We carried out a study of the expression patterns of seven developmental regulatory genes (Lef1, Lhx2, Lhx9, Lhx5, Lmo3, Lmo4, and Prox1), in combination with topological position, to identify the medial pallial derivatives, define its major subdivisions, and compare them between mouse and chicken. In both species, the medial pallium is defined as a pallial sector adjacent to the cortical hem and roof plate/choroid tela, showing moderate to strong ventricular zone expression of Lef1, Lhx2, and Lhx9, but not Lhx5. Based on this, the hippocampal formation (indusium griseum, dentate gyrus, Ammon's horn fields, and subiculum), the medial entorhinal cortex, and part of the amygdalo-hippocampal transition area of mouse appeared to derive from the medial pallium. In the chicken, based on the same position and gene expression profile, we propose that the hippocampus (including the V-shaped area), the parahippocampal area (including its caudolateral part), the entorhinal cortex, and the amygdalo-hippocampal transition area are medial pallial derivatives. Moreover, the combinatorial expression of Lef1, Prox1, Lmo4, and Lmo3 allowed the identification of dentate gyrus/CA3-like, CA1/subicular-like, and medial entorhinal-like comparable sectors in mouse and chicken, and point to the existence of mostly conserved molecular networks involved in hippocampal complex development. Notably, while the mouse medial entorhinal cortex derives from the medial pallium (similarly to the hippocampal formation, both being involved in spatial navigation and spatial memory), the lateral entorhinal cortex (involved in processing non-spatial, contextual information) appears to derive from a distinct dorsolateral caudal pallial sector.
The comparison of gene expression patterns in the embryonic brain of mouse and chicken is being essential for understanding pallial organization. However, the scarcity of gene expression data in reptiles, crucial for understanding evolution, makes it difficult to identify homologues of pallial divisions in different amniotes. We cloned and analyzed the expression of the genes Emx1, Lhx2, Lhx9, and Tbr1 in the embryonic telencephalon of the lacertid lizard Psammodromus algirus. The comparative expression patterns of these genes, critical for pallial development, are better understood when using a recently proposed six‐part model of pallial divisions. The lizard medial pallium, expressing all genes, includes the medial and dorsomedial cortices, and the majority of the dorsal cortex, except the region of the lateral cortical superposition. The latter is rich in Lhx9 expression, being excluded as a candidate of dorsal or lateral pallia, and may belong to a distinct dorsolateral pallium, which extends from rostral to caudal levels. Thus, the neocortex homolog cannot be found in the classical reptilian dorsal cortex, but perhaps in a small Emx1‐expressing/Lhx9‐negative area at the front of the telencephalon, resembling the avian hyperpallium. The ventral pallium, expressing Lhx9, but not Emx1, gives rise to the dorsal ventricular ridge and appears comparable to the avian nidopallium. We also identified a distinct ventrocaudal pallial sector comparable to the avian arcopallium and to part of the mammalian pallial amygdala. These data open new venues for understanding the organization and evolution of the pallium.
Evidence accumulated over the last few decades demonstrates that all reptiles examined thus far continue to add neurons at a high rate and in many regions of the adult brain. This so-called adult neurogenesis has been described in the olfactory bulbs, rostral forebrain, all cortical areas, anterior dorsal ventricular ridge, septum, striatum, nucleus sphericus, and cerebellum. The rate of neuronal production varies greatly among these brain areas. Moreover, striking differences in the rate and distribution of adult neurogenesis have been noted among species. In addition to producing new neurons in the adult brain, lizards, and possibly other reptiles as well, are capable of regenerating large portions of their telencephalon damaged as a result of experimentally-induced injuries, thus exhibiting an enormous potential for neuronal regeneration. Adult neurogenesis and neuronal regeneration take advantage of the same mechanisms that are present during embryonic neurogenesis. New neurons are born in the ependyma lining the ventricles and migrate radially through the brain parenchyma along processes of radial glial cells. Several lines of evidence suggest that radial glial cells also act as stem cells for adult neurogenesis. Once they reach their final destination, the young neurons extend axons that reach appropriate target areas. Tangential migration of neurons alongside the ventricular ependyma has also been reported. Most of these tangentially migrating neurons seem to be destined for the olfactory bulbs and are, thus, part of a system similar to the mammalian rostral migratory stream. The proliferation and recruitment of new neurons appear to result in continuous growth of most areas showing adult neurogenesis. The functional consequences of this continuous generation and integration of new neurons into existing circuits is largely conjectural, but involvement of these phenomena in learning and memory is one likely possibility.
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