Graduation Year


Document Type




Degree Granting Department


Major Professor

Toru Shimizu, Ph.D.

Committee Member

Gary Arendash, Ph.D.

Committee Member

Cynthia Cimino, Ph.D.

Committee Member

Cheryl Kirstein, Ph.D.

Committee Member

Paul Sanberg, Ph.D.


avian, cognition, limbic, striatum, ZENK


Relatively little is known about the existence and traits of a possible nucleus accumbens (Acc) region in non-mammals. The current project investigated a likely candidate for such a structure in pigeons, the medioventral (mvMSt) and mediodorsal (mdMSt) parts of avian medial striatum (MSt). The methods employed were threefold: 1) tract-tracing to determine anatomical connections of the MSt; 2) lesion studies to assess MSt's role in a cognitive task (reversal learning); and 3) measuring an immediate-early gene induced protein, ZENK, in striatal regions during courtship behavior in male pigeons.

The MSt was found to have many forebrain (amygdala, hippocampus, dorsal thalamus) and midbrain (ventral tegmental area, substantia nigra) connections similar to those of Acc. In addition, differences in connection patterns between mvMSt and mdMSt indicated that mvMSt was comparable to the shell of Acc, while the mdMSt showed characteristics of Acc core.

Effects of MSt lesions on pattern discrimination and reversal learning were assessed. Both lesion subjects and controls performed similarly on original discrimination. Furthermore, there were no significant differences in MSt lesioned birds compared to controls. However, there was a tendency for the two groups to make different types of errors. Error patterns indicated that sham-lesioned birds had deficits due to key preference, whereas lesioned birds had fixation on previous reward contingencies (perseverative errors). The performance of the lesioned birds was consistent with Acc lesion effects on reversal learning in mammals.

The expression of ZENK in the mvMSt, mdMSt, lateral MSt, and lateral striatum of male birds exposed to either an empty cage or a live female pigeon was quantified. Higher ZENK expression was found in the live pigeon condition for all the striatal structures. However, the degree of difference between live and empty was much higher in the mvMSt and mdMSt than in the other areas. Therefore, mvMSt and mdMSt appear to play a role in anticipatory sexual behaviors, as has been shown in Acc.

The anatomical and functional data from the current study indicate that avian mMSt has numerous similarities with mammalian Acc. These findings will contribute to understanding the evolution of mammalian Acc and identifying the functional significance of avian MSt.