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However, its effects on retinal cells and its intracellular signaling remain relatively unexplored. During development, expression of GFR\u03b13, a specific receptor for artemin, is strong in the immature retina and gradually decreases during maturation, suggesting a possible role in the formation of retinal connections. Optic nerve damage in mature rats causes levels of GFR\u03b13 mRNA to increase tenfold in the retina within 3 days. GFR\u03b13 mRNA levels continue to rise within the first week and then decline. Artemin, a specific ligand for GFR\u03b13, has a neuroprotective effect on axotomized retinal ganglion cells (RGCs) in vivo and in vitro via activation of the extracellular signal\u2010related kinase\u2212 and phosphoinositide 3\u2010kinase\u2212Akt signaling pathways. Artemin also has a substantial effect on axon regeneration in RGCs both in vivo and in vitro, whereas other GDNF family members do not. 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