Fan Xiaohong, Molotkov Andrei, Manabe Shin-Ichi, Donmoyer Christine M, Deltour Louise, Foglio Mario H, Cuenca Arnold E, Blaner William S, Lipton Stuart A, Duester Gregg
OncoDevelopmental Biology Program. Center for Neuroscience and Aging, Burnham Institute, La Jolla, California 92037, USA.
Mol Cell Biol. 2003 Jul;23(13):4637-48. doi: 10.1128/MCB.23.13.4637-4648.2003.
Genetic studies have shown that retinoic acid (RA) signaling is required for mouse retina development, controlled in part by an RA-generating aldehyde dehydrogenase encoded by Aldh1a2 (Raldh2) expressed transiently in the optic vesicles. We examined the function of a related gene, Aldh1a1 (Raldh1), expressed throughout development in the dorsal retina. Raldh1(-/-) mice are viable and exhibit apparently normal retinal morphology despite a complete absence of Raldh1 protein in the dorsal neural retina. RA signaling in the optic cup, detected by using a RARE-lacZ transgene, is not significantly altered in Raldh1(-/-) embryos at embryonic day 10.5, possibly due to normal expression of Aldh1a3 (Raldh3) in dorsal retinal pigment epithelium and ventral neural retina. However, at E16.5 when Raldh3 is expressed ventrally but not dorsally, Raldh1(-/-) embryos lack RARE-lacZ expression in the dorsal retina and its retinocollicular axonal projections, whereas normal RARE-lacZ expression is detected in the ventral retina and its axonal projections. Retrograde labeling of adult Raldh1(-/-) retinal ganglion cells indicated that dorsal retinal axons project to the superior colliculus, and electroretinography revealed no defect of adult visual function, suggesting that dorsal RA signaling is unnecessary for retinal ganglion cell axonal outgrowth. We observed that RA synthesis in liver of Raldh1(-/-) mice was greatly reduced, thus showing that Raldh1 indeed participates in RA synthesis in vivo. Our findings suggest that RA signaling may be necessary only during early stages of retina development and that if RA synthesis is needed in dorsal retina, it is catalyzed by multiple enzymes, including Raldh1.
遗传学研究表明,视黄酸(RA)信号传导是小鼠视网膜发育所必需的,部分受醛脱氢酶1A2(Raldh2)编码的一种生成RA的醛脱氢酶控制,该酶在视泡中短暂表达。我们研究了一个相关基因醛脱氢酶1A1(Raldh1)的功能,该基因在背侧视网膜的整个发育过程中都有表达。Raldh1基因敲除小鼠是存活的,尽管在背侧神经视网膜中完全没有Raldh1蛋白,但视网膜形态明显正常。在胚胎第10.5天,通过使用RA反应元件 - 乳糖酶Z转基因检测到的视杯中RA信号传导在Raldh1基因敲除胚胎中没有显著改变,这可能是由于醛脱氢酶1A3(Raldh3)在背侧视网膜色素上皮和腹侧神经视网膜中正常表达。然而,在胚胎第16.5天,当Raldh3在腹侧而非背侧表达时,Raldh1基因敲除胚胎在背侧视网膜及其视网膜 - 丘脑轴突投射中缺乏RA反应元件 - 乳糖酶Z表达,而在腹侧视网膜及其轴突投射中检测到正常的RA反应元件 - 乳糖酶Z表达。对成年Raldh1基因敲除视网膜神经节细胞的逆行标记表明,背侧视网膜轴突投射到上丘,视网膜电图显示成年视觉功能无缺陷,这表明背侧RA信号传导对于视网膜神经节细胞轴突生长不是必需的。我们观察到Raldh1基因敲除小鼠肝脏中的RA合成大大减少,从而表明Raldh1确实参与体内RA合成。我们的研究结果表明,RA信号传导可能仅在视网膜发育的早期阶段是必需的,并且如果背侧视网膜需要RA合成,则由多种酶催化,包括Raldh1。