Vitamin A–Not for Your Eyes Only Requirement for Heart Formation Begins Early in Embryogenesis 英文参考文献.docVIP

Nutrients 2010, 2, 532-550; doi:10.3390/nu2050532 OPEN ACCESS nutrients ISSN 2072-6643 /journal/nutrients Review Vitamin A–Not for Your Eyes Only: Requirement for Heart Formation Begins Early in Embryogenesis Maija H. Zile Department of Food Science and Human Nutrition, Michigan State University, East Lansing, MI 48824, USA; E-Mail: zile@; Tel.: +1 (517) 355-8474; ext. 127 Received: 25 March 2010; in revised form: 7 May 2010 / Accepted: 18 May 2010 / Published: 25 May 2010 Abstract: Vitamin A insufficiency has profound adverse effects on embryonic development. Major advances in understanding the role of vitamin A in vertebrate heart formation have been made since the discovery that the vitamin A active form, all-trans- retinoic acid, regulates many genes, including developmental genes. Among the experimental models used, the vitamin A-deficient avian embryo has been an important tool to study the function of vitamin A during early heart formation. A cluster of retinoic acid-regulated developmental genes have been identified that participate in building the heart. In the absence of retinoic acid the embryonic heart develops abnormally leading to embryolethality. Keywords: vitamin A-deficient; quail embryo; retinoic acid; gene regulation; heart morphogenesis Abbreviations RA, all-trans-retinoic acid; VAD, vitamin A-deficient; HH, Hamburger Hamilton developmental stages for avian embryo; ss, somite stage; IFT, inflow tract. 1. Function of Vitamin A, Retinoic Acid, Retinoid Receptors and Gene Regulation It has been more that 80 years since the discovery of vitamin A in 1913 [1,2] and much has been learned about this important vitamin [3-10], but still much more remains to be done to fully understand  Nutrients 2010, 2 533 the complex functions of this micronutrient. The following minireview provides a brief overview of the major questions th