keywords><dates><year>2000</year><pub-dates><date>Sep</date></pub-dates></dates><isbn>0966-842X(Print) 0966-842x</isbn><accession-num>10989307</accession-num><urls></urls><electronic-resource-num>10.1016/s0966-842x(00)01823-0</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[121]。AMP的特异性低，对细菌、真菌、病毒等具有广谱抗性，其水溶性好且大部分耐热性强ADDINEN.CITE<EndNote><Cite><Author>广东农业科学</Author><Year>2012</Year><RecNum>159</RecNum><DisplayText>[122,123]</DisplayText><record><rec-number>159</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613744368">159</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>杜志强%J广东农业科学</author></authors></contributors><titles><title>先天免疫系统中抗菌肽的研究进展</title></titles><dates><year>2012</year></dates><urls></urls></record></Cite><Cite><Author>Aguirre-Guzman</Author><Year>2009</Year><RecNum>160</RecNum><record><rec-number>160</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613744561">160</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Aguirre-Guzman,Gabriel</author><author>Sanchez-Martinez,JesusGenaro</author><author>Campa-Cordova,AngelIsidro</author><author>Luna-Gonzalez,Antonio</author><author>Ascencio,Felipe%JTheThaiJournalofVeterinaryMedicine</author></authors></contributors><titles><title>Penaeidshrimpimmunesystem</title></titles><pages>205-215</pages><volume>39</volume><number>3</number><dates><year>2009</year></dates><isbn>0125-6491</isbn><urls></urls></record></Cite></EndNote>[122,123]。作为一种小分子多肽，AMP通常少于150~200个氨基酸残基。在对虾中，AMP由血细胞产生并储存，已鉴定了几个对虾AMP家族：对虾素（penaeidin，PEN）、溶菌酶（lysozymes，Lyz）、甲壳素（crustins，Cru）、抗脂多糖因子（antilipopo1vsaccharidefactor，ALF）和StylicinADDINEN.CITEADDINEN.CITE.DATA[124,125]。PEN是对虾特有的AMP家族，最初在凡纳滨对虾中发现，现已从9种对虾中分离出来ADDINEN.CITEADDINEN.CITE.DATA[125-129]。PEN具有独特的分子结构，包含一个高度保守的前导肽，N末端有一个富含脯氨酸的结构域（proline-richdomain，PRD），C末端有6个半胱氨酸残基，形成3个二硫键。PEN家族目前由五个相关分子组成，它们对海洋革兰氏阳性细菌和真菌具有抗菌活性。然而，不同类别的PEN由于其序列的可变性而显示出不同的目标特异性ADDINEN.CITE<EndNote><Cite><Author>Tassanakajon</Author><Year>2013</Year><RecNum>137</RecNum><DisplayText>[110]</DisplayText><record><rec-number>137</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613572245">137</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Tassanakajon,A.</author><author>Somboonwiwat,K.</author><author>Supungul,P.</author><author>Tang,S.</author></authors></contributors><auth-address>CenterofExcellenceforMolecularBiologyandGenomicsofShrimp,DepartmentofBiochemistry,FacultyofScience,ChulalongkornUniversity,254PhayathaiRoad,Bangkok10330,Thailand.anchalee.k@chula.ac.th</auth-address><titles><title>Discoveryofimmunemoleculesandtheircrucialfunctionsinshrimpimmunity</title><secondary-title>FishShellfishImmunol</secondary-title><alt-title>Fish&shellfishimmunology</alt-title></titles><periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></periodical><alt-periodical><full-title>FishShellfishImmunol</full-title><abbr-1>Fish&shellfishimmunology</abbr-1></alt-periodical><pages>954-67</pages><volume>34</volume><number>4</number><edition>2012/10/13</edition><keywords><keyword>Animals</keyword><keyword>Hemocytes/immunology</keyword><keyword>Hemolymph/immunology</keyword><keyword>Immunity,Innate/genetics/immunology</keyword><keyword>Penaeidae/genetics/*immunology</keyword></keywords><dates><year>2013</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>1050-4648</isbn><accession-num>23059654</accession-num><urls></urls><electronic-resource-num>10.1016/j.fsi.2012.09.021</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[110]。PEN合成并储存在颗粒细胞中，响应病原体刺激而释放ADDINEN.CITE<EndNote><Cite><Author>Vazquez</Author><Year>2009</Year><RecNum>169</RecNum><DisplayText>[130]</DisplayText><record><rec-number>169</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613794960">169</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Vazquez,L.</author><author>Alpuche,J.</author><author>Maldonado,G.</author><author>Agundis,C.</author><author>Pereyra-Morales,A.</author><author>Zenteno,E.</author></authors></contributors><auth-address>LaboratoriodeLectinas,CIQ,UniversidadAutonomadelEstadodeMorelos,Cuernavaca,Morelos,Mexico.lvazquez@mklinux.dti.uaem.mx</auth-address><titles><title>Review:Immunitymechanismsincrustaceans</title><secondary-title>InnateImmun</secondary-title><alt-title>Innateimmunity</alt-title></titles><periodical><full-title>InnateImmun</full-title><abbr-1>Innateimmunity</abbr-1></periodical><alt-periodical><full-title>InnateImmun</full-title><abbr-1>Innateimmunity</abbr-1></alt-periodical><pages>179-88</pages><volume>15</volume><number>3</number><edition>2009/05/29</edition><keywords><keyword>Animals</keyword><keyword>AntimicrobialCationicPeptides/*immunology/metabolism</keyword><keyword>CatecholOxidase/*immunology/metabolism</keyword><keyword>Crustacea/*immunology/metabolism</keyword><keyword>EnzymePrecursors/*immunology/metabolism</keyword><keyword>Immunity,Innate</keyword><keyword>Lectins/*immunology/metabolism</keyword><keyword>Phagocytosis/*immunology</keyword><keyword>Receptors,PatternRecognition/*immunology/metabolism</keyword></keywords><dates><year>2009</year><pub-dates><date>Jun</date></pub-dates></dates><isbn>1753-4259(Print) 1753-4259</isbn><accession-num>19474211</accession-num><urls></urls><electronic-resource-num>10.1177/1753425909102876</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[130]。除了作为AMP外，PEN还可作为细胞因子发挥免疫调节作用，在体外促进整合素（integrin）介导的颗粒细胞和半颗粒细胞粘附ADDINEN.CITEADDINEN.CITE.DATA[131]。Lyz被分为三类：鸡型Lyz（c型）、鹅型Lyz（g型）和无脊椎动物型Lyz（i型）。在对虾中，仅报道了c型和i型Lyz，而c型Lyz占大多数。Lyz能溶解革兰氏阳性菌和革兰氏阴性菌，对溶藻弧菌和副溶血弧菌均具有很强的抑制作用ADDINEN.CITE<EndNote><Cite><Author>Tassanakajon</Author><Year>2013</Year><RecNum>137</RecNum><DisplayText>[110]</DisplayText><record><rec-number>137</rec-number><foreign-keys><keyapp="EN"db-id="pw2fxtxfdepdfses99t5fzpdtpa0vv0xd909"timestamp="1613572245">137</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Tassanakajon,A.</author><author>Somboonwiwat,K.</author><author>Supungul,P.</author><author>Tang,S.</author></authors></contributors><auth-address>CenterofExc