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This essay re-examines the singular case of the supposedly unique rostrally elongated notochord described classically in amphioxus. We start from our previous observations in hpf 21 larvae [Albuixech-Crespo et al.: PLoS Biol. 2017;15(4):e2001573] indicating that the brain vesicle has rostrally a rather standard hypothalamic molecular configuration. This correlates with the notochord across a possible rostromedian acroterminal hypothalamic domain. The notochord shows some molecular differences that specifically characterize its pre-acroterminal extension beyond its normal rostral end under the mamillary region. We explored an alternative interpretation that the putative extension of this notochord actually represents a variant form of the prechordal plate in amphioxus, some of whose cells would adopt the notochordal typology, but would lack notochordal patterning properties, and might have some (but not all) prechordal ones instead. We survey in detail the classic and recent literature on gastrulation, prechordal plate, and notochord formation in amphioxus, compare the observed patterns with those of some other vertebrates of interest, and re-examine the literature on differential gene expression patterns in this rostralmost area of the head. We noted that previous literature failed to identify the amphioxus prechordal primordia at appropriate stages. Under this interpretation, a consistent picture can be drawn for cephalochordates, tunicates, and vertebrates. Moreover, there is little evidence for an intrinsic capacity of the early notochord to grow rostralwards (it normally elongates caudalwards). Altogether, we conclude that the hypothesis of a prechordal nature of the elongated amphioxus notochord is consistent with the evidence presented., (© 2022 S. Karger AG, Basel.)

In the developing brain, the phenomenon of neurogenesis is manifested heterotopically, that is, much the same neurogenetic steps occur at different places with a different timetable. This is due apparently to early molecular regionalization of the neural tube wall in the anteroposterior and dorsoventral dimensions, in a checkerboard pattern of more or less deformed quadrangular histogenetic areas. Their respective fate is apparently specified by a locally specific combination of active/repressed genes known as "molecular profile." This leads to position-dependent differential control of proliferation, neurogenesis, differentiation, and other aspects, eventually in a heterochronic manner across adjacent areal units with sufficiently different molecular profiles. It is not known how fixed these heterochronic patterns are. We reexamined here comparatively early patterns of forebrain and hindbrain neurogenesis in a lizard (Lacerta gallotia galloti), a bird (the chick), and a mammal (the rat), as demonstrated by activation of acetylcholinesterase (AChE). This is an early marker of postmitotic neurons, which leaves unlabeled the neuroepithelial ventricular cells, so that we can examine cleared wholemounts of the reacted brains to have a birds-eye view of the emergent neuronal pattern at each stage. There is overall heterochrony between the basal and alar plates of the brain, a known fact, but, remarkably, heterochrony occurs even within the precocious basal plate among its final anteroposterior neuromeric subdivisions and their internal microzonal subdivisions. Some neuromeric units or microzones are precocious, while others follow suit without any specific spatial order or gradient; other similar neuromeric units remain retarded in the midst of quite advanced neighbors, though they do produce similar neurogenetic patterns at later stages. It was found that some details of such neuromeric heterochrony are species-specific, possibly related to differential morphogenetic properties. Given the molecular causal underpinning of the updated prosomeric model used here for interpretation, we comment on the close correlation between some genetic patterns and the observed AChE differentiation patterns., (© 2022 S. Karger AG, Basel.)

This essay reviews step by step the conceptual changes of the updated tetrapartite pallium model from its tripartite and early tetrapartite antecedents. The crucial observations in mouse material are explained first in the context of assumptions, tentative interpretations, and literature data. Errors and the solutions offered to resolve them are made explicit. Next, attention is centered on the lateral pallium sector of the updated model, whose definition is novel in incorporating a claustro-insular complex distinct from both olfactory centers (ventral pallium) and the isocortex (dorsal pallium). The general validity of the model is postulated at least for tetrapods. Genoarchitectonic studies performed to check the presence of a claustro-insular field homolog in the avian brain are reviewed next. These studies have indeed revealed the existence of such a complex in the avian mesopallium (though stratified outside-in rather than inside-out as in mammals), and there are indications that the same pattern may be found in reptiles as well. Peculiar pallio-pallial tangential migratory phenomena are apparently shared as well between mice and chicks. The issue of whether the avian mesopallium has connections that are similar to the known connections of the mammalian claustro-insular complex is considered next. Accrued data are consistent with similar connections for the avian insula homolog, but they are judged to be insufficient to reach definitive conclusions about the avian claustrum. An aside discusses that conserved connections are not a necessary feature of field-homologous neural centers. Finally, the present scenario on the evolution of the pallium of sauropsids and mammals is briefly visited, as highlighted by the updated tetrapartite model and present results., (© 2017 S. Karger AG, Basel.)

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.)

Ten years ago, we published the first detailed prosomeric map of the forebrain in lampreys. In the interim, the prosomeric model has been modified and simplified to better explain numerous data on the expression patterns of regulatory genes, as well as data from chemical, hodological and neuroembryological experiments, mostly in amniote vertebrates. In this report we first review the main historical concepts of lamprey forebrain organization, relating them to either columnar- or segmental-influenced models and explicit or implicit axial references. Next, our previous prosomeric model of the lamprey forebrain is updated, postulating some new hypotheses on the organization of the secondary prosencephalon., (2009 S. Karger AG, Basel)

The work examines frog diencephalic subdivisions from a segmental viewpoint and adds a number of details to the atlas of the bullfrog diencephalon reported by Neary and Northcutt [1983]. Acetylcholinesterase histochemistry was performed on brains of Rana perezi, sectioned either sagittally or in a plane roughly parallel to the optic tract to optimize detection of segmented landmarks. This material provided a very detailed picture of individual neurons, neuropils and some fiber tracts expressing the enzyme in diverse patterns characteristic for each diencephalic region. The main diencephalic areas previously recognized in the bullfrog appeared subdivided into smaller AChE-chemoarchitectonic units. Modified subdivisions are proposed for several entities: preoptic, suprachiasmatic, entopeduncular, ventral thalamic, anterior thalamic, pretectal, hypothalamic and tuberculum posterior regions. A number of cell groups are described for the first time in frogs. This mapping is expected to be useful for the interpretation of immunocytochemical and experimental hodologic results in the diencephalon of frogs and opens new possibilities for comparative analysis.

The idea is presented that modern segmental thinking on the forebrain is best conceived of as a scientific paradigm, which is revealing itself to be increasingly advantageous for the interpretation of both causal developmental/genetic data and pitfall-free morphology. Various related or opposed viewpoints are considered briefly, emphasizing their respective assumptions with regard to longitudinal and transverse subdivisions.

Afferents to the habenular complex were studied by means of in vitro horseradish peroxidase retrograde labeling and anterograde control experiments in the lizard Gallotia galloti. The medial habenular nucleus was found to receive abundant afferent fibers from the nucleus of the posterior pallial commissure and the nucleus septalis impar. More restricted input comes from the nucleus eminentiae thalami and the nucleus of the stria medullaris. The lateral habenular nucleus is innervated by various fiber groups originating from the bed nucleus of the anterior commissure, the diagonal band nucleus, the lateral preoptic area, the anterior entopeduncular nucleus, the lateral hypothalamic and mammillary areas, the nucleus of the stria medullaris, the area tegmentalis ventralis and a scattered neuronal subpopulation in the large-celled dorsolateral nucleus of the dorsal thalamus. Habenulopetal fibers generally follow the stria medullaris, but hypothalamic, entopeduncular and dorsal thalamic afferents course through the dorsal peduncle of the lateral forebrain bundle in a transthalamic route. Mesencephalic ventral tegmental afferents ascend through the tractus retroflexus.

In order to test the usefulness of the in vitro HRP-labeling technique in the brains of small lizards, the fasciculus retroflexus was labeled in isolated brains of young specimens of the lizard Gallotia galloti by means of HRP applied with the tip of a micropipette located approximately midway along the descending course of the tract. Cells in the medial and lateral habenular nuclei were labeled retrogradely. Anterograde transport showed the course of the fasciculus retroflexus, first dorsoventrally into the paramedian prerubral tegmentum and then longitudinally into the isthmic nucleus interpeduncularis. Some fibers spread dorsally into the mesencephalic and isthmic tegmental fields. A substantial bundle continues caudally into nucleus raphe parvocellularis. Other fibers diverge rostral-wards from the point of inflexion of the fasciculus, coursing longitudinally in the medial forebrain bundle into the basal telencephalon. Some fibers course rostrally from the nucleus habenularis lateralis in the stria medullaris and could be followed to the dorsal septum.