PMC:4429232 / 23507-35292 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/4429232","sourcedb":"PMC","sourceid":"4429232","source_url":"https://www.ncbi.nlm.nih.gov/pmc/4429232","text":"Similarity in gene expression between PC and neocortical layers II/III and a hypothesis of their evolutionary relationships\nAccording to our hypothesis, the UL neurons of the neocortex may have sister cell types in other pallial regions. To gain insight on this issue and to identify the best candidate regions whose developmental program may have been co-opted in the evolution of the UL, we performed an analysis in the in situ hybridization database of the Mouse Allen Brain Atlas (Lein et al., 2007). In this analysis we compared the lists of the first 500 genes that were enriched relative to the rest of the CNS (contrast structure, gray) in each of the following regions: neocortical layers II/III, layer IV, layer V/VI, piriform cortex, subiculum and cortical subplate (claustro-amygdaloid complex, and endopiriform nucleus; Figure 3, Supplementary data sheet 1A). Layer II/III is closely related to layer IV, with 352 co-expressed genes, and relatively well correlated with layer V/VI, with 250 co-expressed genes (Table 1, Figure 3, Supplementary data sheet 1B, Supplementary Figure 1). Surprisingly, layers II/III cells also shared about 208 enriched genes with the piriform cortex (42%; Table 1, Figures 3, 4, Supplementary Figure 1). The layers V/VI were less related to the piriform cortex with only 143 co-expressed genes (29%; Table 1, Figures 3, 4, Supplementary Figure 1). In addition only 41 genes were exclusively enriched in piriform cortex and layers V/VI but not in layers II/III (29% of PC and layer V/VI common genes). By contrast 106 genes were specifically enriched only in layer II/III and in piriform cortex but not in layer V/VI (51% of PC and layer II/III common genes; Figures 3, 4, Supplementary data sheet 1C). These striking similarities in the gene expression profile raise the intriguing possibility that the developmental program that provided the base for the evolution of the neocortical layers II/III have been that of the olfactory cortex.\nFigure 3 Comparison of gene expression in PC and NC. Representative coronal sections from the Allen Brain Atlas showing the expression of few representative genes selectively enriched in the UL and PC (A–D), in DL and PC (E,F) in DL, UL and PC (G) or only in DL (H). Please note that CUX2, was not present in the list of genes enriched in PC relative to the rest of the gray matter likely because of its relatively widespread expression in the CNS.\nTable 1 Percentage of shared genes among the first 500 genes enriched in each of the indicated pallial sub-regions.\nPercentage of gene co-expression in different pallial regions\nPC NC II/III NC IV NC V/VI C. Sub Subic.\nPC 42 30 29 37 16\nNC II/III 42 70 49 21 16\nNC IV 30 70 51 13 11\nNC V/VI 29 49 51 20 26\nC. Sub 37 21 13 20 38\nSubic 16 16 11 26 38\nPC, Piriform cortex; NC, Neocortex; C. Sub, Cortical subplate; Subic., Subiculum.\nFigure 4 Comparison of genes shared by piriform cortex and laiers II/III or V/VI of the neocortex. The pie chart represents the 500 genes enriched in the piriform cortex. Each sector indicate the number of genes shared only with layers V/VI (violet), only with layer II/III (red) or with both layer II/III and V/VI (green). The fraction of genes that are not enriched neither in layer II/III nor in layer V/VI are in blue. This idea is not new and dates back to the beginning of twentieth century. Indeed, early neuroanatomist proposed that a primordium of the neocortex may be found in the superposition of lateral and dorsal cortex, the so called superpositio lateralis (Figure 5; Kappers and Theunissen, 1908; Kappers, 1909; De Lange, 1911; Schepers, 1948). This superposition is observed only in some species and its extension correlates with the development of the olfactory system (Ulinsky, 1990). Given that most of the increased encephalization of the first mammaliaformes was due to a huge expansion of the olfactory bulbs and olfactory cortex (Rowe et al., 2011), a substantial development of the lateral superposition could have been present in these species and preceded the emergence of the neocortex. In these superpositions the medial edge of the lateral cortex is located on top of the lateral edge of the dorsal cortex giving rise to a rudimentary six layered arrangement (Figure 5A). Indeed, this region have a dense layer II on top of a sparser layer III that are continuous with the olfactory cortex and receive a direct projection from the olfactory bulb (Minelli, 1967; Regidor, 1977; Desan, 1984; Martinez-Garcia et al., 1991), it posses a parvo-cellular layer IV that receive thalamic inputs (Bruce and Butler, 1984; Desan, 1984; Desfilis et al., 2002) and two deep cellular layers (V and VI) that project to various subcortical targets (Minelli, 1967; Ebner, 1976; Regidor, 1977; Desan, 1984; Hoogland and Vermeulen-Vanderzee, 1989). This layer arrangement is closely reminding that of the neocortex and in particular of the lateral peri-allocortical regions (insular and perirhinal cortices; Figure 2A) which receive olfactory information, lacks layer IV and have a dense layer II that is continuous with the priform cortex (Sanides, 1969; Sanides and Sanides, 1972; Shipley and Geinisman, 1984; Haberly, 1990). Previous authors also highlighted the presence of numerous allocortical features in the peri-allocortical ring, and accordingly proposed that it represent the more ancestral part of the neocortex (Abbie, 1940, 1942; Sanides, 1969; Sanides and Sanides, 1972). The cytoarchitectural similarities between the lateral superposition and the neocortex include also their relationships with bordering regions. In particular, on the medial side the deeper layers of the superposition are in continuity with the non-superposed part of the dorsal cortex while DL of the neocortex are in continuity with the subiculum. An homology between the non-superposed part of the dorsal cortex and the subiculum has been previously proposed (Hoogland and Vermeulen-Vanderzee, 1989). Moreover, this latter region share many features with the DL (Ishizuka, 2001) and accordingly our analyses in the Allen Brain Atlas indicated that it has more enriched genes in common with layer V/VI than layer II/III (26 and 16%, respectively, Table 1, Supplementary Figure 1).\nFigure 5 Model for the origin of the neocortex from the superposition of lateral and dorsal cortex. (A) Schematic view of the putative organization of the dorso-lateral part of the telencephalon in an early mammaliaform (pre-mammlian synapsid). In these animals the lateral cortex (LC, violet) may have expanded over the dorsal cortex (DC, green), and at some point some radial glial progenitor (dark cells) may have started to produce both LC and DC cells. Note that since the internal anatomy of these animals is unknown, this scheme was based on modern macrosmatic reptiles. Radial glial cells of other brain regions are omitted for clarity. The proposed homology with neocortical layers is indicated with roman nueral. (B) Tangential expansion of the progenitors of the proto-neocortical column gave rise to the establishement of the neocortex. Note that the more lateral part of the neocortex maintains a direct olfactory input. Abbreviations: H.comm., hippocampal commissure; AC, Anterior commissure. Another interesting aspects of the hypothesis that the neocortex derived from the superposition of lateral and dorsal cortex, is that it may account for some hodological features that the UL shares with the olfactory cortex but not with the reptilian dorsal cortex. For instance, the olfactory cortex of tetrapods possesses homotopic projections to the contralateral hemisphere passing through the anterior commissure (Zeier and Karten, 1973; Butler, 1976; Hoogland and Vermeulen-Vanderzee, 1995; Sassoè-Pognetto et al., 1995; Suárez et al., 2014). Inter-hemispheric projections arising from UL neurons still decussate exclusively through this commissure in monotreme and marsupials, and this is generally thought to represent the basal condition in mammals (Ashwell et al., 1996; Suárez et al., 2014). Nonetheless, in sauropsids inter-hemispheric connections of DP and MP derivatives decussate at the nearby pallial/hippocampal commissure (Butler, 1976; Martinez-Garcia et al., 1990; Atoji et al., 2002). Thus, our hypothesis may account for the strange evolutionary history of the inter-hemispheric connections of the mammalian DP derivatives that at first flipped their direction and coursed a long lateral trip to decussate with fibers of the olfactory cortex at the anterior commissure. Only in eutherian mammals most, but not all, of the neocortical inter-hemispheric connections turned medially again decussating at the corpus callosum (Suárez et al., 2014). Another interesting similarity between the connections of olfactory cortex and UL is that they are both the source of feed-forward projections that flow to a series of hierarchical areas progressively defining sensory objects and ultimately converging on the LEC (Felleman and Van Essen, 1991; Haberly, 2001; Gilbert and Sigman, 2007; Wilson and Sullivan, 2011). In summary, the idea that the six layered neocortex originated from the superposition of lateral and dorsal cortex is consistent with the fossil record and may account not only for the topological position of the neocortex, but also for its basic cytoarchitecthural and hodological features. Unfortunately very little is known about the embryonic development of this putative six-layered primordium in modern reptiles. Guirado and Davila identified radial glial processes crossing both dorsal and lateral cortex in the lateral superposition of the lizard Podarcis Hispanica (Guirado and Dávila, 2002) and we made similar observations in Golgi stains of Lacerta Sicula (Luzzati unpublished observation). These authors raised the possibility that an independent progenitor domain giving rise to neurons of both dorsal and lateral cortex may actually exist in some living reptiles. In contrast to this interpretation however, Ulinsky reported that during development the layer II of the reptilian dorsal and lateral cortex is a continuous stratum of cells that is secondarily ruptured during differentiation (Ulinsky, 1990). Starting from this latter observation, a possible scenario for the evolution of the neocortex may be that in early mammaliaforms the homologs of UL and DL cells organized in a proto-neocortical column that was initially produced by spatially segregated progenitors. At some point a spatial to temporal patterning switch, together with the evolution of the inside-out neurogenic gradient, led to the generation of the proto-neocortical module from a single population of progenitors (Figure 5A). This crucial event enabled the tangential expansion of this module providing the basis for the establishment of the modern neocortex (Rakic, 1995; Lewitus et al., 2014; Figure 5B). According to the growth rings hypothesis of Sanides, during this tangential expansion the internal parts of the neocortical island progressively lost their allocortical features with the addition of stellate cells in layer IV and a reduction of cell density in layer II (Sanides, 1969; Sanides and Sanides, 1972). An intriguing aspect of this model is that it implies that the early neocortex worked as an higher order association cortex and that primary sensory areas appeared only subsequently. This latter idea has been also recently proposed based on functional models of both mammalian and reptilian allocortices (Fournier et al., 2015).\nSeveral crucial questions remain regarding the emergence of the inside out-gradient of neurogenesis, the appearance of layer IV cells and the arrival of the collo-thalamic projections to the dorsal pallial 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