The avian circadian system is composed of multiple inputs, oscillators and outputs. Among its oscillators are the pineal gland, retinae and a hypothalamic structure assumed to be homologous to the mammalian suprachiasmatic nucleus (SCN). Two structures have been suggested as this homolog-the medial SCN (mSCN) and the visual SCN (vSCN). The present study employed biotin dextran amine (BDA) and cholera toxin B subunit (CTB) as anterograde and retrograde tracers to investigate the connectivity of the mSCN and vSCN in order to address this issue. Intravitreal injections of CTB were used to determine whether one or both of these structures receives afferent input from retinal ganglion cells. Both the vSCN and mSCN receive terminal retinal input, with the strongest input terminating in the vSCN. Precise iontophoretic injections of BDA and CTB in the mSCN and vSCN were used to identify efferents and afferents. The avian mSCN and vSCN collectively express more efferents and afferents than does the mammalian SCN. A subset of these connections matches the connections that have been established in rodent species. Individually, both the mSCN and vSCN are similar to the mammalian SCN in terms of their connections. Based on these data and other studies, we present a working model of the avian SCN that includes both the mSCN and vSCN as hypothalamic oscillators. We contend that both structures are involved in a suprachiasmatic complex that, as a functional group, may be homologous to the mammalian SCN.

The vertebrate circadian system is composed of multiple inputs, oscillators, pacemakers, and outputs. In birds, the pineal gland and retinae have been defined as pacemakers within this system. Evidence for a third, hypothalamic pacemaker is abundant. It has been presumed that this pacemaker is homologous to the mammalian suprachiasmatic nucleus (SCN). Two candidate structures have been referred to as the avian SCN--the medial SCN (mSCN) and the visual SCN (vSCN). Previously, we suggested that both structures are involved in a "suprachiasmatic complex." To further explore evidence for an avian SCN, the present study employed several classical techniques to assess intrinsic characteristics of the mSCN and vSCN in the chicken. First, analysis of mSCN and vSCN cytoarchitecture indicated that the mSCN is similar in location and cell population to the mammalian SCN, while the vSCN is more similar in terms of its shape. Second, intravitreal injections of tritiated proline were used to identify hypothalamic retinal terminals. The findings support previous studies identifying the vSCN as the primary retinorecipient hypothalamic structure in birds. Third, analysis of mSCN and vSCN chemoarchitecture suggests that both the mSCN and vSCN display similarity to the mammalian SCN, which displays significant interspecies variation. Finally, a unique astrocytic bridge between the mSCN and vSCN is demonstrated, suggesting that astrocytes play a role within the suprachiasmatic nuclei of birds, similar to the situation in mammals. Our previously presented working model of the avian suprachiasmatic complex is updated to include these data.

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