We compared the combinatorial expression patterns of several LIM domain-containing regulatory genes in the ventrolateral pallium of mouse and chicken, in order to identify the homologues of the ventral pallial amygdala and other olfactory structures in birds. Lmo3, Lmo4, Lhx2, and Lhx9 showed comparable expression patterns in the telencephalon of mouse and chicken, which allowed distinction of the ventrolateral pallium and, particularly, the ventral pallial amygdala and entorhinal cortex. Lmo3 was expressed in most of the ventrolateral pallium in both species, including, in chicken, the piriform cortex and dorsal ventricular ridge (mesopallium, nidopallium, and arcopallium) and, in mouse, the piriform cortex, most of the claustral complex, and the pallial amygdala. Lhx9 was differentially expressed in the ventral pallium, where it was restricted to its rostral (olfactory bulb) and caudal (amygdalar and entorhinal) poles. In the caudal pole, expression of Lhx9 overlapped that of its paralog Lhx2. According to these expression patterns, the chicken ventral pallial amygdala appears to include the caudal dorsolateral pallium, the caudal nidopallium, and the whole arcopallium, and each one relates to a distinct ventricular sector. Finally, the combinatorial expression patterns of Lmo3, Lhx9, and Lmo4 distinguished four distinct subdivisions in the superficial, olfactorecipient area of the chicken ventral pallium, which appear comparable to the piriform, entorhinal, amygdalopiriform, and amygdalar cortices of mammals. The results are discussed in the context of the two existing, opposite views on the homology of the dorsal ventricular ridge of sauropsids and in terms of the evolution of pallial derivatives.

We compared expression of the paralogous LIM-homeodomain genes Lhx1 and Lhx5 in the developing rostral forebrain of mouse and chicken. Both genes are expressed in similar patterns in the septum, preoptic region, and related areas of the basal telencephalon, including the medial septum/diagonal band nuclei and the medial extended amygdala. In the septum, the expression of Lhx5 and Lhx1 appears to be specifically related to the pallial septum and its derivatives in mouse and chicken, and may produce the glutamatergic neurons observed in the diagonal band/medial septum nuclei. The preoptic area expresses both Lhx1 and Lhx5 in mouse and chicken, and appears to produce gamma-aminobutyric acid (GABA)ergic, glutamatergic, and cholinergic cells for the preoptic region and basal telencephalon. In addition, in mouse and chicken Lhx5 is expressed in two extratelencephalic domains that appear to contribute Lhx5-expressing cells to the basal telencephalon, including the supraoptoparaventricular hypothalamic domain and the eminentia thalami. In contrast, there are striking differences in the pallial expression of Lhx1 and Lhx5 between mouse and chicken. Both genes are expressed in Cajal-Retzius cells, and Lhx5 is also present in most pallial sources of Cajal-Retzius cells (including the cortical hem and retrobulbar area) and in the olfactory bulbs in the mouse. In contrast, putative Cajal-Retzius cells, the retrobulbar area, and the olfactory bulb of chicken do not express the paralog genes cLhx1/cLhx5. Moreover, the cortical hem-although it expresses cLhx5-is very tiny in chicken. We discuss the consequences of these differences in Lhx1/Lhx5 expression between mouse and chicken for pallial/cortical evolution.

Although evidence exists for a lateralization of song production, few studies have focused on the perceptual aspect of lateralization in songbirds. In the present study, the authors recorded neuronal responses to a variety of species-specific and artificial, nonspecific stimuli in both hemispheres of awake and anesthetized male starlings (Sturnus vulgaris). Recordings were made in the primary auditory area of the songbird brain, the Field L complex. The right hemisphere exhibited significantly more responsive units than the left hemisphere in awake birds, and this difference was significantly reduced in anesthetized birds. Furthermore, clear hemispheric specialization toward categories of behaviorally relevant stimuli and precise parameters of these stimuli were found. The main auditory area of the starling's brain thus appears to show some degree of lateralization.