Localization of cerebellin-2 in late embryonic chicken brain: implications for a role in synapse formation and for brain evolution.

Cerebellin-1 (Cbln1), the most studied member of the cerebellin family of secreted proteins, is necessary for the formation and maintenance of parallel fiber-Purkinje cell synapses. However, the roles of the other Cblns have received little attention. We previously identified the chicken homolog of Cbln2 and examined its expression in dorsal root ganglia and spinal cord (Yang et al. [2010] J Comp Neurol 518:2818-2840). Interestingly, Cbln2 is expressed by mechanoreceptive and proprioceptive neurons and in regions of the spinal cord where those afferents terminate, as well as by preganglionic sympathetic neurons and their sympathetic ganglia targets. These findings suggest that Cbln2 may demonstrate a tendency to be expressed by synaptically connected neuronal populations. To further assess this possibility, we examined Cbln2 expression in chick brain. We indeed found that Cbln2 is frequently expressed by synaptically connected neurons, although there are exceptions, and we discuss the implications of these findings for Cbln2 function. Cbln2 expression tends to be more common in primary sensory neurons and in second-order sensory regions than it is in motor areas of the brain. Moreover, we found that the level of Cbln2 expression for many regions of the chicken brain is very similar to that of the mammalian homologs, consistent with the view that the expression patterns of molecules playing fundamental roles in processes such as neuronal communication are evolutionarily conserved. There are, however, large differences in the pattern of Cbln2 expression in avian as compared to mammalian telencephalon and in other regions that show the most divergence between the two lineages.