PCP通路调节平面细胞极性的建立
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摘要
平面细胞极性(planar cell polarity,PCP)的建立是胚胎发育过程中的一个关键环节。PCP通路在进化上高度保守,包括"核心"PCP通路和"Ds/Ft"PCP通路,两者共同调节平面细胞极性的建立,导致细胞发生汇聚延伸(convergent extension,CE)运动和定向的细胞分裂(oriented cell division,OCD)。PCP通路的正确激活以及正常的细胞结构需要PCP蛋白的不对称性分布,由Wnt蛋白和Ft-Ds-Fj系统提供极性信号,激活下游效应器而产生。本文综述了PCP通路在平面细胞极性建立过程中的作用及分子机制,旨在为胚胎发育相关疾病积累理论基础。
The establishment of planar cell polarity(PCP) is a key process during the development of the embryo.The PCP pathways are highly conserved in evolution,which include the "core" PCP pathway and the "Ds/Ft" PCP pathway,both of which regulate the establishment of planar cell polarity,and lead to cell occurrence convergent extension(CE) movements and oriented cell division(OCD).The correct activation of the PCP pathways and the normal cellular structure require the asymmetry distribution of the PCP protein,which is depended on the Wnt protein and Ft-Ds-Fj systems to provide the polarity information and to activate the downstream effector.In this paper,the role and molecular mechanism of PCP pathways during the establishment of cell polarity are reviewed.The main aim is to provide a theoretical basis for the advance of embryonic development related diseases.
The establishment of planar cell polarity(PCP) is a key process during the development of the embryo.The PCP pathways are highly conserved in evolution,which include the "core" PCP pathway and the "Ds/Ft" PCP pathway,both of which regulate the establishment of planar cell polarity,and lead to cell occurrence convergent extension(CE) movements and oriented cell division(OCD).The correct activation of the PCP pathways and the normal cellular structure require the asymmetry distribution of the PCP protein,which is depended on the Wnt protein and Ft-Ds-Fj systems to provide the polarity information and to activate the downstream effector.In this paper,the role and molecular mechanism of PCP pathways during the establishment of cell polarity are reviewed.The main aim is to provide a theoretical basis for the advance of embryonic development related diseases.
引文
[1]Chen H,Cheng CY.Planar cell polarity(PCP)proteins and spermatogenesis.Semin Cell Dev Biol,2016,59:99-109
[2]Goodrich LV,Strutt D.Principles of planar polarity in animal development.Development,2011,138(10):1877-1892
[3]Castanon I,Gonzalez-Gaitan M.Oriented cell division in vertebrate embryogenesis.Curr Opin Cell Biol,2011,23(6):697-704
[4]Singh J,Mlodzik M.Planar cell polarity signaling:coordination of cellular orientation across tissues.Wiley Interdiscip Rev Dev Biol,2012,1(4):479-499
[5]Adler PN.The Frizzled/Stan pathway and planar cell polarity in the Drosophila wing.Curr Top Dev Biol,2012,101:1-31
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[7]Bastock R.Strabismus is asymmetrically localised and binds to Prickle and Dishevelled during Drosophila planar polarity patterning.Development,2003,130(13):3007-3014
[8]Nishimura T,Honda H,Takeichi M.Planar cell polarity links axes of spatial dynamics in neural-tube closure.Cell,2012,149(5):1084-1097
[9]Martinez S,Scerbo P,Giordano M,et al.The Ptk7 and Ror2 protein receptors interact in the Vertebrate Wnt/planar cell Polarity(PCP)pathway.J Biol Chem,2015,290(51):30562-30572
[10]Yamanaka H,Moriguchi T,Masuyama N,et al.JNK functions in the non-canonical Wnt pathway to regulate convergent extension movements in vertebrates.EMBO Rep,2002,3(1):69-75
[11]Jung B,Kohler A,Schambony A,et al.PAPC and the Wnt5a/Ror2 pathway control the invagination of the otic placode in Xenopus.BMC Dev Biol,2011,11:36
[12]Levayer R,Lecuit T.Biomechanical regulation of contractility:spatial control and dynamics.Trends Cell Biol,2012,22(2):61-81
[13]Strutt H,Searle E,Thomas-Macarthur V,et al.A Cul-3-BTB ubiquitylation pathway regulates junctional levels and asymmetry of core planar polarity proteins.Development,2013,140(8):1693-1702
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[15]Brittle A,Thomas C,Strutt D.Planar polarity specification through asymmetric subcellular localization of Fat and Dachsous.Curr Biol,2012,22(10):907-914
[16]Staley BK,Irvine KD.Hippo signaling in Drosophila:recent advances and insights.Dev Dyn,2012,241(1):3-15
[17]Mao Y,Rauskolb C,Cho E,et al.Dachs:an unconventional myosin that functions downstream of Fat to regulate growth,affinity and gene expression in Drosophila.Development,2006,133(13):2539-2551
[18]Fanto M.The tumor-suppressor and cell adhesion molecule Fat controls planar polarity via physical interactions with Atrophin,a transcriptional co-repressor.Development,2003,130(4):763-774
[19]Milton CC,Zhang X,Albanese NO,et al.Differential requirement of Salvador-Warts-Hippo pathway members for organ size control in Drosophila melanogaster.Development,2010,137(5):735-743
[20]Sharma P and Mc Neill H.Regulation of long-range planar cell polarity by Fat-Dachsous signaling.Development,2013,140(18):3869-3881
[21]Wu J,Roman AC,Carvajal-Gonzalez JM,et al.Wg and Wnt4 provide long-range directional input to planar cell polarity orientation in Drosophila.Nat Cell Biol,2013,15(9):1045-1055
[22]Kelly LK,Wu J,Yanfeng WA,et al.Frizzled-induced Van Gogh phosphorylation by CK1epsilon promotes asymmetric localization of Core PCP factors in Drosophila.Cell Rep,2016,16(2):344-356
[23]Gao B,Song H,Bishop K,et al.Wnt signaling gradients establish planar cell polarity by inducing Vangl2 phosphorylation through Ror2.Dev Cell,2011,20(2):163-176
[24]Matis M,Russler-Germain DA,Hu Q,et al.Microtubules provide directional information for core PCP function.Elife,2014,3:e02893
[25]Brittle AL,Repiso A,Casal J,et al.Four-jointed modulates growth and planar polarity by reducing the affinity of Dachsous for Fat.Curr Biol,2010,20(9):803-810
[26]Harumoto T,Ito M,Shimada Y,et al.Atypical cadherins Dachsous and Fat control dynamics of noncentrosomal microtubules in planar cell polarity.Dev Cell,2010,19(3):389-401
[27]Casal J,Lawrence PA,Struhl G.Two separate molecular systems,Dachsous/Fat and Starry night/Frizzled,act independently to confer planar cell polarity.Development,2006,133(22):4561-4572
[28]Wang Y,Guo N,Nathans J.The role of Frizzled3 and Frizzled6 in neural tube closure and in the planar polarity of inner-ear sensory hair cells.J Neurosci,2006,26(8):2147-2156
[29]Schulte G and Bryja V.The Frizzled family of unconventional G-protein-coupled receptors.Trends Pharmacol Sci,2007,28(10):518-525
[30]Takeichi M.The cadherin superfamily in neuronal connections and interactions.Nat Rev Neurosci,2007,8(1):11-20
[31]Lei Y,Zhu H,Yang W,et al.Identification of novel Celsr1 mutations in spina bifida.PLo S One,2014,9(3):e92207
[32]Katoh Y,Katoh M.Comparative genomics on Vangl1and Vangl2 genes.Int J Oncol,2005,26(5):1435-1440
[33]Capelluto DG,Kutateladze TG,Habas R,et al.The DIX domain targets dishevelled to actin stress fibres and vesicular membranes.Nature,2002,419(6908):726-729
[34]Liu C,Lin C,Gao C,et al.Null and hypomorph Prickle1alleles in mice phenocopy human Robinow syndrome and disrupt signaling downstream of Wnt5a.Biol Open,2014,3(9):861-870
[35]Carreira-Barbosa F,Concha ML,Takeuchi M,et al.Prickle1regulates cell movements during gastrulation and neuronal migration in Zebrafish.Development,2003,130(17):4037-4046
[36]Das G,Jenny A,Klein TJ,et al.Diego interacts with Prickle and Strabismus/Van Gogh to localize planar cell polarity complexes.Development,2004,131(18):4467-4476
[37]Chen WS,Antic D,Matis M,et al.Asymmetric homotypic interactions of the atypical cadherin flamingo mediate intercellular polarity signaling.Cell,2008,133(6):1093-1105
[38]Vladar EK,Bayly RD,Sangoram AM,et al.Microtubules enable the planar cell polarity of airway cilia.Curr Biol,2012,22(23):2203-2212
[39]Sepich DS,Usmani M,Pawlicki S,et al.Wnt/PCP signaling controls intracellular position of MTOCs during gastrulation convergence and extension movements.Development,2011,138(3):543-552
[40]Olofsson J,Sharp KA,Matis M,et al.Prickle/spiny-legs isoforms control the polarity of the apical microtubule network in planar cell polarity.Development,2014,141(14):2866-2874
[41]Guo Y,Zanetti G,Schekman R.A novel GTP-binding protein-adaptor protein complex responsible for export of Vangl2 from the trans Golgi network.Elife,2013,2:e00160
[42]Karner CM,Chirumamilla R,Aoki S,et al.Wnt9b signaling regulates planar cell polarity and kidney tubule morphogenesis.Nat Genet,2009,41(7):793-799
[43]Randall RM,Shao YY,Wang L,et al.Activation of Wnt planar cell polarity(PCP)signaling promotes growth plate column formation in vitro.J Orthop Res,2012,30(12):1906-1914
[44]Gibbs BC,Damerla RR,Vladar EK,et al.Prickle1 mutation causes planar cell polarity and directional cell migration defects associated with cardiac outflow tract anomalies and other structural birth defects.Biol Open,2016,5(3):323-335
[45]Puvirajesinghe TM,Bertucci F,Jain A,et al.Identification of p62/SQSTM1 as a component of non-canonical Wnt Vangl2-JNK signalling in breast cancer.Nat Commun,2016,7:10318
[46]Copp AJ,Greene ND,Murdoch JN.The genetic basis of mammalian neurulation.Nat Rev Genet,2003,4(10):784-793
[47]Yamaguchi Y,Miura M.How to form and close the brain:insight into the mechanism of cranial neural tube closure in mammals.Cell Mol Life Sci,2013,70(17):3171-3186
[48]Greene ND,Copp AJ.Neural tube defects.Annu Rev Neurosci,2014,37:221-242
[49]Murdoch JN,Damrau C,Paudyal A,et al.Genetic interactions between planar cell polarity genes cause diverse neural tube defects in mice.Dis Model Mech,2014,7(10):1153-1163
[50]Badouel C,Zander MA,Liscio N,et al.Fat1 interacts with Fat4 to regulate neural tube closure,neural progenitor proliferation and apical constriction during mouse brain development.Development,2015,142(16):2781-2791
[2]Goodrich LV,Strutt D.Principles of planar polarity in animal development.Development,2011,138(10):1877-1892
[3]Castanon I,Gonzalez-Gaitan M.Oriented cell division in vertebrate embryogenesis.Curr Opin Cell Biol,2011,23(6):697-704
[4]Singh J,Mlodzik M.Planar cell polarity signaling:coordination of cellular orientation across tissues.Wiley Interdiscip Rev Dev Biol,2012,1(4):479-499
[5]Adler PN.The Frizzled/Stan pathway and planar cell polarity in the Drosophila wing.Curr Top Dev Biol,2012,101:1-31
[6]Narimatsu M,Bose R,Pye M,et al.Regulation of planar cell polarity by Smurf ubiquitin ligases.Cell,2009,137(2):295-307
[7]Bastock R.Strabismus is asymmetrically localised and binds to Prickle and Dishevelled during Drosophila planar polarity patterning.Development,2003,130(13):3007-3014
[8]Nishimura T,Honda H,Takeichi M.Planar cell polarity links axes of spatial dynamics in neural-tube closure.Cell,2012,149(5):1084-1097
[9]Martinez S,Scerbo P,Giordano M,et al.The Ptk7 and Ror2 protein receptors interact in the Vertebrate Wnt/planar cell Polarity(PCP)pathway.J Biol Chem,2015,290(51):30562-30572
[10]Yamanaka H,Moriguchi T,Masuyama N,et al.JNK functions in the non-canonical Wnt pathway to regulate convergent extension movements in vertebrates.EMBO Rep,2002,3(1):69-75
[11]Jung B,Kohler A,Schambony A,et al.PAPC and the Wnt5a/Ror2 pathway control the invagination of the otic placode in Xenopus.BMC Dev Biol,2011,11:36
[12]Levayer R,Lecuit T.Biomechanical regulation of contractility:spatial control and dynamics.Trends Cell Biol,2012,22(2):61-81
[13]Strutt H,Searle E,Thomas-Macarthur V,et al.A Cul-3-BTB ubiquitylation pathway regulates junctional levels and asymmetry of core planar polarity proteins.Development,2013,140(8):1693-1702
[14]Matis M,Axelrod JD.Regulation of PCP by the Fat signaling pathway.Genes Dev,2013,27(20):2207-2220
[15]Brittle A,Thomas C,Strutt D.Planar polarity specification through asymmetric subcellular localization of Fat and Dachsous.Curr Biol,2012,22(10):907-914
[16]Staley BK,Irvine KD.Hippo signaling in Drosophila:recent advances and insights.Dev Dyn,2012,241(1):3-15
[17]Mao Y,Rauskolb C,Cho E,et al.Dachs:an unconventional myosin that functions downstream of Fat to regulate growth,affinity and gene expression in Drosophila.Development,2006,133(13):2539-2551
[18]Fanto M.The tumor-suppressor and cell adhesion molecule Fat controls planar polarity via physical interactions with Atrophin,a transcriptional co-repressor.Development,2003,130(4):763-774
[19]Milton CC,Zhang X,Albanese NO,et al.Differential requirement of Salvador-Warts-Hippo pathway members for organ size control in Drosophila melanogaster.Development,2010,137(5):735-743
[20]Sharma P and Mc Neill H.Regulation of long-range planar cell polarity by Fat-Dachsous signaling.Development,2013,140(18):3869-3881
[21]Wu J,Roman AC,Carvajal-Gonzalez JM,et al.Wg and Wnt4 provide long-range directional input to planar cell polarity orientation in Drosophila.Nat Cell Biol,2013,15(9):1045-1055
[22]Kelly LK,Wu J,Yanfeng WA,et al.Frizzled-induced Van Gogh phosphorylation by CK1epsilon promotes asymmetric localization of Core PCP factors in Drosophila.Cell Rep,2016,16(2):344-356
[23]Gao B,Song H,Bishop K,et al.Wnt signaling gradients establish planar cell polarity by inducing Vangl2 phosphorylation through Ror2.Dev Cell,2011,20(2):163-176
[24]Matis M,Russler-Germain DA,Hu Q,et al.Microtubules provide directional information for core PCP function.Elife,2014,3:e02893
[25]Brittle AL,Repiso A,Casal J,et al.Four-jointed modulates growth and planar polarity by reducing the affinity of Dachsous for Fat.Curr Biol,2010,20(9):803-810
[26]Harumoto T,Ito M,Shimada Y,et al.Atypical cadherins Dachsous and Fat control dynamics of noncentrosomal microtubules in planar cell polarity.Dev Cell,2010,19(3):389-401
[27]Casal J,Lawrence PA,Struhl G.Two separate molecular systems,Dachsous/Fat and Starry night/Frizzled,act independently to confer planar cell polarity.Development,2006,133(22):4561-4572
[28]Wang Y,Guo N,Nathans J.The role of Frizzled3 and Frizzled6 in neural tube closure and in the planar polarity of inner-ear sensory hair cells.J Neurosci,2006,26(8):2147-2156
[29]Schulte G and Bryja V.The Frizzled family of unconventional G-protein-coupled receptors.Trends Pharmacol Sci,2007,28(10):518-525
[30]Takeichi M.The cadherin superfamily in neuronal connections and interactions.Nat Rev Neurosci,2007,8(1):11-20
[31]Lei Y,Zhu H,Yang W,et al.Identification of novel Celsr1 mutations in spina bifida.PLo S One,2014,9(3):e92207
[32]Katoh Y,Katoh M.Comparative genomics on Vangl1and Vangl2 genes.Int J Oncol,2005,26(5):1435-1440
[33]Capelluto DG,Kutateladze TG,Habas R,et al.The DIX domain targets dishevelled to actin stress fibres and vesicular membranes.Nature,2002,419(6908):726-729
[34]Liu C,Lin C,Gao C,et al.Null and hypomorph Prickle1alleles in mice phenocopy human Robinow syndrome and disrupt signaling downstream of Wnt5a.Biol Open,2014,3(9):861-870
[35]Carreira-Barbosa F,Concha ML,Takeuchi M,et al.Prickle1regulates cell movements during gastrulation and neuronal migration in Zebrafish.Development,2003,130(17):4037-4046
[36]Das G,Jenny A,Klein TJ,et al.Diego interacts with Prickle and Strabismus/Van Gogh to localize planar cell polarity complexes.Development,2004,131(18):4467-4476
[37]Chen WS,Antic D,Matis M,et al.Asymmetric homotypic interactions of the atypical cadherin flamingo mediate intercellular polarity signaling.Cell,2008,133(6):1093-1105
[38]Vladar EK,Bayly RD,Sangoram AM,et al.Microtubules enable the planar cell polarity of airway cilia.Curr Biol,2012,22(23):2203-2212
[39]Sepich DS,Usmani M,Pawlicki S,et al.Wnt/PCP signaling controls intracellular position of MTOCs during gastrulation convergence and extension movements.Development,2011,138(3):543-552
[40]Olofsson J,Sharp KA,Matis M,et al.Prickle/spiny-legs isoforms control the polarity of the apical microtubule network in planar cell polarity.Development,2014,141(14):2866-2874
[41]Guo Y,Zanetti G,Schekman R.A novel GTP-binding protein-adaptor protein complex responsible for export of Vangl2 from the trans Golgi network.Elife,2013,2:e00160
[42]Karner CM,Chirumamilla R,Aoki S,et al.Wnt9b signaling regulates planar cell polarity and kidney tubule morphogenesis.Nat Genet,2009,41(7):793-799
[43]Randall RM,Shao YY,Wang L,et al.Activation of Wnt planar cell polarity(PCP)signaling promotes growth plate column formation in vitro.J Orthop Res,2012,30(12):1906-1914
[44]Gibbs BC,Damerla RR,Vladar EK,et al.Prickle1 mutation causes planar cell polarity and directional cell migration defects associated with cardiac outflow tract anomalies and other structural birth defects.Biol Open,2016,5(3):323-335
[45]Puvirajesinghe TM,Bertucci F,Jain A,et al.Identification of p62/SQSTM1 as a component of non-canonical Wnt Vangl2-JNK signalling in breast cancer.Nat Commun,2016,7:10318
[46]Copp AJ,Greene ND,Murdoch JN.The genetic basis of mammalian neurulation.Nat Rev Genet,2003,4(10):784-793
[47]Yamaguchi Y,Miura M.How to form and close the brain:insight into the mechanism of cranial neural tube closure in mammals.Cell Mol Life Sci,2013,70(17):3171-3186
[48]Greene ND,Copp AJ.Neural tube defects.Annu Rev Neurosci,2014,37:221-242
[49]Murdoch JN,Damrau C,Paudyal A,et al.Genetic interactions between planar cell polarity genes cause diverse neural tube defects in mice.Dis Model Mech,2014,7(10):1153-1163
[50]Badouel C,Zander MA,Liscio N,et al.Fat1 interacts with Fat4 to regulate neural tube closure,neural progenitor proliferation and apical constriction during mouse brain development.Development,2015,142(16):2781-2791