【《根细胞类型特异性启动子综述》2000字】

-0-根细胞类型特异性启动子综述在高等植物的不同器官中，初生根是非常优良的发育模型，因为其细胞类型的构造简单且已定型ADDINEN.CITEADDINEN.CITE.DATA(Dolanetal.1993;Marjoleinetal.2005;Sarkaretal.2007)。启动子功能高度依赖于环境，如跨不同组织和细胞类型的基因特异性表达。细胞类型特异性启动子调控是每个细胞独特的转录机制组成部分的功能。因此，为了在特定细胞类型中获得高水平的转录活性，必须专门设计合成启动子以利用这些细胞中离散的可用转录因子库ADDINEN.CITE<EndNote><Cite><Author>Brown</Author><Year>2017</Year><RecNum>82</RecNum><DisplayText>(Brownetal.2017)</DisplayText><record><rec-number>82</rec-number><foreign-keys><keyapp="EN"db-id="2p0s2z5pwfrpauextzhprsswz00ps20w0rxe"timestamp="1609247424">82</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Brown,AdamJ.</author><author>James,DavidC.</author></authors></contributors><titles><title>ConstructingStrongCellType-SpecificPromotersThroughInformedDesign</title></titles><dates><year>2017</year></dates><urls></urls></record></Cite></EndNote>(Brownetal.2017)。多细胞生物由许多细胞类型组成，这些细胞类型通过在时间和空间上精确控制的基因表达模式来得知其特定命运。但是，许多基因的功能通常是通过使用哺乳动物细胞中的巨细胞病毒（CMV）或植物中的花椰菜花叶病毒（CaMV35S）产生的强病毒启动子来产生转基因生物来进行研究的。但虽然这些启动子可以驱动大多数细胞类型中的高表达水平，但阻止了器官特异性或细胞类型特异性作用的分析。要了解高度特异化的细胞类型在整个生物体发育过程中的作用，就需要分离出不同的细胞类型。目前，细胞类型特异性启动子已运用于研究转录组，蛋白质组或分析不同根细胞类型中染色质的变化ADDINEN.CITEADDINEN.CITE.DATA(Birnbaumetal.2003;Bradyetal.2007;Dealetal.2010;Fridmanetal.2014;Kristinaetal.2015;Mustrophetal.2009)，极大地提高了我们对高等植物细胞组织的分辨率。过去二十年来，已经在拟南芥中鉴定出了几种根细胞类型特异性启动子。这些特定的启动子已被用来驱动目的基因在根的离散区域中表达，或响应特定信号和变化来研究其在参与其他表达区域时发挥的作用，或阐明其作为移动信号的功能ADDINEN.CITEADDINEN.CITE.DATA(Duanetal.2013;Fridmanetal.2014;Gonzalez-Garciaetal.2011;Hachametal.2011)。此外，在拟南芥根中已用FRET糖传感器检测到葡萄糖和蔗糖沿浓度梯度的进出口ADDINEN.CITEADDINEN.CITE.DATA(Chaudhurietal.2008)。但是，SUT和其他已知的单糖转运蛋白在相关细胞中表达水平较低ADDINEN.CITE<EndNote><Cite><Author>Slewinski</Author><Year>2011</Year><RecNum>200</RecNum><DisplayText>(Slewinski2011)</DisplayText><record><rec-number>200</rec-number><foreign-keys><keyapp="EN"db-id="2p0s2z5pwfrpauextzhprsswz00ps20w0rxe"timestamp="1609922047">200</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Slewinski,T.L.</author></authors></contributors><auth-address>DepartmentofPlantBiology,CornellUniversity,262PlantScienceBuilding,Ithaca,NY14853,USA.tls98@</auth-address><titles><title>Diversefunctionalrolesofmonosaccharidetransportersandtheirhomologsinvascularplants:aphysiologicalperspective</title><secondary-title>MolPlant</secondary-title><alt-title>Molecularplant</alt-title></titles><alt-periodical><full-title>MolecularPlant</full-title></alt-periodical><pages>641-62</pages><volume>4</volume><number>4</number><edition>2011/07/13</edition><keywords><keyword>GeneExpressionRegulation,Plant</keyword><keyword>MonosaccharideTransportProteins/genetics/*metabolism</keyword><keyword>*MultigeneFamily</keyword><keyword>PlantProteins/genetics/*metabolism</keyword><keyword>Plants/genetics/*metabolism</keyword></keywords><dates><year>2011</year><pub-dates><date>Jul</date></pub-dates></dates><isbn>1674-2052</isbn><accession-num>21746702</accession-num><urls></urls><electronic-resource-num>10.1093/mp/ssr051</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>(Slewinski2011)。1.1PIN2启动子ThepotatoProteinaseInhibitor=2\*ROMANII基因（PIN2）家族是系统性基因激活分析的最适系统ADDINEN.CITEADDINEN.CITE.DATA(Ludevidetal.1990;McElroyetal.1991;Xuetal.1993)。在拟南芥中，PIN基因家族由8个成员组成，研究表明，这些成员参与了各种不同的发育过程ADDINEN.CITEADDINEN.CITE.DATA(Petráseketal.2006;Wangetal.2015)。定位于质膜上的PIN1，PIN2，PIN3，PIN14和PIN7与热带反应和根系生长有关。相比之下，定位于内质网（ER）的PIN包括PIN5，PIN6和PIN8主要参与维持细胞内生长素的稳态ADDINEN.CITEADDINEN.CITE.DATA(Dingetal.2012;Mravecetal.2009)。在所有PIN家庭成员中，功能缺失的pin1和pin2突变体表现出明显的表型差异，因此非常适合研究生长素依赖的发育过程ADDINEN.CITEADDINEN.CITE.DATA(Guoetal.2015)。PIN1主要参与维持胚胎的生长素梯度，并且pin1突变体中器官萌发过程受到严重影响，从而进一步导致形成裸露的花序茎ADDINEN.CITE<EndNote><Cite><Author>Alonso-Peral</Author><Year>2006</Year><RecNum>91</RecNum><DisplayText>(Alonso-Peraletal.2006)</DisplayText><record><rec-number>91</rec-number><foreign-keys><keyapp="EN"db-id="2p0s2z5pwfrpauextzhprsswz00ps20w0rxe"timestamp="1609248777">91</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Alonso-Peral</author><author>M.,M.%JDevelopment</author></authors></contributors><titles><title>TheHVE/CAND1geneisrequiredfortheearlypatterningofleafvenationinArabidopsis</title><secondary-title>Development</secondary-title></titles><periodical><full-title>Development</full-title></periodical><pages>3755</pages><volume>133</volume><number>19</number><dates><year>2006</year></dates><urls></urls></record></Cite></EndNote>(Alonso-Peraletal.2006)。Constitutivetripleresponse1（CTR1），是一种在乙烯受体下游起作用的蛋白质，是乙烯信号的负调节剂。PIN2在CTR1的下游起作用并主要调节中主要负责调节根的向地性ADDINEN.CITE<EndNote><Cite><Author>Alonso-Peral</Author><Year>2006</Year><RecNum>91</RecNum><DisplayText>(Alonso-Peraletal.2006)</DisplayText><record><rec-number>91</rec-number><foreign-keys><keyapp="EN"db-id="2p0s2z5pwfrpauextzhprsswz00ps20w0rxe"timestamp="1609248777">91</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Alonso-Peral</author><author>M.,M.%JDevelopment</author></authors></contributors><titles><title>TheHVE/CAND1geneisrequiredfortheearlypatterningofleafvenationinArabidopsis</title><secondary-title>Development</secondary-title></titles><periodical><full-title>Development</full-title></periodical><pages>3755</pages><volume>133</volume><number>19</number><dates><year>2006</year></dates><urls></urls></record></Cite></EndNote>(Alonso-Peraletal.2006)。有研究表明，PIN基因在激素信号传导和非生物应激反应的整合中起作用。对拟南芥的遗传分析也表明了PIN基因的功能，其中，AtPIN1,AtPIN2,AtPIN3,AtPIN4和AtPIN7参与了根系中细胞间的生长素转运ADDINEN.CITEADDINEN.CITE.DATA(Liuetal.2017)，特别是AtPIN1和AtPIN3被证明是主要的生长素运输促进剂，介导从芽到根尖的极性生长素重新分配ADDINEN.CITEADDINEN.CITE.DATA(Adamowskietal.2015;Blilouetal.2005;Liuetal.2017)。这两个PIN基因和AtPIN2在根中表达量很高，从而决定了分生组织的大小，以及主根的生长速度ADDINEN.CITEADDINEN.CITE.DATA(Blilouetal.2005;Omelyanchuketal.2016)。除了研究伤口信号转导途径，具有编码蛋白酶抑制剂功能和系统伤口诱导能力的PIN2启动子成为潜在的研究昆虫和病原体及转基因植物的抗药性的有效工具ADDINEN.CITE<EndNote><Cite><Author>Ludevid</Author><Year>1990</Year><RecNum>202</RecNum><DisplayText>(Ludevidetal.1990)</DisplayText><record><rec-number>202</rec-number><foreign-keys><keyapp="EN"db-id="2p0s2z5pwfrpauextzhprsswz00ps20w0rxe"timestamp="1609922181">202</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Ludevid,M.D.</author><author>Ruiz-Avila,L.</author><author>Vallés,M.P.</author><author>Stiefel,V.</author><author>Torrent,M.</author><author>Torné,J.M.</author><author>Puigdomènech,P.</author></authors></contributors><auth-address>DepartamentodeGenéticaMolecular,CentrodeInvestigaciónyDesarrollo,CSIC,JordiGirona,18,E-08034,Barcelona,Spain.</auth-address><titles><title>Expressionofgenesforcell-wallproteinsindividingandwoundedtissuesofZeamaysL</title><secondary-title>Planta</secondary-title><alt-title>Planta</alt-title></titles><periodical><full-title>Planta</full-title><abbr-1>Planta</abbr-1></periodical><alt-periodical><full-title>Planta</full-title><abbr-1>Planta</abbr-1></alt-periodical><pages>524-9</pages><volume>180</volume><number>4</number><edition>1990/03/01</edition><dates><year>1990</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0032-0935(Print) 0032-0935</isbn><accession-num>24202097</accession-num><urls></urls><electronic-resource-num>10.1007/bf02411450</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>(Ludevidetal.1990)。PIN2启动子片段在整个根部具有广泛活性，即不仅在根分生组织中而且在根的分化部分中，以及表皮和皮质中均有活性。Sieberer的研究中，以GUS为报告基因对PIN2启动子及PIN2蛋白表达域进行了比较分析，发现PIN2启动子本身并不具有组织特异性ADDINEN.CITEADDINEN.CITE.DATA(Siebereretal.2000)。通过对PIN2启动子活性的分析表明，它对表皮和皮层细胞具有特异性，而蛋白质的稳定性可能限制了PIN2蛋白对这些组织的分生组织的作用。此外，研究还发现PIN2启动子在侧根帽中活跃，而在小柱中不活跃。1.2WER启动子拟南芥（Arabidopsisthaliana）根表皮中的根毛和非毛细胞由一组转录调节因子编码。其中的两个分别是WEREWOLF（WER）和CAPRICE（CPC），WEREWOLF（WER）是拟南芥中表皮细胞模式的调节器。因此，WER是根和下胚轴（而不是叶）表皮细胞模式形成以及同源基因GL2正确表达所必需的。此外，WER蛋白在玉米R基因编码的bHLH蛋白的下游或与其平行作用，以影响GL2表达和表皮发育，并与CPC相对ADDINEN.CITE<EndNote><Cite><Author>Lee</Author><Year>1999</Year><RecNum>98</RecNum><DisplayText>(Leeetal.1999)</DisplayText><record><rec-number>98</rec-number><foreign-keys><keyapp="EN"db-id="2p0s2z5pwfrpauextzhprsswz00ps20w0rxe"timestamp="1609250118">98</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Lee,M.M.</author><author>Schiefelbein,J.</author></authors></contributors><auth-address>UnivMichigan,DeptBiol,AnnArbor,MI48109USA</auth-address><titles><title>WEREWOLF,aMYB-relatedproteininarabidopsis,isaposition-dependentregulatorofepidermalcellpatterning</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>473-483</pages><volume>99</volume><number>5</number><keywords><keyword>roothairdevelopment</keyword><keyword>geneencodes</keyword><keyword>thaliana</keyword><keyword>differentiation</keyword><keyword>plants</keyword><keyword>sequence</keyword><keyword>glabra2</keyword><keyword>family</keyword><keyword>leaves</keyword><keyword>fate</keyword></keywords><dates><year>1999</year><pub-dates><date>Nov24</date></pub-dates></dates><isbn>0092-8674</isbn><accession-num>WOS:000083986300007</accession-num><urls><related-urls><url><GotoISI>://WOS:000083986300007</url></related-urls></urls><electronic-resource-num>Doi10.1016/S0092-8674(00)81536-6</electronic-resource-num><language>English</language></record></Cite></EndNote>(Leeetal.1999)。而且，WER与在拟南芥中寻找MYB基因时发现的几个不完整序列相同ADDINEN.CITEADDINEN.CITE.DATA(Kranzetal.1998;Oppenheimeretal.1991;Romeroetal.1998)。这些发现表明，WER是拟南芥根表皮中细胞类型模式的关键决定因素ADDINEN.CITE<EndNote><Cite><Author>Lee</Author><Year>1999</Year><RecNum>98</RecNum><DisplayText>(Leeetal.1999)</DisplayText><record><rec-number>98</rec-number><foreign-keys><keyapp="EN"db-id="2p0s2z5pwfrpauextzhprsswz00ps20w0rxe"timestamp="1609250118">98</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Lee,M.M.</author><author>Schiefelbein,J.</author></authors></contributors><auth-address>UnivMichigan,DeptBiol,AnnArbor,MI48109USA</auth-address><titles><title>WEREWOLF,aMYB-relatedproteininarabidopsis,isaposition-dependentregulatorofepidermalcellpatterning</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>473-483</pages><volume>99</volume><number>5</number><keywords><keyword>roothairdevelopment</keyword><keyword>geneencodes</keyword><keyword>thaliana</keyword><keyword>differentiation</keyword><keyword>plants</keyword><keyword>sequence</keyword><keyword>glabra2</keyword><keyword>family</keyword><keyword>leaves</keyword><keyword>fate</keyword></keywords><dates><year>1999</year><pub-dates><date>Nov24</date></pub-dates></dates><isbn>0092-8674</isbn><accession-num>WOS:000083986300007</accession-num><urls><related-urls><url><GotoISI>://WOS:000083986300007</url></related-urls></urls><electronic-resource-num>Doi10.1016/S0092-8674(00)81536-6</electronic-resource-num><language>English</language></record></Cite></EndNote>(Leeetal.1999)。WER和CPC分别编码R2R3类的MYB转录因子，分别促进非毛细胞命运和毛细胞命运。已有研究发现拟南芥根尖中的WER过表达足以直接诱导GLABRA2（GL2）基因表达而使表皮细胞采用非毛细胞命运。该研究还表明，GLABRA3（GL3）和GLABRA3（EGL3）的增强子是过表达WER的作用所必需的，并且WER与这些bHLHs在植物细胞中相互作用。此外，他们还发现CPC抑制了WER过表达的表型。这些结果表明，WER通过与GL3/EGL3共同作用以诱导GL2的表达，并且WER和CPC相互竞争以定义拟南芥根表皮中的细胞命运ADDINEN.CITE<EndNote><Cite><Author>Song</Author><Year>2011</Year><RecNum>99</RecNum><DisplayText>(Songetal.2011)</DisplayText><record><rec-number>99</rec-number><foreign-keys><keyapp="EN"db-id="2p0s2z5pwfrpauextzhprsswz00ps20w0rxe"timestamp="1609250215">99</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Song,S.K.</author><author>Ryu,K.H.</author><author>Kang,Y.H.</author><author>Song,J.H.</author><author>Cho,Y.H.</author><author>Yoo,S.D.</author><author>Schiefelbein,J.</author><author>Lee,M.M.</author></authors></contributors><auth-address>YonseiUniv,DeptSystBiol,Seoul120749,SouthKorea SungkyunkwanUniv,DeptBiolSci,Suwon440746,SouthKorea UnivMichigan,DeptMolCellular&DevBiol,AnnArbor,MI48109USA</auth-address><titles><title>CellFateintheArabidopsisRootEpidermisIsDeterminedbyCompetitionbetweenWEREWOLFandCAPRICE</title><secondary-title>PlantPhysiology</secondary-title><alt-title>PlantPhysiol</alt-title></titles><periodical><full-title>PlantPhysiology</full-title></periodical><alt-periodical><full-title>PlantPhysiol</full-title></alt-periodical><pages>1196-1208</pages><volume>157</volume><number>3</number><keywords><keyword>receptor-likekinase</keyword><keyword>trichomedevelopment</keyword><keyword>lateralinhibition</keyword><keyword>hairdevelopment</keyword><keyword>gene-expression</keyword><keyword>mybprotein</keyword><keyword>differentiation</keyword><keyword>thaliana</keyword><keyword>position</keyword><keyword>gl3</keyword></keywords><dates><year>2011</year><pub-dates><date>Nov</date></pub-dates></dates><isbn>0032-0889</isbn><accession-num>WOS:000296722300019</accession-num><urls><related-urls><url><GotoISI>://WOS:000296722300019</url></related-urls></urls><electronic-resource-num>10.1104/pp.111.185785</electronic-resource-num><language>English</language></record></Cite></EndNote>(Songetal.2011)，它们为整个拟南芥植物中表皮发育的调控提供了新的方向。因此，综上所述，WER是毛状细胞特异性的启动子，它对于驱动无根毛或有根毛的根表皮细胞中基因表达有显著作用ADDINEN.CITEADDINEN.CITE.DATA(Bruexetal.2012;Leeetal.1999)，它可以导致根毛在几乎每个根表皮上形成。图1.4PIN2和WER启动子的表达模式ADDINEN.CITE<EndNote><Cite><Author>Maria</Author><Year>2016</Year><RecNum>102</RecNum><DisplayText>(Mariaetal.2016)</DisplayText><record><rec-number>102</rec-number><foreign-keys><keyapp="EN"db-id="2p0s2z5pwfrpauextzhprsswz00ps20w0rxe"timestamp="1609250549">102</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Maria</author><author>Mar</author><author>Marquès-Bueno</author><author>Ana</author><author>K.</author><author>Morao</author><author>Anne</author><author>Cayrel</au