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However, bladder development and its embryonic function remain poorly understood. Here, we report that in both humans and mice the bladder emerges from the cloaca as a rostral outgrowth, which is flanked by the umbilical arteries. Arrest of the outgrowth, as seen in Shh-null mouse embryos, causes bladder agenesis and unexpected severe defects of the umbilical arteries, suggesting that the outgrowth epithelial signal Shh coordinates development of both the bladder and the umbilical arteries. The signaling molecule Wnt2 is restricted to the rostral outgrowth mesenchyme and its expression is dramatically downregulated in Shh-null mutants. Furthermore, Wnt2-null mutants develop a small bladder and single umbilical artery defects. Collectively, we uncover an unexpected co-development mechanism of the bladder and umbilical arteries orchestrated by the cloacal outgrowth signals. The co-development paradigm offers a conceptual framework to understand evolution and development of the bladder.<\/jats:p>","DOI":"10.1242\/dev.205398","type":"journal-article","created":{"date-parts":[[2026,7,10]],"date-time":"2026-07-10T08:10:09Z","timestamp":1783671009000},"source":"Crossref","is-referenced-by-count":0,"title":["The cloacal outgrowth orchestrates co-development of the bladder and umbilical arteries"],"prefix":"10.1242","volume":"153","author":[{"given":"Xing","family":"Ye","sequence":"first","affiliation":[{"name":"Samuel Oschin Comprehensive Cancer Institute, Cedars-Sinai Medical Center, Cedars-Sinai Health Sciences University 1 Department of Medicine, Department of Biomedical Sciences , , 8700 Beverly Blvd, Los Angeles, CA 90048 ,","place":["USA"]},{"name":"Department of Urology, Boston Children's Hospital, Harvard Medical School 2 , 300 Longwood Ave, Boston, MA 02115 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liguang","family":"Xia","sequence":"additional","affiliation":[{"name":"Department of Urology, Boston Children's Hospital, Harvard Medical School 2 , 300 Longwood Ave, Boston, MA 02115 ,","place":["USA"]},{"name":"The Second Affiliated Hospital and Yuying Children's Hospital of Wenzhou Medical University 3 Department of Pediatric Surgery , , Wenzhou 325027, Zhejiang Province ,","place":["People's Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xianfa","family":"Yang","sequence":"additional","affiliation":[{"name":"Guangzhou Institutes of Biomedicine and Health, Chinese Academy of Sciences 4 Guangdong Provincial Key Laboratory of Stem Cell and Regenerative Medicine , , Guangzhou 510005 ,","place":["People's Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ruirong","family":"Tan","sequence":"additional","affiliation":[{"name":"Samuel Oschin Comprehensive Cancer Institute, Cedars-Sinai Medical Center, Cedars-Sinai Health Sciences University 1 Department of Medicine, Department of Biomedical Sciences , , 8700 Beverly Blvd, Los Angeles, CA 90048 ,","place":["USA"]},{"name":"Department of Urology, Boston Children's Hospital, Harvard Medical School 2 , 300 Longwood Ave, Boston, MA 02115 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haochuan","family":"Zhang","sequence":"additional","affiliation":[{"name":"Department of Urology, Boston Children's Hospital, Harvard Medical School 2 , 300 Longwood Ave, Boston, MA 02115 ,","place":["USA"]},{"name":"The Second Affiliated Hospital and Yuying Children's Hospital of Wenzhou Medical University 3 Department of Pediatric Surgery , , Wenzhou 325027, Zhejiang Province ,","place":["People's Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liming","family":"Lei","sequence":"additional","affiliation":[{"name":"Department of Urology, Boston Children's Hospital, Harvard Medical School 2 , 300 Longwood Ave, Boston, MA 02115 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jungang","family":"Huang","sequence":"additional","affiliation":[{"name":"Department of Urology, Boston Children's Hospital, Harvard Medical School 2 , 300 Longwood Ave, Boston, MA 02115 ,","place":["USA"]},{"name":"Guangdong Cardiovascular Institute, Guangdong Provincial People's Hospital, Guangdong Academy of Medical Sciences, Southern Medical University 5 , Guangzhou, Guangdong 510100 ,","place":["People's Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1580-2781","authenticated-orcid":false,"given":"Yingsheng","family":"Zhang","sequence":"additional","affiliation":[{"name":"Samuel Oschin Comprehensive Cancer Institute, Cedars-Sinai Medical Center, Cedars-Sinai Health Sciences University 1 Department of Medicine, Department of Biomedical Sciences , , 8700 Beverly Blvd, Los Angeles, CA 90048 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Edward","family":"Morrisey","sequence":"additional","affiliation":[{"name":"University of Pennsylvania 6 Department of Medicine, Department of Cell and Developmental Biology , , Philadelphia, PA 19104 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4276-9593","authenticated-orcid":false,"given":"Ping","family":"Zhu","sequence":"additional","affiliation":[{"name":"Guangdong Cardiovascular Institute, Guangdong Provincial People's Hospital, Guangdong Academy of Medical Sciences, Southern Medical University 5 , Guangzhou, Guangdong 510100 ,","place":["People's Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhongrong","family":"Li","sequence":"additional","affiliation":[{"name":"The Second Affiliated Hospital and Yuying Children's Hospital of Wenzhou Medical University 3 Department of Pediatric Surgery , , Wenzhou 325027, Zhejiang Province ,","place":["People's Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Naihe","family":"Jing","sequence":"additional","affiliation":[{"name":"Guangzhou Institutes of Biomedicine and Health, Chinese Academy of Sciences 4 Guangdong Provincial Key Laboratory of Stem Cell and Regenerative Medicine , , Guangzhou 510005 ,","place":["People's Republic of China"]},{"name":"Institute for Stem Cell and Regeneration, Chinese Academy of Sciences 7 , Beijing 100101 ,","place":["People's Republic of 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