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While signaling pathways are known to control organ and body size, the downstream effectors that mediate their effects remain poorly understood. In the nematode Caenorhabditis elegans, a Bone Morphogenetic Protein (BMP)-related signaling pathway is the major regulator of growth and body size. We investigated the transcriptional network through which the BMP pathway regulates body size and identified cuticle collagen genes as major effectors of growth control. We demonstrate that cuticle collagens can act as positive regulators (col-41), negative regulators (col-141), or dose-sensitive regulators (rol-6) of body size. Moreover, we find a requirement of BMP signaling for stage-specific expression of cuticle collagen genes. We show that the Smad signal transducers directly bind conserved Smad-binding elements in regulatory regions of col-141 and col-142, but not of col-41. Hence, cuticle collagen genes may be directly and indirectly regulated via the BMP pathway. Our work thus connects a conserved signaling pathway with its critical downstream effectors, advancing insight into how body size is specified. Since collagen mutations and misregulation are implicated in numerous human genetic disorders and injury sequelae, understanding how collagen gene expression is regulated has broad implications.<\/jats:p>","DOI":"10.1534\/genetics.118.301631","type":"journal-article","created":{"date-parts":[[2018,10,1]],"date-time":"2018-10-01T18:55:14Z","timestamp":1538420114000},"page":"1355-1367","source":"Crossref","is-referenced-by-count":50,"title":["BMP Signaling Determines Body Size via Transcriptional Regulation of Collagen Genes in\n                    <i>Caenorhabditis elegans<\/i>"],"prefix":"10.1093","volume":"210","author":[{"given":"Uday","family":"Madaan","sequence":"first","affiliation":[{"name":"Department of Biology, Queens College, The City University of New York, Flushing, New York 11367"},{"name":"PhD Program in Biology, The Graduate Center, The City University of New York 10016, New York"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Edlira","family":"Yzeiraj","sequence":"additional","affiliation":[{"name":"Department of Biology, Queens College, The City University of New York, Flushing, New York 11367"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Meade","sequence":"additional","affiliation":[{"name":"Department of Biology, Queens College, The City University of New York, Flushing, New York 11367"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James F","family":"Clark","sequence":"additional","affiliation":[{"name":"Department of Biology, Queens College, The City University of New York, Flushing, New York 11367"},{"name":"PhD Program in Biology, The Graduate Center, The City University of New York 10016, New York"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christine A","family":"Rushlow","sequence":"additional","affiliation":[{"name":"Department of Biology, New York University, New York 10003"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cathy","family":"Savage-Dunn","sequence":"additional","affiliation":[{"name":"Department of Biology, Queens College, The City University of New York, Flushing, New York 11367"},{"name":"PhD Program in Biology, The Graduate Center, The City University of New York 10016, New York"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2018,10,1]]},"reference":[{"key":"2022011706420447000_bib1","doi-asserted-by":"publisher","first-page":"27678","DOI":"10.1074\/jbc.272.44.27678","article-title":"TGF-\u03b2 RI phosphorylation of Smad2 on Ser465 and Ser467 is required for Smad2-Smad4 complex formation and signaling.","volume":"272","author":"Abdollah","year":"1997","journal-title":"J. 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