LMO4 is a LIM zinc-binding domain-containing protein. LMO4 cDNA was first isolated from a breast tumor cDNA library. This transcriptional modulator is overexpressed in several epithelial cancers such as prostate, pancreas, and breast. LMO4 exhibits pro-oncogenic activities by inducing centrosome amplification and mitotic spindle abnormalities. LMO4 is also expressed in the brain, in regions involved in learning and the regulation of motivated behavior. In the basolateral amygdala, LMO4 functions to negatively regulate fear learning. Furthermore, in the nucleus accumbens, LMO4 was found to regulate the behavioral effects of cocaine.