【《果蝇的机械力感受细胞研究文献综述》4000字】

果蝇的机械力感受细胞研究文献综述果蝇成虫和幼虫根据其生活环境和发育状态不同，具有不同偏好类型的力敏感受体。果蝇成虫体表及体内分布了大量的外感受器官和弦音受体ADDINEN.CITE<EndNote><Cite><Author>Lorimer</Author><Year>2015</Year><RecNum>611</RecNum><DisplayText>(Lorimeretal.,2015)</DisplayText><record><rec-number>611</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614304104">611</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Lorimer,T.</author><author>Gomez,F.</author><author>Stoop,R.</author></authors></contributors><auth-address>UnivZurich,InstNeuroinformat,CH-8057Zurich,Switzerland UnivZurich,InstComputatSci,CH-8057Zurich,Switzerland UnivZurich,CH-8057Zurich,Switzerland</auth-address><titles><title>Mammaliancochleaasaphysicsguidedevolution-optimizedhearingsensor</title><secondary-title>ScientificReports</secondary-title><alt-title>SciRep-Uk</alt-title></titles><periodical><full-title>ScientificReports</full-title></periodical><volume>5</volume><keywords><keyword>outerhair-cells</keyword><keyword>convergentevolution</keyword><keyword>pitchsensation</keyword><keyword>insecthearing</keyword><keyword>organs</keyword><keyword>amplification</keyword><keyword>drosophila</keyword></keywords><dates><year>2015</year><pub-dates><date>Jul28</date></pub-dates></dates><isbn>2045-2322</isbn><accession-num>WOS:000358586600001</accession-num><urls><related-urls><url><GotoISI>://WOS:000358586600001</url></related-urls></urls><electronic-resource-num>ARTN12492 10.1038/srep12492</electronic-resource-num><language>English</language></record></Cite></EndNote>(Lorimeretal.,2015)。外部感觉器官的毛受体和钟型受体，均具有特化的表皮结构。一种为短毛，另一种为突出于体表的穹顶结构ADDINEN.CITE<EndNote><Cite><Author>Keil</Author><Year>1997</Year><RecNum>612</RecNum><DisplayText>(Keil,1997)</DisplayText><record><rec-number>612</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614304838">612</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Keil,T.A.</author></authors></contributors><auth-address>ArbeitsgruppeKaissling,Max-Planck-InstitutfurVerhaltensphysiologie,Seewiesen,Germany.</auth-address><titles><title>Functionalmorphologyofinsectmechanoreceptors</title><secondary-title>MicroscResTech</secondary-title></titles><periodical><full-title>MicroscResTech</full-title></periodical><pages>506-31</pages><volume>39</volume><number>6</number><edition>1998/01/23</edition><keywords><keyword>Animals</keyword><keyword>Insecta/*cytology/*physiology</keyword><keyword>Mechanoreceptors/*cytology/*physiology/ultrastructure</keyword><keyword>Neurons/*physiology</keyword></keywords><dates><year>1997</year><pub-dates><date>Dec15</date></pub-dates></dates><isbn>1059-910X(Print) 1059-910X(Linking)</isbn><accession-num>9438251</accession-num><urls><related-urls><url>/pubmed/9438251</url></related-urls></urls><electronic-resource-num>10.1002/(SICI)1097-0029(19971215)39:6<506::AID-JEMT5>3.0.CO;2-B</electronic-resource-num></record></Cite></EndNote>(Keil,1997)。这些特化的结构可以将外界的机械刺激通过短毛或体表结构的变形，引起内部结构的变化，继而将力转导到神经元树突的顶端ADDINEN.CITE<EndNote><Cite><Author>Keil</Author><Year>1997</Year><RecNum>612</RecNum><DisplayText>(Keil,1997)</DisplayText><record><rec-number>612</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614304838">612</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Keil,T.A.</author></authors></contributors><auth-address>ArbeitsgruppeKaissling,Max-Planck-InstitutfurVerhaltensphysiologie,Seewiesen,Germany.</auth-address><titles><title>Functionalmorphologyofinsectmechanoreceptors</title><secondary-title>MicroscResTech</secondary-title></titles><periodical><full-title>MicroscResTech</full-title></periodical><pages>506-31</pages><volume>39</volume><number>6</number><edition>1998/01/23</edition><keywords><keyword>Animals</keyword><keyword>Insecta/*cytology/*physiology</keyword><keyword>Mechanoreceptors/*cytology/*physiology/ultrastructure</keyword><keyword>Neurons/*physiology</keyword></keywords><dates><year>1997</year><pub-dates><date>Dec15</date></pub-dates></dates><isbn>1059-910X(Print) 1059-910X(Linking)</isbn><accession-num>9438251</accession-num><urls><related-urls><url>/pubmed/9438251</url></related-urls></urls><electronic-resource-num>10.1002/(SICI)1097-0029(19971215)39:6<506::AID-JEMT5>3.0.CO;2-B</electronic-resource-num></record></Cite></EndNote>(Keil,1997)。弦音受体没有外部特化的表皮结构，通常情况下这类受体是嵌在表皮之下。幼虫主要的机械感受神经元主要为II型机械感受器，分布于体表，包括1型（classI）、2型（classII）、3型（classIII）和4型（classIV）神经元，这类神经元存在于幼虫体表的每个节段内，并位于表皮层。它们有复杂的分支树突结构和一个向中枢神经系统提供信息的轴突。其中研究较为广泛的为3型和4型神经元ADDINEN.CITEADDINEN.CITE.DATA(Chengetal.,2010;Robertsonetal.,2013;Tsubouchietal.,2012)，它们在机械感受中的作用也被揭示：3型神经元对轻微触碰刺激敏感ADDINEN.CITEADDINEN.CITE.DATA(Yanetal.,2013a)，而4型神经元对强烈触碰刺激敏感ADDINEN.CITEADDINEN.CITE.DATA(Kimetal.,ab2012;Zhongetal.,2010)。ab图1.4果蝇不同机械敏感受体示例图a.果蝇成虫体表受体，包括最大的弦音受体(1)黑色箭头所示、体表的毛受体(2)和平衡棒上的钟型受体阵列(3)。b.果蝇幼虫体表的Ⅱ型机械力敏感神经元（上图是Ⅲ型神经元，下图是Ⅳ型神经元）ADDINEN.CITE<EndNote><Cite><Author>Liang</Author><Year>2017</Year><RecNum>595</RecNum><DisplayText>(Liang,2017)</DisplayText><record><rec-number>595</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614248128">595</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Liang,X.,Sun,L.,andLiu,Z</author></authors></contributors><titles><title><2017_Book_MechanosensoryTransductionInDr.pdf></title></titles><dates><year>2017</year></dates><urls></urls></record></Cite></EndNote>(Liang,2017)。毛受体和弦音器概述果蝇的毛受体覆盖于整个体表，主要分为两种，较粗大的称为巨毛类，较细小的称为微毛类。巨毛类刚毛通常位于果蝇体表几个固定的位置，例如果蝇头部、背腹部刚毛和前角质鳞化刚毛等。微毛类则几乎分布于整个蝇体。这些毛受体可以感知到由环境中机械力所引起的感受器结构变形的刺激，例如触觉、气流、重力等ADDINEN.CITE<EndNote><Cite><Author>Gaffal</Author><Year>1975</Year><RecNum>623</RecNum><DisplayText>(Gaffaletal.,1975)</DisplayText><record><rec-number>623</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614316788">623</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Gaffal,K.P.</author><author>Tichy,H.</author><author>Theiss,J.</author><author>Seelinger,G.</author></authors></contributors><auth-address>UnivRegensburg,ZoolInst,D-8400Regensburg,FedRepGer</auth-address><titles><title>StructuralPolaritiesinMechanosensitiveSensillaandTheirInfluenceonStimulusTransmission(Arthropoda)</title><secondary-title>Zoomorphologie</secondary-title><alt-title>Zoomorphologie</alt-title></titles><periodical><full-title>Zoomorphologie</full-title><abbr-1>Zoomorphologie</abbr-1></periodical><alt-periodical><full-title>Zoomorphologie</full-title><abbr-1>Zoomorphologie</abbr-1></alt-periodical><pages>79-103</pages><volume>82</volume><number>2-3</number><dates><year>1975</year></dates><accession-num>WOS:A1975BF61700001</accession-num><urls><related-urls><url><GotoISI>://WOS:A1975BF61700001</url></related-urls></urls><electronic-resource-num>Doi10.1007/Bf00993585</electronic-resource-num><language>English</language></record></Cite></EndNote>(Gaffaletal.,1975)。例如当果蝇生存环境中有螨虫时，果蝇翅膀下的刚毛会参与其机械感受过程，将螨虫从体表踢下从而起到防御功能(Lietal.2016)。毛受体的结构的角质层端口处呈无定形，多为内向沟槽状。毛发通过弹性关节膜连接到身体角质层的圆形或椭圆形凹窝ADDINEN.CITE<EndNote><Cite><Author>Blackwell</Author><Year>1980</Year><RecNum>622</RecNum><DisplayText>(BlackwellandWeih,1980)</DisplayText><record><rec-number>622</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614316653">622</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Blackwell,J.</author><author>Weih,M.A.</author></authors></contributors><titles><title>Structureofchitin-proteincomplexes:ovipositoroftheichneumonflyMegarhyssa</title><secondary-title>JMolBiol</secondary-title></titles><periodical><full-title>JMolBiol</full-title></periodical><pages>49-60</pages><volume>137</volume><number>1</number><edition>1980/02/15</edition><keywords><keyword>Animals</keyword><keyword>*Chitin</keyword><keyword>Female</keyword><keyword>*Hymenoptera/anatomy&histology</keyword><keyword>*Models,Molecular</keyword><keyword>*Models,Structural</keyword><keyword>Oviposition</keyword><keyword>ProteinConformation</keyword><keyword>*Proteins</keyword><keyword>X-RayDiffraction</keyword></keywords><dates><year>1980</year><pub-dates><date>Feb15</date></pub-dates></dates><isbn>0022-2836(Print) 0022-2836(Linking)</isbn><accession-num>7365796</accession-num><urls><related-urls><url>/pubmed/7365796</url></related-urls></urls><electronic-resource-num>10.1016/0022-2836(80)90156-4</electronic-resource-num></record></Cite></EndNote>(BlackwellandWeih,1980)。大多数的毛受体的毛发都是以一级杠杆的构造形式存在。树突顶端仅通过树突鞘间接与毛发基部相连，主要由皮下槽状隔膜支撑ADDINEN.CITE<EndNote><Cite><Author>Gaffal</Author><Year>1975</Year><RecNum>623</RecNum><DisplayText>(Gaffaletal.,1975)</DisplayText><record><rec-number>623</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614316788">623</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Gaffal,K.P.</author><author>Tichy,H.</author><author>Theiss,J.</author><author>Seelinger,G.</author></authors></contributors><auth-address>UnivRegensburg,ZoolInst,D-8400Regensburg,FedRepGer</auth-address><titles><title>StructuralPolaritiesinMechanosensitiveSensillaandTheirInfluenceonStimulusTransmission(Arthropoda)</title><secondary-title>Zoomorphologie</secondary-title><alt-title>Zoomorphologie</alt-title></titles><periodical><full-title>Zoomorphologie</full-title><abbr-1>Zoomorphologie</abbr-1></periodical><alt-periodical><full-title>Zoomorphologie</full-title><abbr-1>Zoomorphologie</abbr-1></alt-periodical><pages>79-103</pages><volume>82</volume><number>2-3</number><dates><year>1975</year></dates><accession-num>WOS:A1975BF61700001</accession-num><urls><related-urls><url><GotoISI>://WOS:A1975BF61700001</url></related-urls></urls><electronic-resource-num>Doi10.1007/Bf00993585</electronic-resource-num><language>English</language></record></Cite></EndNote>(Gaffaletal.,1975)。在受体的体内部分，都包含一个纤毛化的双极化感觉神经元。该神经元具有一个无分支的树突、一个胞体和一个轴突。神经元树突远端的2~3μm区域特化为一个变态纤毛modifiedcilium）结构，具有经典的“9+0”微管轴丝。在纤毛区中段，轴突处基于“9+0”的轴丝转变为一种特殊的高度有序的微管结构，呈现扩张膨大状(在部分结构中，呈现花托状)，这个结构称为管状体（tubularbody，TB）ADDINEN.CITEADDINEN.CITE.DATA(Liangetal.,2013;Liangetal.,2014)。管状体的远端与毛发结构直接连接，当毛发发生位置改变时，会引起管状体内微管等相应结构产生生理变化，因此，它被认为是机械转导的位置，其中包含机械感受器，将机械刺激信号转导为细胞内信号ADDINEN.CITEADDINEN.CITE.DATA(Liangetal.,2017b;Walkeretal.,2000)。钟型受体中的神经元具有和毛受体类似的内部结构，在下1.1.4章节将作重点介绍。图1.5果蝇毛受体结构示例图果蝇毛受体具有典型的纤毛化神经元结构。a.毛受体神经元（GFP标记的神经元）的荧光图像。b.毛受体结构模式图，毛受体感觉神经元有胞体、树突、刚毛、基体、轴突、神经外支持细胞等结构，受体神经元外周被支持细胞所包围。ADDINEN.CITE<EndNote><Cite><Author>Liang</Author><Year>2017</Year><RecNum>595</RecNum><DisplayText>(Keil,1997;Liang,2017)</DisplayText><record><rec-number>595</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614248128">595</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Liang,X.,Sun,L.,andLiu,Z</author></authors></contributors><titles><title><2017_Book_MechanosensoryTransductionInDr.pdf></title></titles><dates><year>2017</year></dates><urls></urls></record></Cite><Cite><Author>Keil</Author><Year>1997</Year><RecNum>612</RecNum><record><rec-number>612</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614304838">612</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Keil,T.A.</author></authors></contributors><auth-address>ArbeitsgruppeKaissling,Max-Planck-InstitutfurVerhaltensphysiologie,Seewiesen,Germany.</auth-address><titles><title>Functionalmorphologyofinsectmechanoreceptors</title><secondary-title>MicroscResTech</secondary-title></titles><periodical><full-title>MicroscResTech</full-title></periodical><pages>506-31</pages><volume>39</volume><number>6</number><edition>1998/01/23</edition><keywords><keyword>Animals</keyword><keyword>Insecta/*cytology/*physiology</keyword><keyword>Mechanoreceptors/*cytology/*physiology/ultrastructure</keyword><keyword>Neurons/*physiology</keyword></keywords><dates><year>1997</year><pub-dates><date>Dec15</date></pub-dates></dates><isbn>1059-910X(Print) 1059-910X(Linking)</isbn><accession-num>9438251</accession-num><urls><related-urls><url>/pubmed/9438251</url></related-urls></urls><electronic-resource-num>10.1002/(SICI)1097-0029(19971215)39:6<506::AID-JEMT5>3.0.CO;2-B</electronic-resource-num></record></Cite></EndNote>(Keil,1997;Liang,2017)。弦音器（Chordotonalorgans）又称为剑梢感受器（Scolopoidsensilla）是存在于昆虫体内可以感知声音、风力和重力等机械刺激的一种外感受器，同时还参与关节运动等本体感受ADDINEN.CITEADDINEN.CITE.DATA(KavlieandAlbert,2013;Keil,1997)。它们由227个弦音受体单元(scolopidium)所构成，每个受体单元中包含2~3个神经元及其附属细胞。不同类型的昆虫，以及不同种类的弦音器所包含的弦音细胞数量也不同，果蝇中一般有1-3个ADDINEN.CITEADDINEN.CITE.DATA(Ghysenetal.,1986;KavlieandAlbert,2013)。果蝇幼虫的弦音感受器没有体外结构，体内结构同前部分所述的毛受体结构类似，弦音感觉细胞是包含树突特化微管结构的一个双极神经元细胞ADDINEN.CITE<EndNote><Cite><Author>Boo</Author><Year>1975</Year><RecNum>706</RecNum><DisplayText>(BooandRichards,1975)</DisplayText><record><rec-number>706</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614326607">706</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Boo,K.S.</author><author>Richards,A.G.</author></authors></contributors><auth-address>UnivMinnesota,DeptEntomolFisheries&Wildlife,StPaul,Mn55108</auth-address><titles><title>Fine-StructureofScolopidiainJohnstonsOrganofFemaleAedes-AegyptiComparedwithThatofMale</title><secondary-title>JournalofInsectPhysiology</secondary-title><alt-title>JInsectPhysiol</alt-title></titles><periodical><full-title>JournalofInsectPhysiology</full-title><abbr-1>JInsectPhysiol</abbr-1></periodical><alt-periodical><full-title>JournalofInsectPhysiology</full-title><abbr-1>JInsectPhysiol</abbr-1></alt-periodical><pages>1129-1139</pages><volume>21</volume><number>5</number><dates><year>1975</year></dates><isbn>0022-1910</isbn><accession-num>WOS:A1975AA96700019</accession-num><urls><related-urls><url><GotoISI>://WOS:A1975AA96700019</url></related-urls></urls><electronic-resource-num>Doi10.1016/0022-1910(75)90126-2</electronic-resource-num><language>English</language></record></Cite></EndNote>(BooandRichards,1975)。弦音感觉细胞纤毛化的“9+0”的轴丝结构要远远长于毛受体和钟型受体细胞。弦音感觉细胞基体位于神经元树突中间位置，将其分为内段和外段。内段（innersegment）包含纤毛根丝结构，外段（outersegment）是纤毛结构。纤毛区远端的1/3处会形成一个膨胀结构，称为纤毛扩张区。在这个区域内，可以观察到存在一种由未知物质组成的副晶体形式的电子致密结构。纤毛中的微管在纤毛扩张区弯曲，并在扩张远端变为纤毛结构ADDINEN.CITEADDINEN.CITE.DATA(Boekhoff-FalkandEberl,2014;BokoliaandMishra,2015)。树突帽被附着细胞所包裹，最终连接到感觉细胞和皮肤的角质层。在弦音器中，树突顶端是感觉细胞的机械转导位点，来自于周围的机械刺激信号将通过其附着细胞、顶端细胞和其它分子来完成机械转导功能（图1.6）ADDINEN.CITEADDINEN.CITE.DATA(KavlieandAlbert,2013;Khan,2018)。图1.6果蝇幼虫弦音受体结构示例图果蝇弦音受体具有胞体（soma）、树突（dendrite）、基体（basalbody）和较长的cilium结构并且有不同于其它感受器的纤毛膨大区（ciliarydilation），包裹神经元的导音杆细胞（scolopalecell），冠细胞（capcell）连接弦音器和体表ADDINEN.CITE<EndNote><Cite><Author>Kavlie</Author><Year>2013</Year><RecNum>703</RecNum><DisplayText>(KavlieandAlbert,2013)</DisplayText><record><rec-number>703</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614323277">703</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Kavlie,R.G.</author><author>Albert,J.T.</author></authors></contributors><auth-address>UCL,EarInst,LondonWC1X8EE,England</auth-address><titles><title>Chordotonalorgans</title><secondary-title>CurrentBiology</secondary-title><alt-title>CurrBiol</alt-title></titles><periodical><full-title>CurrentBiology</full-title></periodical><alt-periodical><full-title>CurrBiol</full-title></alt-periodical><pages>R334-R335</pages><volume>23</volume><number>9</number><keywords><keyword>johnstonsorgan</keyword></keywords><dates><year>2013</year><pub-dates><date>May6</date></pub-dates></dates><isbn>0960-9822</isbn><accession-num>WOS:000318750900002</accession-num><urls><related-urls><url><GotoISI>://WOS:000318750900002</url></related-urls></urls><electronic-resource-num>DOI10.1016/j.cub.2013.03.048</electronic-resource-num><language>English</language></record></Cite></EndNote>(KavlieandAlbert,2013)。位于果蝇成虫触角第二节的Johnston’sOrgan是其最大的弦音感受器ADDINEN.CITEADDINEN.CITE.DATA(Boekhoff-FalkandEberl,2014;BokoliaandMishra,2015;Jarmanetal.,1995)。该受体是由约480个神经元汇聚到排列规则的200多个弦音受体单元中ADDINEN.CITEADDINEN.CITE.DATA(Chalfie,2009;Kamikouchietal.,2009a)，85%的弦音受体单元中具有2个神经元，其它的包含3个神经元ADDINEN.CITEADDINEN.CITE.DATA(Todietal.,2004)。JO神经元具有单一不分叉的树突和轴突结构，树突中包含经典的“9+0”纤毛轴丝结构。Johnston’sOrgan是果蝇重要的听觉器官，它有助于果蝇对重力和风的感知ADDINEN.CITEADDINEN.CITE.DATA(Budellietal.,2019;Kamikouchietal.,2009b;Yorozuetal.,2009)以及温度感知和昼夜节律相关ADDINEN.CITE<EndNote><Cite><Author>Sehadova</Author><Year>2009</Year><RecNum>798</RecNum><DisplayText>(Sehadovaetal.,2009)</DisplayText><record><rec-number>798</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614400130">798</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Sehadova,H.</author><author>Glaser,F.T.</author><author>Gentile,C.</author><author>Simoni,A.</author><author>Giesecke,A.</author><author>Albert,J.T.</author><author>Stanewsky,R.</author></authors></contributors><auth-address>QueenMaryUnivLondon,SchBiol&ChemSci,LondonE14NS,England UnivRegensburg,InstZool,D-93040Regensburg,Germany UCL,EarInst,LondonWC1X8EE,England</auth-address><titles><title>TemperatureEntrainmentofDrosophila'sCircadianClockInvolvestheGenenocteandSignalingfromPeripheralSensoryTissuestotheBrain</title><secondary-title>Neuron</secondary-title><alt-title>Neuron</alt-title></titles><periodical><full-title>Neuron</full-title></periodical><alt-periodical><full-title>Neuron</full-title></alt-periodical><pages>251-266</pages><volume>64</volume><number>2</number><keywords><keyword>rhythms</keyword><keyword>neurons</keyword><keyword>light</keyword><keyword>synchronization</keyword><keyword>transcription</keyword><keyword>cryptochrome</keyword><keyword>oscillator</keyword><keyword>encodes</keyword><keyword>system</keyword><keyword>period</keyword></keywords><dates><year>2009</year><pub-dates><date>Oct29</date></pub-dates></dates><isbn>0896-6273</isbn><accession-num>WOS:000271889900014</accession-num><urls><related-urls><url><GotoISI>://WOS:000271889900014</url></related-urls></urls><electronic-resource-num>10.1016/j.neuron.2009.08.026</electronic-resource-num><language>English</language></record></Cite></EndNote>(Sehadovaetal.,2009)，同时在果蝇交配时，雌蝇的JO神经元还能帮助其听到“求爱之歌”ADDINEN.CITE<EndNote><Cite><Author>Dornan</Author><Year>2008</Year><RecNum>829</RecNum><DisplayText>(DornanandGoodwin,2008)</DisplayText><record><rec-number>829</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614413451">829</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Dornan,A.J.</author><author>Goodwin,S.F.</author></authors></contributors><auth-address>UnivGlasgow,IBLS,DivMolGenet,GlasgowG116NU,Lanark,Scotland</auth-address><titles><title>Flycourtshipsong:Triggeringthelightfantastic</title><secondary-title>Cell</secondary-title><alt-title>Cell</alt-title></titles><periodical><full-title>Cell</full-title></periodical><alt-periodical><full-title>Cell</full-title></alt-periodical><pages>210-212</pages><volume>133</volume><number>2</number><keywords><keyword>neuralcircuitry</keyword><keyword>drosophila</keyword><keyword>fruitless</keyword><keyword>behavior</keyword></keywords><dates><year>2008</year><pub-dates><date>Apr18</date></pub-dates></dates><isbn>0092-8674</isbn><accession-num>WOS:000255052000010</accession-num><urls><related-urls><url><GotoISI>://WOS:000255052000010</url></related-urls></urls><electronic-resource-num>10.1016/j.cell.2008.04.008</electronic-resource-num><language>English</language></record></Cite></EndNote>(DornanandGoodwin,2008)。图1.7果蝇头部触觉感受器示意图a.果蝇触觉感受器Johnston’sOrgan扫描电镜示意图，该结构具有三个组成部分。b.Johnston’sOrgan中包含数以百计的规律排列的神经元，每一个接收外界刺激的感觉纤毛神经元位于Johnston’sOrgan的A2段，Johnston’sOrgan可以感知风力、重力以及声音震动等ADDINEN.CITE<EndNote><Cite><Author>Eatock</Author><Year>2009</Year><RecNum>1309</RecNum><DisplayText>(Eatock,2009)</DisplayText><record><rec-number>1309</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1617094950">1309</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Eatock,R.A.</author></authors></contributors><titles><title>Neuroscience:Up,down,flyingaround</title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title></periodical><pages>156-7</pages><volume>458</volume><number>7235</number><edition>2009/03/13</edition><keywords><keyword>*AirMovements</keyword><keyword>Animals</keyword><keyword>Drosophila/anatomy&histology/*physiology</keyword><keyword>*GravitySensing</keyword><keyword>SensoryReceptorCells/physiology</keyword></keywords><dates><year>2009</year><pub-dates><date>Mar12</date></pub-dates></dates><isbn>1476-4687(Electronic) 0028-0836(Linking)</isbn><accession-num>19279623</accession-num><urls><related-urls><url>/pubmed/19279623</url></related-urls></urls><electronic-resource-num>10.1038/458156a</electronic-resource-num></record></Cite></EndNote>(Eatock,2009)。1.1.4果蝇钟型受体钟型感受器目前仅存在于昆虫中ADDINEN.CITE<EndNote><Cite><Author>Keil</Author><Year>1997</Year><RecNum>612</RecNum><DisplayText>(Keil,1997)</DisplayText><record><rec-number>612</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614304838">612</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Keil,T.A.</author></authors></contributors><auth-address>ArbeitsgruppeKaissling,Max-Planck-InstitutfurVerhaltensphysiologie,Seewiesen,Germany.</auth-address><titles><title>Functionalmorphologyofinsectmechanoreceptors</title><secondary-title>MicroscResTech</secondary-title></titles><periodical><full-title>MicroscResTech</full-title></periodical><pages>506-31</pages><volume>39</volume><number>6</number><edition>1998/01/23</edition><keywords><keyword>Animals</keyword><keyword>Insecta/*cytology/*physiology</keyword><keyword>Mechanoreceptors/*cytology/*physiology/ultrastructure</keyword><keyword>Neurons/*physiology</keyword></keywords><dates><year>1997</year><pub-dates><date>Dec15</date></pub-dates></dates><isbn>1059-910X(Print) 1059-910X(Linking)</isbn><accession-num>9438251</accession-num><urls><related-urls><url>/pubmed/9438251</url></related-urls></urls><electronic-resource-num>10.1002/(SICI)1097-0029(19971215)39:6<506::AID-JEMT5>3.0.CO;2-B</electronic-resource-num></record></Cite></EndNote>(Keil,1997)。在果蝇体表，它主要分布于不同身体部位之间的关节处，例如腿部、翅膀和平衡棒等处。钟型受体的外形呈无毛的穹顶结构，当果蝇爬行、飞行和跳跃时，它们能感觉到角质层的变形。果蝇翅膀基部有一小片的钟型力感受器ADDINEN.CITE<EndNote><Cite><Author>Keil</Author><Year>1997</Year><RecNum>612</RecNum><DisplayText>(Keil,1997)</DisplayText><record><rec-number>612</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614304838">612</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Keil,T.A.</author></authors></contributors><auth-address>ArbeitsgruppeKaissling,Max-Planck-InstitutfurVerhaltensphysiologie,Seewiesen,Germany.</auth-address><titles><title>Functionalmorphologyofinsectmechanoreceptors</title><secondary-title>MicroscResTech</secondary-title></titles><periodical><full-title>MicroscResTech</full-title></periodical><pages>506-31</pages><volume>39</volume><number>6</number><edition>1998/01/23</edition><keywords><keyword>Animals</keyword><keyword>Insecta/*cytology/*physiology</keyword><keyword>Mechanoreceptors/*cytology/*physiology/ultrastructure</keyword><keyword>Neurons/*physiology</keyword></keywords><dates><year>1997</year><pub-dates><date>Dec15</date></pub-dates></dates><isbn>1059-910X(Print) 1059-910X(Linking)</isbn><accession-num>9438251</accession-num><urls><related-urls><url>/pubmed/9438251</url></related-urls></urls><electronic-resource-num>10.1002/(SICI)1097-0029(19971215)39:6<506::AID-JEMT5>3.0.CO;2-B</electronic-resource-num></record></Cite></EndNote>(Keil,1997)。这些感受器是果蝇中面积最大的，该区域称平衡棒（haltere）。平衡棒是退化的后翅，属于附肢ADDINEN.CITEADDINEN.CITE.DATA(HerranzandMorata,2001;Raadetal.,2016)。这些钟型感受器对果蝇的飞行行为极为重要ADDINEN.CITEADDINEN.CITE.DATA(Liangetal.,2014;Pringle,1948)。果蝇具有非常好的飞行能力。它们的翅膀可以在空中非常快速的扇动ADDINEN.CITEADDINEN.CITE.DATA(Chanetal.,1998;RochandAkam,2000)。然而果蝇具有稳定的飞行能力的前提是机动飞行信号可以在中枢神经系统和运动系统中正确的识别和传递，否则果蝇会失去身体平衡ADDINEN.CITE<EndNote><Cite><Author>Thurm</Author><Year>1983</Year><RecNum>795</RecNum><DisplayText>(Thurmetal.,1983)</DisplayText><record><rec-number>795</rec-number><foreign-keys><keyapp="EN"db-id="vrwdadt98e5druedp2b5e2dc9wz2des0w22f"timestamp="1614399323">795</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Thurm,U.</author><author>Erler,G.</author><author>Godde,J.</author><author>Kastrup,H.</author><author>Keil,T.</author><author>Volker,W.</author><author>Vohwinkel,B.</author></authors></contributors><titles><title>CiliaSpecializedforMechanoreception</title><secondary-title>JournalofSubmicroscopicCytologyandPathology</secondary-title><alt-title>JSubmicrCytolPath</alt-title></titles><periodical><full-title>JournalofSubmicroscopicCytologyandPathology</full-title><abbr-1>JSubmicrCytolPath</abbr-1></periodical><alt-periodical><full-title>JournalofSubmicroscopicCytologyandPathology</full-title><abbr-1>JSubmicrCytolPath</abbr-1></alt-periodical><pages>151-155</pages><volume>15</volume><number>1</number><dates><year>1983</year></dates><isbn>1122-9497</isbn><accession-num>WOS:A1983QB17600028</accession-num><urls><related-urls><url><GotoISI>://WOS:A1983QB17600028</url></related-urls></urls><language>English</language></record></Cite></EndNote>(Thurmetal.,1983)。因此，果蝇需要一个类似陀螺仪的结构来保持身体的稳定。翅膀根部的平衡棒对果蝇正是起着类似陀螺仪的作用ADDINEN.CITEADDINEN.CITE.DATA(Liang,2017;Pringle,1948;RochandAkam,2000)。平衡棒上第二节平衡棒茎处大约有100个钟型受体，这些受体在翅膀或者身体发生位移和震动时，可以将刺激信号反馈到中枢神经