Bibliography from NCBI

Selected papers


Sokol S., Wong G.G., Melton D.A. (1990). A mouse macrophage factor induces head structures and organizes a body axis in Xenopus. Science 249, 561-564.

Sokol, S., and Melton, D.A. (1991). Pre-existent pattern in Xenopus animal pole cells revealed by induction with activin. Nature 351, 409-411.

Sokol, S. , Christian J., Moon, R.T., and D.A. Melton (1991). Injected Wnt RNA induces a complete body axis in Xenopus embryos. Cell 67, 741-752.

Sokol, S., and Melton, D.A. (1992). Interaction of Wnt and activin in dorsal mesoderm induction in Xenopus . Dev. Biol. 154 , 348-355.

Sokol, S. (1993). Mesoderm formation in Xenopus ectodermal explants overexpressing Xwnt8: Evidence for a cooperating signal reaching animal pole by gastrulation. Development 118 , 1335-1342.

Itoh, K., and Sokol, S. (1994). Heparan sulfate proteoglycans are required for mesoderm formation in Xenopus embryos. Development 120, 2703-2711.

Sokol, S. (1994). The pregastrula establishment of gene expression pattern in Xenopus embryos: requirements for local cell interactions and for protein synthesis. Dev. Biol. 166, 782-788.

Itoh, K., Tang, T.L., Neel, B.G., and Sokol, S. (1995). Specific modulation of ectodermal cell fates in Xenopus embryos by glycogen synthase kinase. Development 121, 3979-3988.

Tang, T.L., Freeman, R.M., O’Reilly, A.M., Neel, B.G., and Sokol, S. (1995). The SH2-containing protein-tyrosine phosphatase SH-PTP2 is required upstream of MAP kinase for early Xenopus development. Cell 80, 473-483.

Sokol, S., and Itoh, K. (1995). The role of proteoglycans in Xenopus gastrulation. In: Proceedings of the Workshop ‘Interplay of genetic and physical processes in the development of biological form’, ed. Beysens, D., Les Houches Series, World Scientific, Singapore, pp.106-113.

Sokol, S., Klingensmith, J., Perrimon, N., and Itoh, K. (1995). Dorsalizing and neuralizing activities of Xdsh, a maternally expressed Xenopus homolog of dishevelled. Development 121 (6), 1637-1647.

Dominguez, I., Itoh, K., and Sokol, S. (1995). Role of glycogen synthase kinase 3 beta as a negative regulator of dorsoventral axis formation in Xenopus embryos. Proc. Natl. Acad. Sci. USA 92 , 8498-8502.


Sokol, S. (1996). Analysis of Dishevelled signaling pathways during Xenopus development. Curr. Biol. 6, 1456-1467.

Itoh K. and S. Sokol. (1997). Graded amounts ofXenopus dishevelled specify discrete anteroposterior cell fates in prospective ectoderm. Mech. Dev. 61, 113-125.

Fan, M. J., Sokol S.Y. (1997). A role for Siamois in Spemann organizer formation. Development 124, 2581-2589.

Landesman, Y. and S. Sokol (1997). Xwnt-2b is a novel axis-inducing Xenopus Wnt, which is expressed in embryonic brain. Mech.Dev. 63, 199-209.

Fan, M. J., Gruning, W., Walz, G., and Sokol, S. (1998). Wnt signaling and transcriptional control of Siamois in Xenopus embryos. Proc. Natl. Acad. Sci. 95, 5626-5631.

Itoh, K., Krupnik, V. and S.Y. Sokol (1998). Axis determination in Xenopus involves biochemical interactions of axin, glycogen synthase kinase 3 and beta-catenin. Curr.Biol. 8, 591-594.

Sokol, S.Y., and K. Itoh (1998). A network of cell interactions mediated by Frizzled is essential for gastrulation and anteroposterior axis determination in Xenopus embryos. In: “Dynamical networks in Physics and Biology”, Editions de Physique- Springer Verlag, 227-236.

Xu, Q., D’Amore, P. A. and S. Y. Sokol (1998). Functional and biochemical interactions of Wnts with FrzA, a secreted Wnt antagonist. Development Nov 9; 125(Pt 23), 4767-4776.

Kim, E., Arnould, T., Fan, M. J., Gruning, W., Sokol, S. Y., Drummond, I., and Walz, G. (1999). Polycystin modulates Wnt signaling. J Biol Chem. 274, 4947-53.

Itoh, K., Jacob, J. and Sokol, S.Y. (1998). A role for Xenopus Frizzled 8 in dorsal development. Mech. Dev. 74, 145-157.


Abu-Jawdeh, G., Comella, N., Tomita, Y., Brown, L.F., Tognazzi, K., Sokol, S.Y., Kocher, O. (1999). Differential expression of frpHE: a novel human stromal protein of the secreted frizzled gene family, during the endometrial cycle and malignacy. Lab. Invest. 79, 439-337.

Itoh, K., and Sokol, S. Y. (1999) . Axis determination by inhibition of Wnt signaling in Xenopus embryos. Genes Dev.

Krupnik VE, Sharp JD, Jiang C, Robison K, Chickering TW, Amaravadi L, Brown DE, Guyot D, Mays G, Leiby K, Chang B, Duong T, Goodearl AD, Gearing DP, Sokol SY, McCarthy SA. (1999). Functional and structural diversity of the human Dickkopf gene family. Gene. 238(2), 301-13.

Sokol, S.Y. (1999). Wnt signaling and dorsoventral axis specification in vertebrates. Curr. Opin. Genet. Develop. 9, 405-410.

Holowacz, T. and S.Y. Sokol (1999) FGF is required for posterior neural patterning, but not for neural induction. Dev. Biol. 205(2), 296-308.

Broun M, Sokol S, Bode HR (1999) Cngsc, a homologue of goosecoid, participates in the patterning of the head, and is expressed in the organizer region of Hydra. Development. 126(23), 5245-54.

Itoh, K., Antipova, A., Ratcliffe, M.J., Sokol, S.Y. (2000). Interaction of dishevelled and Xenopus axin-related protein is required for Wnt signal transduction. Mol. Cell. Biol. 20, 2228-2238.

Ratcliffe MJ, Itoh K, Sokol SY. (2000) A positive role for the PP2A catalytic subunit in Wnt signal transduction. J. Biol. Chem. 275(46), 35680-3.

Sokol, S. Y. (2000). A role for Wnts in morphogenesis and tissue polarity. Nat. Cell Biol. 2(7), E124-5.

Semenov MV, Tamai K, Brott BK, Kuhl M, Sokol S, He X.(2001) Head inducer Dickkopf-1 is a ligand for Wnt coreceptor LRP6. Curr Biol. 11, 951-61.

Sokol, S. Y. , Li, Z., and Sacks D. (2001). The effect of IQGAP1 on Xenopus embryonic ectoderm requires Cdc42. J Biol Chem. , Oct 2 .


Brott, B. and Sokol S. Y. (2002). Regulation of Wnt/LRP signaling by distinct domains of Dickkopf proteins. Mol Cell Biol., 22, 6100-10.

Lisovsky, M., Itoh, K., and Sokol S. Y. (2002). Frizzled receptors activate a novel JNK-dependent pathway that may lead to apoptosis. Curr Biol., 12(1), 53-8.

Gloy, J., and Sokol S. Y. (2002). Frodo interacts with Dishevelled to transduce Wnt signals. Nat Cell Biol, 4, 351-7.

Hanai, J., Karumanchi, S. A. Gloy, J., Ramchandran, R., Kale, S., Jha, V., Sukhatme, V. P., and Sokol, S. Y. (2002). Endostatin is a potential inhibitor of Wnt signaling. J Cell Biol. 2002 , 158, 529-39.


Weber J.R., Sokol S.Y. (2003). Identification of a phylogenetically conserved activin-responsive enhancer in the Zic3 gene. Mech. Dev.(in press).


Gillhouse M, Wagner Nyholm M, Hikasa H, Sokol SY, Grinblat Y. (2004). Two Frodo/Dapper homologs are expressed in the developing brain and mesoderm of zebrafish. Dev Dyn. 230:403-9.

Hikasa, H. and Sokol, S. Y. (2004). The involvement of Frodo in TCF-dependent signaling and neural tissue development. Development 131, 4725-34.


Itoh, K., Brott, B. K., Bae, G., Ratcliffe, M. R., and Sokol, S. Y. (2005). Nuclear localization is required for Dishevelled function in Wnt/beta-catenin signaling. J. Biol. 4, 3.

Brott, B. K., and Sokol, S. Y. (2005). A vertebrate homologue of the cell cycle regulator Dbf4 is a Wnt inhibitor required for heart development. Dev. Cell 8, 703-715.

Itoh K., Lisovsky M., Hikasa H., and Sokol S. Y. (2005). Reorganization of actin cytoskeleton by FRIED, a Frizzled-8 associated protein tyrosine phosphatase. Dev Dyn. 234:90-101.

Ku M., Sokol S. Y., Wu J., Tussie-Luna M. I., Roy A. L., Hata A. (2005). Positive and negative regulation of the transforming growth factor beta/activin target gene goosecoid by the TFII-I family of transcription factors. Mol Cell Biol. 25:7144-57.

Ossipova O, Dhawan S, Sokol S, Green JB. (2005). Distinct PAR-1 proteins function in different branches of Wnt signaling during vertebrate development. Dev. Cell 8:829-41.

Brott, B. K., and Sokol, S. Y. (2005). Frodo proteins: novel regulators of signaling pathways in vertebrate embryos. Differentiation 73:323-329.

Dollar, G., Weber, U., Mlodzik, M., and Sokol, S. Y. (2005). Regulation of Lethal giant larvae by Dishevelled. Nature 437:1376-1389.


Park J.I., Ji H., Jun S., Gu D., Hikasa H., Li L, Sokol S. Y., McCrea P. D. (2006). Frodo links Dishevelled to the p120-catenin/ Kaiso pathway: distinct catenin sub-families promote Wnt signals. Dev. Cell 11, 683.

Hunter, N. L., Hikasa, H., Dymecki, S. M, and Sokol, S. Y. (2006). Vertebrate homologues of Frodo are dynamically expressed in tissues undergoing extensive morphogenetic movements. Dev. Dyn. 235: 279-284.

Kibardin, A., Ossipova, O., and Sokol, S. Y. (2006). Metastasis-associated kinase modulates Wnt signaling to regulate brain patterning and morphogenesis. Development 133, 2845-54.


Sokol, S. Y. and Wharton, Jr., K. A. (2007). Wnters in La Jolla. Development 134, 3393-9.

Itoh, K, and Sokol S. Y. (2007) Early development of epidermis and neural tissue. In: Principles of Developmental Genetics, S. A. Moody, Ed., Elsevier Press, 241-258.

Ossipova, O., Tabler, J., Green, J. B., and Sokol, S. Y. (2007). PAR-1 specifies ciliated cells in vertebrate ectoderm downstream of aPKC. Development 134, 4297-4306.


Pan W, Choi SC, Wang H, Qin Y, Volpicelli-Daley L, Swan L, Lucast L, Khoo C, Zhang X, Li L, Abrams CS, Sokol S. Y., Wu D. (2008). Wnt3a-mediated formation of phosphatidylinositol 4,5-bisphosphate regulates LRP6 phosphorylation. Science 321, 1350-3.


Lake, B.B., and Sokol, S.Y. (2009). Strabismus regulates asymmetric cell divisions and cell fate determination in the mouse brain. J Cell Biol. 185, 59-66.

Itoh, K., Jenny, A., M. Mlodzik, M., and Sokol, S. Y. (2009). Centrosomal localization of Diversin and its relevance to Wnt signaling. J. Cell Sci. (in press).

Ossipova, O., Ezan, J., and Sokol, S. Y. (2009). PAR-1 phosphorylates Mind bomb to promote vertebrate neurogenesis.
Dev. Cell 17, 222-233.


Ossipova, O., and Sokol, S. Y. (2010). Cell polarity, Notch signaling and neurogenesis. Cell Cycle 9,1-2.

Hikasa, H, Ezan, J, Itoh K, Li, X, Klymkowsky, M. W., Sokol, S. Y. (2010) Regulation of TCF3 by Wnt-Dependent Phosphorylation during Vertebrate Axis Specification. Dev Cell 19: 521-32.


Itoh, K., and Sokol, S. Y. (2011). Polarized translocation of fluorescent proteins in Xenopus ectoderm in response to Wnt signaling. J. Vis. Exp. 51 doi: 10.3791/2700.

Hikasa, H., and Sokol, S. Y. (2011) Phosphorylation of TCF proteins by homeodomain-interacting protein kinase 2. J. Biol. Chem. 286:12093-100.

Yasunaga, T, Itoh, K., Sokol, S. Y. (2011). Regulation of basal body and ciliary functions by Diversin. Mechanisms of Development 128, 376-386.

Sokol, S. Y.(2011). Wnt signal transduction through T-cell factor phosphorylation. Cell Research 21:1002-12.

Ossipova, O., and Sokol, S. Y. (2011). Neural crest specification in Xenopus embryos by noncanonical Wnt signaling and PAR-1. Development 138, 5441-50.

Sokol, S. Y. (2011). Maintaining embryonic stem cell pluripotency with Wnt signaling. Development 138, 4341-4350.


Hong, H., Sokol, S. Y. and McCrea, P. (2012). Down-syndrome related kinase Dyrk1A modulates the p120-catenin/Kaiso trajectory of the Wnt signaling pathway. J. Cell Sci., (in press).

H.-S. Lee, Sokol, S. Y. , Moody, S. A., and I. O. Daar, (2012). Using 32-cell stage Xenopus embryos to probe PCP signaling. In : PCP Methods and Protocols, ed. Daar, I.

Kim K., Lake B. B., Haremaki T., Weinstein D. C., and Sokol S. Y.(2012). Rab11 regulates planar polarity and migratory behavior of multiciliated cells in Xenopus embryonic epidermis. Dev Dyn. Jul 9. doi: 10.1002/dvdy.23826.

Kalinichenko S. V., Itoh K., Korobko E.V., Sokol S. Y., Buchman V. L., Korobko I. V.(2012) Identification of Nedd4 E3 ubiquitin ligase as a binding partner and regulator of MAK-V protein kinase. PLoS One. 2012;7(6):e39505. Epub 2012 Jun 20.


Hikasa, H., and Sokol, S. Y. (2013). Wnt signaling in vertebrate axis specification. Cold Spring Harb Perspect Biol. 5: a007955.

Chu, C., Gerstenzang, E., Ossipova, O., and Sokol, S. Y. (2013). Lulu regulates Shroom-induced apical constriction during neural tube closure. PLOS One 8:e81854.


Itoh, K., Ossipova. O., and Sokol, S. Y. (2014). GEFH1 functions in apical constriction and cell intercalations and is essential for vertebrate neural tube closure. J. Cell Sci. 127: 2542-53. doi: 10.1242/jcs.146811.

Ossipova, O., Kim, K., Lake, B. B., Itoh, K., Ioannou, A., and Sokol, S. Y. (2014). Role of Rab11 in planar cell polarity and apical constriction during vertebrate neural tube closure. Nat. Commun. 5:3734, doi: 10.1038/ncomms4734 (2014).

Itoh, K., and Sokol, S. Y. (2014). Expression cloning of camelid nanobodies specific for Xenopus embryonic antigens. PLOS One, 9: e107521, doi: 10.1371/journal.pone.0107521.


Itoh K, Sokol SY.  Early Development of Epidermis and Neural Tissue.  In: ‘Principles of Developmental Genetics’ Second Edition ed. Moody SA, editor. San Diego, California: Elsevier, Academic Press; 2015. Chapter 11, 189-201p.

Ossipova O, Chuykin I, Chu CW, Sokol SY. Vangl2 cooperates with Rab11 and Myosin V to regulate apical constriction during vertebrate gastrulation. Development (Cambridge, England). 2015; 142(1):99-107.

Kim K, Ossipova O, Sokol SY.  Neural crest specification by inhibition of the ROCK/Myosin II pathway.  Stem cells (Dayton, Ohio). 2015; 33(3):674-85.

Ossipova O, Kim K, Sokol SY.  Planar polarization of Vangl2 in the vertebrate neural plate is controlled by Wnt and Myosin II signaling.  Biology open. 2015; 4(6):722-30.

Sokol SY.  Spatial and temporal aspects of Wnt signaling and planar cell polarity during vertebrate embryonic development.  Seminars in cell & developmental biology. 2015; 42:78-85.

Ossipova O, Chu CW, Fillatre J, Brott BK, Itoh K, Sokol SY.  The involvement of PCP proteins in radial cell intercalations during Xenopus embryonic development.  Developmental Biology. 2015; 408(2):316-27.


Sokol SY.  Mechanotransduction During Vertebrate Neurulation.
Current topics in developmental biology. 2016; 117:359-76.

Gaston-Massuet C, et al.  Transcription factor 7-like 1 is involved in hypothalamo-pituitary axis development in mice and humans. Proc. Natl. Acad.Sci. USA. 2016; 113(5):E548-57.

Chu CW, Ossipova O, Ioannou A, Sokol SY.  Prickle3 synergizes with Wtip to regulate basal body organization and cilia growth. Scientific Reports. 2016; 6:24104.

Chu CW, Sokol SY.Wnt proteins can direct planar cell polarity in vertebrate ectoderm. eLife. 2016; 5.


Sokol SY. At the Crossroads between Cell Polarity and Adhesion in Neocortical Development. Developmental Cell. 2017; 41(5):453-454.

Ossipova O, Kerney R, Saint-Jeannet JP, Sokol SY. Regulation of neural crest development by the formin family protein Daam1. Genesis (New York, N.Y.:2000); :e23108.

Chu CW, Xiang B, Ossipova O, Ioannou A, Sokol SY. The Ajuba family protein Wtip regulates actomyosin contractility during vertebrate neural tube closure. J. Cell.Sci. 131(10).

Chuykin I, Ossipova O, Sokol SY. Par3 interacts with Prickle3 to generate apical PCP complexes in the vertebrate neural plate. Elife. 2018 Sep 26;7. pii: e37881. doi: 10.7554/eLife.37881.

Itoh K, Reis AH, Hayhurst A, Sokol SY. Isolation of nanobodies against Xenopus embryonic antigens using immune and non-immune phage display libraries. PLoS One. 2019 May 2;14(5):e0216083. doi: 10.1371/journal.pone.0216083. eCollection 2019.PMID: 31048885


Reis AH, Sokol SY. Rspo2 antagonizes FGF signaling during vertebrate mesoderm formation and patterning. Development. 2020 May 27;147(10):dev189324. doi: 10.1242/dev.189324.PMID: 32366679

Ossipova O, Itoh K, Radu A, Ezan J, Sokol SY. Pinhead signaling regulates mesoderm heterogeneity via the FGF receptor-dependent pathway. Development. 2020,147(17):dev188094. doi: 10.1242/dev.188094.PMID: 32859582

Mancini P, Ossipova O, Sokol SY. The dorsal blastopore lip is a source of signals inducing planar cell polarity in the Xenopus neural plate.
Biol Open. 2021 Jul 15;10(7):bio058761. doi: 10.1242/bio.058761. Epub 2021 Jul 19.PMID: 34259326

Ossipova O, Sokol SY. Cryosectioning and Immunostaining of Xenopus Embryonic Tissues. Cold Spring Harb Protoc. 2021 Sep 1;2021(9):pdb.prot107151. doi: 10.1101/pdb.prot107151.PMID: 34244349

Reis AH, Sokol SY. Rspo2 inhibits TCF3 phosphorylation to antagonize Wnt signaling during vertebrate anteroposterior axis specification. Sci Rep. 2021 Jun 28;11(1):13433. doi: 10.1038/s41598-021-92824-6.PMID: 34183732

Itoh K, Ossipova O, Sokol SY. Pinhead antagonizes Admp to promote notochord formation. iScience. 2021 May 7;24(6):102520. doi: 10.1016/j.isci.2021.102520. eCollection 2021 Jun 25.PMID: 34142034

Sokol SY. Preface. Curr Top Dev Biol. 2021;145:xi-xiii. doi: 10.1016/S0070-2153(21)00065-X.PMID: 34074538 No abstract available.

Matsuda M, Sokol SY. Xenopus neural tube closure: A vertebrate model linking planar cell polarity to actomyosin contractions. Curr Top Dev Biol. 2021;145:41-60. doi: 10.1016/bs.ctdb.2021.04.001. Epub 2021 May 4.PMID: 34074535

Torban E, Sokol SY. Planar cell polarity pathway in kidney development, function and disease. Nat Rev Nephrol. 2021,17(6):369-385. doi: 10.1038/s41581-021-00395-6. Epub 2021 Feb 5.PMID: 33547419 Review.

Chuykin I, Itoh K, Kim K, and Sokol SY (2021). Frizzled3 inhibits Vangl2-Prickle3 association to establish planar cell polarity in the vertebrate neural plate. BioArxiv https://doiorg/101101/20210502442356.

Reis AH, Xiang B, Itoh K, Sokol SY. (2021). Identification SSX2IP/Msd1 as Wtip-binding partner by a targeted proximity biotinylation approach. BioRxiv/2021/444380.

Matsuda M, Chu CW, and Sokol SY (2021).  LMO7-dependent apical constriction requires the binding of the Myosin heavy chain. BioRxiv.