{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T11:54:57Z","timestamp":1791806097051,"version":"4.3.4"},"reference-count":153,"publisher":"Wiley","issue":"12","license":[{"start":{"date-parts":[[2016,4,13]],"date-time":"2016-04-13T00:00:00Z","timestamp":1460505600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2016,4,13]],"date-time":"2016-04-13T00:00:00Z","timestamp":1460505600000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["The Journal of Physiology"],"published-print":{"date-parts":[[2016,6,15]]},"abstract":"<jats:sec>\n                    <jats:title>Key points<\/jats:title>\n                    <jats:p>\n                      <jats:list list-type=\"bullet\">\n                        <jats:list-item>\n                          <jats:p>\n                            <jats:italic>Xenopus laevis<\/jats:italic>\n                            craniofacial development is a good system for the study of Andersen\u2013Tawil Syndrome (ATS)\u2010associated craniofacial anomalies (CFAs) because (1)\n                            <jats:italic>Kcnj2<\/jats:italic>\n                            is expressed in the nascent face; (2) molecular\u2010genetic and biophysical techniques are available for the study of ion\u2010dependent signalling during craniofacial morphogenesis; (3) as in humans, expression of variant\n                            <jats:italic>Kcnj2<\/jats:italic>\n                            forms in embryos causes a muscle phenotype; and (4) variant forms of\n                            <jats:italic>Kcnj2<\/jats:italic>\n                            found in human patients, when injected into frog embryos, cause CFAs in the same cell lineages.\n                          <\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>\n                            Forced expression of WT or variant\n                            <jats:italic>Kcnj2<\/jats:italic>\n                            changes the normal pattern of\n                            <jats:italic>V<\/jats:italic>\n                            <jats:sub>mem<\/jats:sub>\n                            (resting potential) regionalization found in the ectoderm of neurulating embryos, and changes the normal pattern of expression of ten different genetic regulators of craniofacial development, including markers of cranial neural crest and of placodes.\n                          <\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>\n                            Expression of other potassium channels and two different light\u2010activated channels, all of which have an effect on\n                            <jats:italic>V<\/jats:italic>\n                            <jats:sub>mem<\/jats:sub>\n                            , causes CFAs like those induced by injection of\n                            <jats:italic>Kcnj<\/jats:italic>\n                            2 variants. In contrast, expression of\n                            <jats:italic>Slc9A<\/jats:italic>\n                            (NHE3), an electroneutral ion channel, and of\n                            <jats:italic>GlyR<\/jats:italic>\n                            , an inactive Cl\n                            <jats:sup>\u2212<\/jats:sup>\n                            channel, do not cause CFAs, demonstrating that correct craniofacial development depends on a pattern of bioelectric states, not on ion\u2010 or channel\u2010specific signalling.\n                          <\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>\n                            Using optogenetics to control both the location and the timing of ion flux in developing embryos, we show that affecting\n                            <jats:italic>V<\/jats:italic>\n                            <jats:sub>mem<\/jats:sub>\n                            of the ectoderm and no other cell layers is sufficient to cause CFAs, but only during early neurula stages. Changes in\n                            <jats:italic>V<\/jats:italic>\n                            <jats:sub>mem<\/jats:sub>\n                            induced late in neurulation do not affect craniofacial development.\n                          <\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>\n                            We interpret these data as strong evidence, consistent with our hypothesis, that ATS\u2010associated CFAs are caused by the effect of variant\n                            <jats:italic>Kcnj2<\/jats:italic>\n                            on the\n                            <jats:italic>V<\/jats:italic>\n                            <jats:sub>mem<\/jats:sub>\n                            of ectodermal cells of the developing face. We predict that the critical time is early during neurulation, and the critical cells are the ectodermal cranial neural crest and placode lineages. This points to the potential utility of extant, ion flux\u2010modifying drugs as treatments to prevent CFAs associated with channelopathies such as ATS.\n                          <\/jats:p>\n                        <\/jats:list-item>\n                      <\/jats:list>\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      Variants in potassium channel\n                      <jats:italic>KCNJ2<\/jats:italic>\n                      cause Andersen\u2013Tawil Syndrome (ATS); the induced craniofacial anomalies (CFAs) are entirely unexplained. We show that\n                      <jats:italic>KCNJ2<\/jats:italic>\n                      is expressed in\n                      <jats:italic>Xenopus<\/jats:italic>\n                      and mouse during the earliest stages of craniofacial development. Misexpression in\n                      <jats:italic>Xenopus<\/jats:italic>\n                      of\n                      <jats:italic>KCNJ2<\/jats:italic>\n                      carrying ATS\u2010associated mutations causes CFAs in the same structures affected in humans, changes the normal pattern of membrane voltage potential regionalization in the developing face and disrupts expression of important craniofacial patterning genes, revealing the endogenous control of craniofacial patterning by bioelectric cell states. By altering cells\u2019 resting potentials using other ion translocators, we show that a change in ectodermal voltage, not tied to a specific protein or ion, is sufficient to cause CFAs. By adapting optogenetics for use in non\u2010neural cells in embryos, we show that developmentally patterned K\n                      <jats:sup>+<\/jats:sup>\n                      flux is required for correct regionalization of the resting potentials and for establishment of endogenous early gene expression domains in the anterior ectoderm, and that variants in\n                      <jats:italic>KCNJ2<\/jats:italic>\n                      disrupt this regionalization, leading to the CFAs seen in ATS patients.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1113\/jp271930","type":"journal-article","created":{"date-parts":[[2016,2,10]],"date-time":"2016-02-10T23:59:04Z","timestamp":1455148744000},"page":"3245-3270","source":"Crossref","is-referenced-by-count":129,"title":["Bioelectric signalling via potassium channels: a mechanism for craniofacial dysmorphogenesis in KCNJ2\u2010associated Andersen\u2013Tawil Syndrome"],"prefix":"10.1113","volume":"594","author":[{"given":"Dany Spencer","family":"Adams","sequence":"first","affiliation":[{"name":"Department of Biology and Tufts Centre for Regenerative and Developmental Biology Tufts University  200 Boston Avenue Medford MA 02155 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sebastien G. 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Crotty","family":"Yelick","sequence":"additional","affiliation":[{"name":"Department of Orthodontics, Division of Craniofacial and Molecular Genetics Tufts University School of Dental Medicine  Boston MA 02111 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Adrian","family":"Devitt\u2010Lee","sequence":"additional","affiliation":[{"name":"Department of Biology and Tufts Centre for Regenerative and Developmental Biology Tufts University  200 Boston Avenue Medford MA 02155 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jean\u2010Francois","family":"Pare","sequence":"additional","affiliation":[{"name":"Department of Biology and Tufts Centre for Regenerative and Developmental Biology Tufts University  200 Boston Avenue Medford MA 02155 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Levin","sequence":"additional","affiliation":[{"name":"Department of Biology and Tufts Centre for Regenerative and 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