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Understanding the pathogenesis of <jats:styled-content style=\"fixed-case\">HPE<\/jats:styled-content> requires knowledge of the relationship between the developing brain and the facial structures during embryogenesis. A number of signaling pathways control and coordinate the development of the brain and face, including Sonic hedgehog, Bone morphogenetic protein, Fibroblast growth factor, and Nodal signaling. Mutations in these pathways have been identified in animal models of <jats:styled-content style=\"fixed-case\">HPE<\/jats:styled-content> and human patients. Because of incomplete penetrance and variable expressivity of <jats:styled-content style=\"fixed-case\">HPE<\/jats:styled-content>, patients carrying defined mutations may not manifest the disease at all, or have a spectrum of defects. It is currently unknown what drives manifestation of <jats:styled-content style=\"fixed-case\">HPE<\/jats:styled-content> in genetically at\u2010risk individuals, but it has been speculated that other gene mutations and environmental factors may combine as cumulative insults. <jats:styled-content style=\"fixed-case\">HPE<\/jats:styled-content> can be diagnosed <jats:italic>in utero<\/jats:italic> by a high\u2010resolution prenatal ultrasound or a fetal magnetic resonance imaging, sometimes in combination with molecular testing from chorionic villi or amniotic fluid sampling. Currently, there are no effective preventive methods for <jats:styled-content style=\"fixed-case\">HPE<\/jats:styled-content>. Better understanding of the mechanisms of gene\u2013environment interactions in <jats:styled-content style=\"fixed-case\">HPE<\/jats:styled-content> would provide avenues for such interventions. <jats:italic>WIREs Dev Biol<\/jats:italic> 2015, 4:17\u201332. doi: 10.1002\/wdev.161<\/jats:p><jats:p>This article is categorized under:\n<jats:list list-type=\"explicit-label\">\n<jats:list-item><jats:p>Nervous System Development &gt; Vertebrates: General Principles<\/jats:p><\/jats:list-item>\n<jats:list-item><jats:p>Birth Defects &gt; Craniofacial and Nervous System Anomalies<\/jats:p><\/jats:list-item>\n<\/jats:list><\/jats:p>","DOI":"10.1002\/wdev.161","type":"journal-article","created":{"date-parts":[[2014,10,22]],"date-time":"2014-10-22T10:36:34Z","timestamp":1413974194000},"page":"17-32","source":"Crossref","is-referenced-by-count":93,"title":["Holoprosencephaly: signaling interactions between the brain and the face, the environment and the genes, and the phenotypic variability in animal models and humans"],"prefix":"10.1002","volume":"4","author":[{"given":"Anna","family":"Petryk","sequence":"first","affiliation":[{"name":"Department of Pediatrics University of Minnesota  Minneapolis MN USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel","family":"Graf","sequence":"additional","affiliation":[{"name":"School of Dentistry University of Alberta  Edmonton Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ralph","family":"Marcucio","sequence":"additional","affiliation":[{"name":"Department of Orthopaedic Surgery, Orthopaedic Trauma Institute University of California, San Francisco  San Francisco CA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2014,10,22]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1172\/JCI38937"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.c.30238"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ydbio.2003.11.015"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.1320140411"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.c.30239"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.c.30236"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.c.30233"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1038\/ng1196-357"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.10258"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0959-437X(00)00084-8"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1002\/humu.21016"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.c.30248"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1086\/344412"},{"key":"e_1_2_9_15_1","first-page":"390","article-title":"Aberrant forebrain signaling during early development underlies the generation of holoprosencephaly and coloboma","volume":"2011","author":"Gongal PA","year":"1812","journal-title":"Biochim Biophys Acta"},{"key":"e_1_2_9_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0070-2153(08)00603-0"},{"key":"e_1_2_9_17_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.026583"},{"key":"e_1_2_9_18_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.00397"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.1002\/tera.1420510605"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1006\/dbio.1997.8589"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.162356199"},{"key":"e_1_2_9_22_1","doi-asserted-by":"publisher","DOI":"10.1002\/tera.1420400304"},{"key":"e_1_2_9_23_1","doi-asserted-by":"publisher","DOI":"10.3174\/ajnr.A2123"},{"key":"e_1_2_9_24_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.c.30249"},{"key":"e_1_2_9_25_1","doi-asserted-by":"publisher","DOI":"10.1093\/cercor\/bhp009"},{"key":"e_1_2_9_26_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.c.30241"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1098-2779(2000)6:1<15::AID-MRDD3>3.0.CO;2-8"},{"key":"e_1_2_9_28_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajmg.10672"},{"key":"e_1_2_9_29_1","first-page":"196","article-title":"Perspectives on holoprosencephaly: Part II. 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