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In E9.5 embryos, expressed in the rhombencephalon, metencephalon, and in the cephalic mesoderm surround the optic vesicle. By E12.5, expression continues in the mesenchyme and also begins in subsets of cells in the neuroretina, becoming expressed in the retinal inner neuroblast layers by E16.5. Expressed in developing bipolar cells during retinal development starting at postnatal day 5, and expressed in a subset of cone bipolar cells in the mature retina. Also expressed along the spinal cord, in the ventricular layer, in motor neurons and in the proximal limb buds during embryonic development. Expressed in the developing heart in the endocardium that lines the heart chambers.