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The amygdaloid complex plays a crucial role in socio-emotional conduct, learning, survival, and reproductive behaviors. It is constituted by a set of nuclei presenting a great cellular heterogeneity and embryonic origin diversity (pallial, subpallial, and even extra-telencephalic). In the last two decades, the tetrapartite pallial paradigm defined the pallial portion of the amygdala as a derivative of the lateroventral pallium. However, the pallial conception is currently being reanalyzed and one of these new proposals is to consider the mouse pallial amygdala as a radial histogenetic domain independent from the rest of the pallial subdomains. In anamniotes, and particularly in amphibian anurans, the amygdaloid complex was described as a region with pallial and subpallial components similar to those described in amniotes. In the present study carried out in Xenopus laevis, after a detailed analysis of the orientation of the amygdalar radial glia, we propose an additional amygdala derived from the pallial region. It is independent of the vomeronasal/olfactory amygdaloid nuclei described in anurans, expresses markers such as Lhx9 present in the mammalian pallial amygdala, and lacks Otp-expressing cells, detected in the adjacent medial amygdala. Further studies are needed to clarify the functional involvement of this area, and whether it is a derivative of the adjacent ventral pallium or an independent pallial domain. [ABSTRACT FROM AUTHOR]

Cajal-Retzius cells are essential for cortical development in mammals, and their involvement in the evolution of this structure has been widely postulated, but very little is known about their progenitor domains in non-mammalian vertebrates. Using in situhybridization and immunofluorescence techniques we analyzed the expression of some of the main Cajal-Retzius cell markers such as Dbx1, Ebf3, ER81, Lhx1, Lhx5, p73, Reelin, Wnt3a, Zic1, and Zic2 in the forebrain of the anuran Xenopus laevis, because amphibians are the only class of anamniote tetrapods and show a tetrapartite evaginated pallium, but no layered or nuclear organization. Our results suggested that the Cajal-Retzius cell progenitor domains were comparable to those previously described in amniotes. Thus, at dorsomedial telencephalic portions a region comparable to the cortical hem was defined in Xenopus based on the expression of Wnt3a, p73, Reelin, Zic1, and Zic2. In the septum, two different domains were observed: a periventricular dorsal septum, at the limit between the pallium and the subpallium, expressing Reelin, Zic1, and Zic2, and a related septal domain, expressing Ebf3, Zic1, and Zic2. In the lateral telencephalon, the ventral pallium next to the pallio-subpallial boundary, the lack of Dbx1 and the unique expression of Reelin during development defined this territory as the most divergent with respect to mammals. Finally, we also analyzed the expression of these markers at the prethalamic eminence region, suggested as Cajal-Retzius progenitor domain in amniotes, observing there Zic1, Zic2, ER81, and Lhx1 expression. Our data show that in anurans there are different subtypes and progenitor domains of Cajal-Retzius cells, which probably contribute to the cortical regional specification and territory-specific properties. This supports the notion that the basic organization of pallial derivatives in vertebrates follows a comparable fundamental arrangement, even in those that do not have a sophisticated stratified cortical structure like the mammalian cerebral cortex. [ABSTRACT FROM AUTHOR]

In the developing cerebral cortex, how progenitors that seemingly display limited diversity end up producing a vast array of neurons remains a puzzling question. The prevailing model suggests that temporal maturation of progenitors is a key driver in the diversification of the neuronal output. However, temporal constraints are unlikely to account for all diversity, especially in the ventral and lateral pallium where neuronal types significantly differ from their dorsal neocortical counterparts born at the same time. In this study, we implemented single-cell RNAseq to sample the diversity of progenitors and neurons along the dorso-ventral axis of the early developing pallium. We first identified neuronal types, mapped them on the tissue and determined their origin through genetic tracing. We characterised progenitor diversity and disentangled the gene modules underlying temporal versus spatial regulations of neuronal specification. Finally, we reconstructed the developmental trajectories followed by ventral and dorsal pallial neurons to identify lineage-specific gene waves. Our data suggest a model by which discrete neuronal fate acquisition from a continuous gradient of progenitors results from the superimposition of spatial information and temporal maturation. [ABSTRACT FROM AUTHOR]

The amygdala in mammals plays a key role in emotional processing and learning, being subdivided in pallial and subpallial derivatives. Recently, the cortical ring model and the pallial amygdalar radial model (Puelles et al. 2019; Garcia-Calero et al. 2020) described the pallial amygdala as an histogenetic field external to the allocortical ring, and subdivided it in five major radial domains called lateral, basal, anterior, posterior and retroendopiriform units. The anterior radial unit, whose cells typically express the Lhx9 gene (see molecular profile in Garcia-Calero et al. 2020), is located next to the pallial/subpallial boundary. This radial domain shows massive radial translocation and accumulation of its derivatives into its intermediate and superficial strata, with only a glial palisade representing its final periventricular domain. To better understand the development of this singular radial domain, not described previously, we followed the expression of Lhx9 during mouse amygdalar development in the context of the postulated radial subdivisions of the pallial amygdala and other telencephalic developmental features. [ABSTRACT FROM AUTHOR]

The neocortex (NCx) generates at the dorsal region of the pallium in the forebrain. Several adjacent structures also contribute with neurons to NCx. Ventral pallium (VP) is considered to generate several populations of neurons that arrive through tangential migration to the NCx. Amongst them are the Cajal-Retzius cells and some transient pyramidal neurons. However, the specific site and timing of generation, trajectory of migration and actual contribution to the pyramidal population remains elusive. Here, we investigate the spatio-temporal origin of neuronal populations from VP in an in vivo model, using a transposase mediated in utero electroporation method in embryonic mouse. From E11 to E14 cells born at the lateral corner of the neocortical neuroepithelium including the VP migrated ventro-laterally to settle all areas of the ventral telencephalon. Specifically, neurons migrated into amygdala (Ag), olfactory cortices, and claustrum (Cl). However, we found no evidence for any neurons migrating tangentially toward the NCx, regardless the antero-posterior level and developmental time of the electroporation. Our results challenge the described ventral-pallial origin of the transient pyramidal neuron population. In order to find the exact origin of cortical neurons that were previously Dbx1-fate mapped we used the promoter region of the murine Dbx1 locus to selectively target Dbx1-expressing progenitors and label their lineage. We found these progenitors in low numbers in all pallial areas, and not only in the ventral pallial ventricular zone. Our findings on the local cortical origin of the Dbx1-derived pyramidal neurons reconcile the observation of Dbx1-derived neurons in the cortex without evidence of dorsal tangential migration from VP and provide a new framework for the origin of the transient Dbx1-derived pyramidal neuron population. We conclude that these neurons are born locally within the dorsal pallial neuroepithelium.

In rodents, the medial nucleus of the amygdala receives direct inputs from the accessory olfactory bulbs and is mainly implicated in pheromone-mediated reproductive and defensive behaviors. The principal neurons of the medial amygdala are GABAergic neurons generated principally in the caudo-ventral medial ganglionic eminence and preoptic area. Beside GABAergic neurons, the medial amygdala also contains glutamatergic Otp-expressing neurons cells generated in the lateral hypothalamic neuroepithelium and a non-well characterized Pax6-positive population. In the present work, we describe a novel glutamatergic Ebf3-expressing neuronal subpopulation distributed within the periphery of the postero-ventral medial amygdala. These neurons are generated in a pallial domain characterized by high expression of Gdf10. This territory is topologically the most caudal tier of the ventral pallium and accordingly, we named it Caudo-Ventral Pallium (CVP). In the absence of Pax6, the CVP is disrupted and Ebf3-expressing neurons fail to be generated. Overall, this work proposes a novel model of the neuronal composition of the medial amygdala and unravels for the first time a new novel pallial subpopulation originating from the CVP and expressing the transcription factor Ebf3. [ABSTRACT FROM AUTHOR]

ABSTRACT Studies on gene expression in the developing claustrum of the mouse have clarified the relationships and identity of the claustrum proper and related endopiriform nuclei. The cells of the claustrum primordium express Nr4a2; they are formed in combination with the Nr4a2-labeled subplate cells in the lateral pallium at the site of the future insular cortex. The insular cortex cells, which are born later and which are Nr4a2-negative, migrate through the claustrum toward the pial surface to form layers (2-6a) of the insular cortex. The claustrum is made up of distinct deep (subplate-like) and superficial (principal) parts. The cells of the dorsal endopiriform nucleus (which are also Nr4a2-positive) are formed in the deep part of the claustrum primordium in the lateral pallium, but they migrate ventrally to reach the ventral pallium deep to the piriform cortex at E14.5 in the mouse. On the other hand, the ventral endopiriform nucleus is formed by radially migrating Nr4a2-negative cells in the ventral pallium; it is therefore developmentally distinct from the Nr4a2-postive dorsal endopiriform nucleus, which is a lateral pallial derivative. J. Comp. Neurol. 525:1499-1508, 2017. © 2016 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

The progeny of Dbx1 -expressing progenitors was studied in the developing mouse pallium, using two transgenic mouse lines: (1) Dbx1 nlslacZ mice, in which the gene of the β-galactosidase reporter (LacZ) is inserted directly under the control of the Dbx1 promoter, allowing short-term lineage tracing of Dbx1 -derived cells; and (2) Dbx1 CRE mice crossed with a Cre-dependent reporter strain ( ROSA26 loxP-stop-loxP-LacZ ), in which the Dbx1 -derived cells result permanently labeled ( Bielle et al., 2005 ). We thus examined in detail the derivatives of the postulated longitudinal ventral pallium (VPall) sector, which has been defined among other features by its selective ventricular zone expression of Dbx1 (the recent ascription by Puelles, 2014 of the whole olfactory cortex primordium to the VPall was tested). Earlier notions about a gradiental caudorostral reduction of Dbx1 signal were corroborated, so that virtually no signal was found at the olfactory bulb and the anterior olfactory area. The piriform cortex was increasingly labeled caudalwards. The only endopiriform grisea labeled were the ventral endopiriform nucleus and the bed nucleus of the external capsule. Anterior and basolateral parts of the whole pallial amygdala also were densely marked, in contrast to the negative posterior parts of these pallial amygdalar nuclei (leaving apart medial amygdalar parts ascribed to subpallial or extratelencephalic sources of Dbx1 -derived GABAergic and non-GABAergic neurons). Alternative tentative interpretations are discussed to explain the partial labeling obtained of both olfactory and amygdaloid structures. This includes the hypothesis of an as yet undefined part of the pallium, potentially responsible for the posterior amygdala, or the hypothesis that the VPall may not be wholly characterized by Dbx1 expression (this gene not being necessary for VPall molecular distinctness and histogenetic potency), which would leave a dorsal Dbx1 -negative VPall subdomain of variable size that might contribute partially to olfactory and posterior amygdalar structures. [ABSTRACT FROM AUTHOR]

The transcription factor Nr4a2 was recently revealed as a very early developmental marker of the claustrum (CL) proper in the mouse. The earliest claustral primordium was identified superficially, dorsal to the olfactory cortex, and was subsequently covered by the Nr4a2-negative cells of the insular cortex. Some tangentially migrating claustral derivatives (subplate cells and some endopiriform elements) also expressed this marker. The present study employs the same genetic marker to explore the presence of a comparable pallial division in chicken in which, in principle, the same pallial sectors exist as in mammals. We were indeed able to delineate an early-developing Nr4a2-positive mantle domain at the expected topologic position within the developing chicken lateral pallium. In the chicken as well as in the turtle (from data in the literature), the earliest postmitotic lateropallial cells likewise express Nr4a2 and occupy a corticoid superficial stratum of the mesopallium, which is clearly comparable in spatial and chronological profile to the mouse CL. Other cells produced in this pallial sector include various tangentially migrating Nr4a2-labeled derivatives as well as Nr4a2-negative and Nr4a2-positive local deeper subpopulations that partially interdigitate, forming mesopallial core and shell populations. We hold that the deep avian and reptilian mesopallial formation developing under the superficial corticoid CL homolog represents a field homolog of the insula, although additional studies are required to underpin this hypothesis. [ABSTRACT FROM AUTHOR]

This article proposes a comprehensive view of the origin of the mammalian brain. We discuss i) from which region in the brain of a reptilian-like ancestor did the isocortex originate, and ii) the origin of the multilayered structure of the isocortex from a simple-layered structure like that observed in the cortex of present-day reptiles. Regarding question i there have been two alternative hypotheses, one suggesting that most or all the isocortex originated from the dorsal pallium, and the other suggesting that part of the isocortex originated from a ventral pallial component. The latter implies that a massive tangential migration of cells from the ventral pallium to the dorsal pallium takes place in isocortical development, something that has not been shown. Question ii refers to the origin of the six-layered isocortex from a primitive three-layered cortex. It is argued that the superficial isocortical layers can be considered to be an evolutionary acquisition of the mammalian brain, since no equivalent structures can be found in the reptilian brain. Furthermore, a characteristic of the isocortex is that it develops according to an inside-out neurogenetic gradient, in which late-produced cells migrate past layers of early-produced cells. It is proposed that the inside-out neurogenetic gradient was partly achieved by the activation of a signaling pathway associated with the Cdk5 kinase and its activator p35, while an extracellular protein called reelin (secreted in the marginal zone during development) may have prevented migrating cells from penetrating into the developing marginal zone (future layer I).

The three-layered piriform cortex, an integral part of the olfactory system, processes odor information relayed by olfactory bulb mitral cells. Specifically, mitral cell axons form the lateral olfactory tract (LOT) by targeting lateral olfactory tract (lot) guidepost cells in the piriform cortex. While lot cells and other piriform cortical neurons share a pallial origin, the factors that specify their precise phenotypes are poorly understood. Here we show that in mouse, the proneural genes Neurog1 and Neurog2 are coexpressed in the ventral pallium, a progenitor pool that first gives rise to Cajal-Retzius (CR) cells, which populate layer I of all cortical domains, and later to layer II/III neurons of the piriform cortex. Using loss-of-function and gain-of-function approaches, we find that Neurog1 has a unique early role in reducing CR cell neurogenesis by tempering Neurog2's proneural activity. In addition, Neurog1 and Neurog2 have redundant functions in the ventral pallium, acting in two phases to first specify a CR cell fate and later to specify layer II/III piriform cortex neuronal identities. In the early phase, Neurog1 and Neurog2 are also required for lot cell differentiation, which we reveal are a subset of CR neurons, the loss of which prevents mitral cell axon innervation and LOT formation. Consequently, mutation of Trp73, a CR-specific cortical gene, results in lot cell andLOTaxon displacement. Neurog1 and Neurog2 thus have unique and redundant functions in the piriform cortex, controlling the timing of differentiation of early-born CR/lot cells and specifying the identities of later-born layer II/III neurons. [ABSTRACT FROM AUTHOR]

The cytoarchitecture of the telencephalon of the Comoran coelacanth, Latimeria chalumnae, was analyzed in the context of recent advances in our understanding of telencephalic organization in lungfishes and amphibians, which constitute the sister group to coelacanths. In coelacanths, the telencephalon is divided into pendunculated olfactory bulbs, paired hemispheres, and an unevaginated telencephalon impar. The hemispheres consist of a ventrally located subpallium and, dorsally, a greatly expanded pallium. Traditionally, the subpallium in coelacanths has been divided into a medial septal area and a lateral striatum. Re-examination of the lateral subpallial wall, however, suggests that the striatum is more restricted than previously believed, and it is replaced dorsally by a more scattered plate of cells, which appears to represent the ventral pallium. The putative ventral pallium is continuous with a ventromedial pallial formation, which appears to receive input from the lateral olfactory tract and should be considered a possible homologue of the lateral pallium in tetrapods. The putative lateral pallium is replaced by a more dorsomedial pallial formation, which may represent the dorsal pallium. This formation is replaced, in turn by an extensive lateral pallial formation, which appears to be homologous to the medial pallium of tetrapods. An expanded medial pallium in coelacanths, lepidosirenid lungfishes, and amphibians may be related to well developed spatial learning. Traditionally, the telencephalon impar of coelacanths, has been interpreted as an enlarged preoptic area, but reanalysis indicates that the so-called superior preoptic nucleus actually consists of the medial amygdalar nucleus.

Observations accruing in recent years imply that the areal patterning and size dimensioning of the mammalian neocortex are influenced by diverse sets of tangentially migrating glutamatergic neurons that invade the cortical plate and, in so doing, modify the properties of the neopallial proliferative compartments. This developmental scenario sheds new light upon the old issue of how the mammalian neocortex evolved its more complex structure from nonmammalian antecedent forms. In reviewing these novelties, I first point out the topological position of the neopallial island as a central component of the pallium in all gnathostomes, surrounded by a ring of prospective allocortical pallial regions and a more distant set of peripheral neighboring forebrain areas. Early patterning arises from the periphery via passive planar signaling. This process probably establishes the pallium field and its basic island plus allocortical ring organization, as well as a rough prepatterning of some regional subareas. Afterwards, patterning and modulated growth are also actively influenced by the convergence of separate streams of tangentially migrating subpial cells (partly peripheral and partly allocortical in origin) which collectively form the Cajal-Retzius neuronal population in layer I. Effects of these cells include the inside-out stratification of the cortical plate and they may also contribute to the evolutionary emergence of the 6-layered neocortical structure. The most recent addition to our knowledge of pallio-pallial migrations is the existence of a subsequent deep tangential migration of ventropallial cells into the neopallial primordium, whose signaling influence upon local progenitors magnifies the cortex population by 20%. These glutamatergic cells dispersedly invade the entire cortex but largely die postnatally. The crucial implications of these data for comparative thinking on mammalian neocortex evolution and interpretation of potential homologs in sauropsids are explored. Finally, a new conjecture regarding a possible role of the hitherto disregarded lateral pallium is advanced. Copyright © 2011 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

Abstract: The neocortex is the most representative and elaborated structure of the mammalian brain and is related to the achievement of complex cognitive capabilities, which are disturbed following malformation or lesion. Searching for the evolutionary origin of this structure continues to be one of the most important and challenging questions in comparative neurobiology. However, this is extremely difficult because of the highly divergent evolution of the pallium in different vertebrates, which has obscured the comparison. Herein, we review developmental neurobiology data for trying to understand the genetic factors that define and underlie the parcellation of homologous pallial subdivisions in different vertebrates. According to these data, the pallium in all tetrapods parcellates during development into four major histogenetic subdivisions, which are homologous as fields across species. The neocortex derives from the dorsal pallium and, as such, is only comparable to the sauropsidian dorsal pallium (avian hyperpallium and lizard/turtle dorsal cortex). We also tried to identify developmental changes in phylogeny that may be responsible of pallial divergent evolution. In particular, we point out to evolutionary differences regarding the cortical hem (an important signaling center for pallial patterning, that also is a source of Cajal–Retzius cells, which are involved in cortical lamination), which may be behind the distinct organization of the pallium in mammals and non-mammals. In addition, we mention recent data suggesting a correlation between the appearance and elaboration of the subventricular zone (a new germinative cell layer of the developing neocortex), and the evolution of novel cell layers (the supragranular layers) and interneuron subtypes. Finally, we comment on epigenetic factors that modulate the developmental programs, leading to changes in the formation of functional areas in the pallium (within some constraints). [Copyright &y& Elsevier]

Abstract: In this study, we review data on the existence of comparable divisions and subdivisions in the telencephalon of different groups of tetrapods based on expression of some developmental regulatory genes, having a particular focus in the comparison of the anuran amphibian Xenopus and the mouse. The available data on Xenopus, mouse, chick and turtle indicate that apparently all tetrapod groups possess the same molecularly distinct divisions and subdivisions in the telencephalon. This basic organization was likely present in the telencephalon of stem tetrapods. Each division/subdivision is characterized by expression of a unique combination of developmental regulatory genes, and appears to represent a self-regulated and topologically constant histogenetic brain compartment that gives rise to specific groups of cells. This interpretation has an important consequence for searching homologies, since a basic condition for cell groups in different vertebrates to be considered homologous is that they originate in the same compartment. However, evolution may allow individual cell groups derived from comparable (field homologous) subdivisions to be either similar or dissimilar across the vertebrate groups, giving rise to several possible scenarios of evolution, which include both the evolutionary conservation of similar (homologous) cells or the production of novel cell groups. Finally, available data in the lamprey, a jawless fish, suggest that not all telencephalic subdivisions were present at the origin of vertebrates, raising important questions about their evolution. [Copyright &y& Elsevier]

We showed previously that the orphan nuclear receptor Tlx is required for the correct establishment of the pallio-subpallial boundary. Loss of Tlx results in a dorsal expansion of ventral markers (e.g., the homeodomain protein GSH2) into the ventralmost pallial region, i.e., the ventral pallium. We also observed a disproportionate reduction in the size of the Tlx mutant lateral ganglionic eminence (LGE) from embryonic day 14.5 onward. Here we show that this reduction is caused, at least in large part, by a proliferation defect. Interestingly, in Tlx mutants, the LGE derivatives are differentially affected. Although the development of the Tlx mutant striatum is compromised, an apparently normal number of olfactory bulb interneurons are observed. Consistent with this observation, we found that Tlx is required for the normal establishment of the ventral LGE that gives rise to striatal projection neurons. This domain is reduced by the dorsal and ventral expansion of molecular markers normally confined to progenitor domains flanking the ventral LGE. Finally, we investigated possible genetic interactions between Gsh2 and Tlx in lateral telencephalic development. Our results show that, although Gsh2 and Tlx have additive effects on striatal development, they differentially regulate the establishment of ventral pallial identity. [ABSTRACT FROM AUTHOR]

The isocortex is a distinctive feature of mammalian brains, which has no clear counterpart in the cerebral hemispheres of other amniotes. This paper speculates on the evolutionary processes giving rise to the isocortex. As a first step, we intend to identify what structure may be ancestral to the isocortex in the reptilian brain. Then, it is necessary to account for the transformations (developmental, connectional, and functional) of this ancestral structure, which resulted in the origin of the isocortex. One long-held perspective argues that part of the isocortex derives from the ventral pallium of reptiles, whereas another view proposes that the isocortex originated mostly from the dorsal pallium. We consider that, at this point, evidence tends to favor correspondence of the isocortex with the dorsal cortex of reptiles. In any case, the isocortex may have originated partly as a consequence of an overall "dorsalizing" effect (that is, an expansion of the territories expressing dorsal-specific genes) during pallial development. Furthermore, expansion of the dorsal pallium may have been driven by selective pressures favoring the development of associative networks between the dorsal cortex, the olfactory cortex, and the hippocampus, which participated in spatial or episodic memory in the early mammals. In this context, sensory projections that in reptiles end in the ventral pallium, are observed to terminate in the isocortex (dorsal pallium) of mammals, perhaps owing to their participation in these associative networks. [ABSTRACT FROM AUTHOR]

The present paper describes the distribution of three calcium-binding proteins (calbindin D28k, calretinin, and parvalbumin) in the mouse dorsal claustrum and endopiriform nucleus. The three calcium-binding proteins were distinctly expressed in structures of both the claustrum and the endopiriform nucleus. Calbindin was the calcium-binding protein showing the highest expression in the claustrum and the endopiriform nucleus. In contrast, calretinin-immunoreactive structures, particularly cell bodies, were very scarce in these regions. Both calbindin-immunoreactive and parvalbumin-immunoreactive neurons were more abundant in the claustrum than in the endopiriform nucleus, and more in rostral than in caudal levels. Nevertheless, calcium-binding protein immunoreactive neurons constitute a minority population of claustral neurons. The colocalization study of calbindin and parvalbumin immunoreactivities has demonstrated that both calcium-binding proteins are mostly expressed by separate claustral neurons in the mouse. On the other hand, our results on parvalbumin and calretinin immunoreactivity match a novel subdivision of the mouse claustrum mostly based on the pattern of cadherin expression [Neuroscience 106 (2001) 505]. In this sense, we propose that a specific zone of the dorsal claustrum with cell bodies that strongly express Rcad and cadherin-8 would be the selective target for parvalbumin-expressing fibers, and that they would be mostly avoided by calretinin-expressing axons. [Copyright &y& Elsevier]

The isocortex is a distinctive feature of the mammalian brain, with no clear counterpart in other amniotes. There have been long controversies regarding possible homologues of this structure in reptiles and birds. The brains of the latter are characterized by the presence of a structure termed dorsal ventricular ridge (DVR), which receives ascending auditory and visual projections, and has been postulated to be homologous to parts of the mammalian isocortex (i.e., the auditory and the extrastriate visual cortices). Dissenting views, now supported by molecular evidence, claim that the DVR originates from a region termed ventral pallium, while the isocortex may arise mostly from the dorsal pallium (in mammals, the ventral pallium relates to the claustroamygdaloid complex). Although it is possible that in mammals the embryonic ventral pallium contributes cells to the developing isocortex, there is no evidence yet supporting this alternative. The possibility is raised that the expansion of the cerebral cortex in the origin of mammals was product of a generalized dorsalizing influence in pallial development, at the expense of growth in ventral pallial regions. Importantly, the evidence suggests that organization of sensory projections is significantly different between mammals and sauropsids. In reptiles and birds, some sensory pathways project to the ventral pallium and others project to the dorsal pallium, while in mammals sensory projections end mainly in the dorsal pallium. We suggest a scenario for the origin of the mammalian isocortex which relies on the development of associative circuits between the olfactory, the dorsal and the hippocampal cortices in the earliest mammals. [Copyright &y& Elsevier]

To further understand the organization and evolution of the telencephalon, we analyzed in the frog Xenopus laevis the expression of the genes Distal-less-4 (Xdll-4, comparable to the mouse gene Dlx2) and GAD-67 (XGAD-67, expressed in GABAergic cells), and compared this with calretinin immunostaining and the cytoarchitecture of the telencephalon. Our results show that like in other vertebrates, the telencephalon of the frog Xenopus is divided into two major territories: a basal, subpallial region showing a high density of cells expressing Xdll-4 and XGAD-67, and a dorsal, pallial region showing only few, dispersed cells expressing these genes. The subpallial territory of the frog Xenopus includes the septum, the amphibian basal ganglia, some basal forebrain cholinergic cell groups and some amygdala nuclei. In the pallium of the frog Xenopus, medial, dorsal, lateral, and ventral parts could be distinguished, similar to those described in amniotes . In summary, the amphibian telencephalon shows a basic morphogenetic organization similar to that of amniotes, which suggests that this organization is common to the telencephalon of all tetrapods. [Copyright &y& Elsevier]

The organization of the cerebral hemispheres of mammals is characterized by corticostriatal glutamatergic projections and striatopallidal GABAergic ones, plus the descending projections of the pallium and subpallium to extratelencephalic targets. The present review of the available neuroanatomical data on the forebrain of lizards suggests that the telencephalon of reptiles also follows this basic pattern of connectivity. In addition, we show that this basic circuitry includes a pallido-cortical projection, therefore forming a cortico-striato-pallido-cortical circuit. The analysis of this circuitry for the medial, dorsal, lateral, and ventral pallial divisions in reptiles and mammals leads to the following conclusions: (1) The medial and dorsal cortices of lizards together appear to be equivalent to the medial pallium of mammals. (2) The projection from the lacertilian dorsal cortex to the striatum proper resembles the subiculo-striatal projection of mammals, rather than the isocortical projection to the caudatus-putamen. (3) Most of the dorsal striatum of reptiles is engaged in the corticostriatal circuit corresponding to the ventral pallium (the anterior dorsal ventricular ridge), and therefore, it is not equivalent to the mammalian caudatus-putamen, which is involved in the circuit of the dorsal pallium. (4) The main and accessory olfactory bulbs also follow this pattern of connections. [Copyright &y& Elsevier]