哺乳动物着床前胚胎中细胞系的分化及调控机制
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摘要
哺乳动物着床前发育是指精卵结合后胚胎在子宫着床之前的发育。这一阶段的发育时间短(小鼠为4.5 d),但是发生了一系列形态学和发育生物学上的变化。形态上,小鼠在着床前胚胎经历了3个重要变化:8细胞阶段的致密化和极性化、内细胞团和滋养层的分化及原始内胚层和外胚层的分离;在发育生物学方面,受精后会发生染色质重编程、母源因子降解、胚胎基因组激活等事件。本文以小鼠着床前胚胎为例,重点介绍了影响胚胎形态变化的调控机制及参与早期胚胎细胞分化的信号通路。对这些调控机制的理解有助于开发相关技术,从而提高动物繁殖效率或哺乳动物辅助生殖效率。
Preimplantation development refers to the early embryonic development between fertilization and implantation. During this short developmental window(4.5 days of mouse), a series of morphological and developmental changes take place. In morphology, three important changes occurred during preimplantation development in mouse embryos: compaction and polarity at 8-cell stage; inner cell mass and trophectoderm differentiation at early blastocyst stage; epiblast and primitive endoderm segregation at late blastocyst stage. In developmental biology, chromatin reprogramming, degradation of maternal factor, and embryo genomic activation take place after fertilization. Taken preimplantation embryos of mouse as an example, the present review aims to highlight the regulatory mechanisms responsible for the change of embryo morphology, and related signaling pathways of early embryonic cell differentiation, which are essential for the lineage specification. Understanding the fundamental mechanisms underlying the preimplantation development would greatly enhance our capability to improve reproductive performance of domestic animals or increase the efficiency of assisted reproductive technologies in mammals.
Preimplantation development refers to the early embryonic development between fertilization and implantation. During this short developmental window(4.5 days of mouse), a series of morphological and developmental changes take place. In morphology, three important changes occurred during preimplantation development in mouse embryos: compaction and polarity at 8-cell stage; inner cell mass and trophectoderm differentiation at early blastocyst stage; epiblast and primitive endoderm segregation at late blastocyst stage. In developmental biology, chromatin reprogramming, degradation of maternal factor, and embryo genomic activation take place after fertilization. Taken preimplantation embryos of mouse as an example, the present review aims to highlight the regulatory mechanisms responsible for the change of embryo morphology, and related signaling pathways of early embryonic cell differentiation, which are essential for the lineage specification. Understanding the fundamental mechanisms underlying the preimplantation development would greatly enhance our capability to improve reproductive performance of domestic animals or increase the efficiency of assisted reproductive technologies in mammals.
引文
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[27]LORTHONGPANICH C,MESSERSCHMIDT D M,CHANS W,et al.Temporal reduction of LATS kinases in the early preimplantation embryo prevents ICM lineage differentiation.Genes&Development,2013,27(13):1441-1446.
[28]COCKBURN K,BIECHELE S,GARNER J,et al.The Hippo pathway member Nf2 is required for inner cell mass specification.Current Biology,2013,23(13):1195-1201.
[29]HIRATE Y,HIRAHARA S,INOUE K,et al.Polaritydependent distribution of angiomotin localizes Hippo signaling in preimplantation embryos.Current Biology,2013,23(13):1181-1194.
[30]WICKLOW E,BLIJ S,FRUM T.HIPPO pathway members restrict SOX2 to the inner cell mass where it promotes ICMfates in the mouse blastocyst.PLo S Genetics,2014,10(10):e1004618.
[31]DE LAS HERAS J M,CASANOVA J.Spatially distinct downregulation of Capicua repression and tailless activation by the Torso RTK pathway in the Drosophila embryo.Mechanisms of Development,2006,123(6):481-486.
[32]CHAZAUD C,YAMANAKA Y,PAWSON T,et al.Early lineage segregation between epiblast and primitive endoderm in mouse blastocysts through the Grb2-MAPKpathway.Developmental Cell,2006,10(5):615-624.
[33]MCKAY M M,MORRISON D K.Integrating signals from RTKs to ERK/MAPK.Oncogene,2007,26(22):3113-3121.
[34]SCHRODE N,SAIZ N,DI TALIA S,et al.GATA6 levels modulate primitive endoderm cell fate choice and timing in the mouse blastocyst.Developmental Cell,2014,29(4):454-467.
[35]TAKAOKA K,HAMADA H.Cell fate decisions and axis determination in the early mouse embryo.Development,2012,139(1):3-14.
[36]KOCH U,LEHAL R,RADTKE F.Stem cells living with a Notch.Development,2013,140(4):689-704.
[37]WANG H F,ZANG C Z,LIU X S,et al.The role of Notch receptors in transcriptional regulation.Journal of Cellular Physiology,2015,230(5):982-988.
[38]RAYON T,MENCHERO S,NIETO A,et al.Notch and Hippo converge on Cdx2 to specify the trophectoderm lineage in the mouse blastocyst.Developmental Cell,2014,30(4):410-422.
[2]MIHAJLOVIC A I,BRUCE A W.The first cell-fate decision of mouse preimplantation embryo development:Integrating cell position and polarity.Open Biology,2017,7(11):170210.
[3]JOHNSON M H,MCCONNELL J M L.Lineage allocation and cell polarity during mouse embryogenesis.Seminars in Cell&Developmental Biology,2004,15(5):583-597.
[4]CHAZAUD C,YAMANAKA Y.Lineage specification in the mouse preimplantation embryo.Development,2016,143(7):1063-1074.
[5]DE VRIES W N,EVSIKOV A V,HAAC B E,et al.Maternalβ-catenin and E-cadherin in mouse development.Development,2004,131(18):4435-4445.
[6]LIU G Y,ZHANG X,LIN H Y,et al.Effects of E-cadherin on mouse embryo implantation and expression of matrix metalloproteinase-2 and-9.Biochemical and Biophysical Research Communications,2006,343(3):832-838.
[7]REEVE W J D,ZIOMEK C A.Distribution of microvilli on dissociated blastomeres from mouse embryos:Evidence for surface polarization at compaction.Development,1981,62(1):339-350.
[8]LOUVET S,AGHION J,SANTA-MARIA A,et al.Ezrin becomes restricted to outer cells following asymmetrical division in the preimplantation mouse embryo.Developmental Biology,1996,177(2):568-579.
[9]VINOT S,LE T,OHNO S,et al.Asymmetric distribution of PAR proteins in the mouse embryo begins at the 8-cell stage during compaction.Developmental Biology,2005,282(2):307-319.
[10]ALARCON V B.Cell polarity regulator PARD6B is essential for trophectoderm formation in the preimplantation mouse embryo.Biology of Reproduction,2010,83(3):347-358.
[11]YAMANAKA Y,RALSTON A,STEPHENSON R O,et al.Cell and molecular regulation of the mouse blastocyst.Developmental Dynamics,2006,235(9):2301-2314.
[12]WILEY L M.Cavitation in the mouse preimplantation embryo:Na/K-ATPase and the origin of nascent blastocoele fluid.Developmental Biology,1984,105(2):330-342.
[13]TARKOWSKI A K,WROBLEWSKA J.Development of blastomeres of mouse eggs isolated at the 4-and 8-cell stage.Journal of Embryology and Experimental Morphology,1967,18(1):155-180.
[14]JOHNSON M H,ZIOMEK C A.The foundation of two distinct cell lineages within the mouse morula.Cell,1981,24(1):71-80.
[15]SAMARAGE C R,WHITE M D,áLVAREZ Y D,et al.Cortical tension allocates the first inner cells of the mammalian embryo.Developmental Cell,2015,34(4):435-447.
[16]STRUMPF D,MAO C A,YAMANAKA Y,et al.Cdx2 is required for correct cell fate specification and differentiation of trophectoderm in the mouse blastocyst.Development,2005,132(9):2093-2102.
[17]GOOLAM M,SCIALDONE A,GRAHAM S J L,et al.Heterogeneity in Oct4 and Sox2 targets biases cell fate in 4-cell mouse embryos.Cell Report,2016,165(1):61-74.
[18]CHEN L Y,YABUUCHI A,EMINLI S,et al.Crossregulation of the Nanog and Cdx2 promoters.Cell Research,2009,19(9):1052-1061.
[19]NIWA H,TOYOOKA Y,SHIMOSATO D,et al.Interaction between Oct3/4 and Cdx2 determines trophectoderm differentiation.Cell,2005,123(5):917-929.
[20]YUAN P,HAN J Y,GUO G J,et al.Eset partners with Oct4to restrict extraembryonic trophoblast lineage potential in embryonic stem cells.Genes&Development,2009,23(21):2507-2520.
[21]ZHANG K,HAVERSAT J M,MAGER J.CTR9/PAF1c regulates molecular lineage identity,histone H3K36trimethylation and genomic imprinting during preimplantation development.Developmental Biology,2013,383(1):15-27.
[22]KANG M J,GARG V,HADJANTONAKIS A K.Lineage establishment and progression within the inner cell mass of the mouse blastocyst requires FGFR1 and FGFR2.Developmental Cell,2017,41(5):496-510.e5.
[23]KANG M J,PILISZEK A,ARTUS J,et al.FGF4 is required for lineage restriction and salt-and-pepper distribution of primitive endoderm factors but not their initial expression in the mouse.Development,2013,140(2):267-279.
[24]NISHIOKA N,INOUE K,ADACHI K,et al.The Hippo signaling pathway components Lats and Yap pattern Tead4activity to distinguish mouse trophectoderm from inner cell mass.Developmental Cell,2009,16(3):398-410.
[25]HIRATE Y,COCKBURN K,ROSSANT J,et al.Tead4 is constitutively nuclear,while nuclear vs.cytoplasmic Yap distribution is regulated in preimplantation mouse embryos.Proceedings of the National Academy of Sciences of the USA,2012,109(50):E3389-E3390.
[26]SASAKI H.Roles and regulations of Hippo signaling during preimplantation mouse development.Development,Growth&Differentiation,2017,59(1):12-20.
[27]LORTHONGPANICH C,MESSERSCHMIDT D M,CHANS W,et al.Temporal reduction of LATS kinases in the early preimplantation embryo prevents ICM lineage differentiation.Genes&Development,2013,27(13):1441-1446.
[28]COCKBURN K,BIECHELE S,GARNER J,et al.The Hippo pathway member Nf2 is required for inner cell mass specification.Current Biology,2013,23(13):1195-1201.
[29]HIRATE Y,HIRAHARA S,INOUE K,et al.Polaritydependent distribution of angiomotin localizes Hippo signaling in preimplantation embryos.Current Biology,2013,23(13):1181-1194.
[30]WICKLOW E,BLIJ S,FRUM T.HIPPO pathway members restrict SOX2 to the inner cell mass where it promotes ICMfates in the mouse blastocyst.PLo S Genetics,2014,10(10):e1004618.
[31]DE LAS HERAS J M,CASANOVA J.Spatially distinct downregulation of Capicua repression and tailless activation by the Torso RTK pathway in the Drosophila embryo.Mechanisms of Development,2006,123(6):481-486.
[32]CHAZAUD C,YAMANAKA Y,PAWSON T,et al.Early lineage segregation between epiblast and primitive endoderm in mouse blastocysts through the Grb2-MAPKpathway.Developmental Cell,2006,10(5):615-624.
[33]MCKAY M M,MORRISON D K.Integrating signals from RTKs to ERK/MAPK.Oncogene,2007,26(22):3113-3121.
[34]SCHRODE N,SAIZ N,DI TALIA S,et al.GATA6 levels modulate primitive endoderm cell fate choice and timing in the mouse blastocyst.Developmental Cell,2014,29(4):454-467.
[35]TAKAOKA K,HAMADA H.Cell fate decisions and axis determination in the early mouse embryo.Development,2012,139(1):3-14.
[36]KOCH U,LEHAL R,RADTKE F.Stem cells living with a Notch.Development,2013,140(4):689-704.
[37]WANG H F,ZANG C Z,LIU X S,et al.The role of Notch receptors in transcriptional regulation.Journal of Cellular Physiology,2015,230(5):982-988.
[38]RAYON T,MENCHERO S,NIETO A,et al.Notch and Hippo converge on Cdx2 to specify the trophectoderm lineage in the mouse blastocyst.Developmental Cell,2014,30(4):410-422.