TL;DR: The efferent connections of the hippocampal formation of the rat have been re‐examined autoradiographically following the injection of small quantities of 3H‐amino acids into different parts of Ammon's horn and the adjoining structures to indicate quite clearly that each component of the hippocampusal formation has a distinctive pattern ofefferent connections.
Abstract: The efferent connections of the hippocampal formation of the rat have been re-examined autoradiographically following the injection of small quantities of 3H-amino acids (usually 3H-proline) into different parts of Ammon's horn and the adjoining structures. The findings indicate quite clearly that each component of the hippocampal formation has a distinctive pattern of efferent connections and that each component of the fornix system arises from a specific subdivision of the hippocampus or the adjoining cortical fields. Thus, the precommissural fornix has been found to originate solely in fields CA1-3 of the hippocampus proper and from the subiculum; the projection to the anterior nuclear complex of the thalamus arises more posteriorly in the pre- and/or parasubiculum and the postsubicular area; the projection to the mammillary complex which comprises a major part of the descending columns of the fornix has its origin in the dorsal subiculum and the pre- and/or parasubiculum; and finally, the medial cortico-hypothalamic tract arises from the ventral subiculum. The lateral septal nuclei (and the adjoining parts of the posterior septal complex) constitute the only subcortical projection field of the pyramidal cells in fields CA1-3 of Ammon's horn. There is a rostral extension of the pre-commissural fornix to the bed nucleus of the stria terminalis, the nucleus accumbens, the medial and posterior parts of the anterior olfactory nucleus, the taenia tecta, and the infralimbic area, which appears to arise from the temporal part of field CA, or the adjacent part of the ventral subiculum. The projection of Ammon's horn upon the lateral septal complex shows a high degree of topographic organization (such that different parts of fields CA1 and CA3 project in an ordered manner to different zones within the lateral septal nucleus). The septal projection of “CA2” and field CA3 is bilateral, while that of field CA1 is strictly unilateral. In addition to its subcortical projections, the hippocampus has been found to give rise to a surprisingly extensive series of intracortical association connections. For example, all parts of fields CA1, CA2 and CA3 project to the subiculum, and at least some parts of these fields send fibers to the pre- and parasubiculum, and to the entorhinal, perirhinal, retrosplenial and cingulate areas. From the region of the preand parasubiculum there is a projection to the entorhinal cortex and the parasubiculum of both sides. That part of the postsubiculum (= dorsal part of the presubiculum) which we have examined has been found to project to the cingulate and retrosplenial areas ipsilaterally, and to the entorhinal cortex and parasubiculum bilaterally.
TL;DR: The existence of an entorhinal cell type that fires when an animal is close to the borders of the proximal environment is reported, and border cells may be instrumental in planning trajectories and anchoring grid fields and place fields to a geometric reference frame.
Abstract: We report the existence of an entorhinal cell type that fires when an animal is close to the borders of the proximal environment. The orientation-specific edge-apposing activity of these "border cells" is maintained when the environment is stretched and during testing in enclosures of different size and shape in different rooms. Border cells are relatively sparse, making up less than 10% of the local cell population, but can be found in all layers of the medial entorhinal cortex as well as the adjacent parasubiculum, often intermingled with head-direction cells and grid cells. Border cells may be instrumental in planning trajectories and anchoring grid fields and place fields to a geometric reference frame.
TL;DR: The heaviest projections from the amygdala to the hippocampal formation and the parahippocampal areas originate in the lateral, basal, accessory basal, and posterior cortical nuclei, and the underlying principles of organization of these projections are discussed.
Abstract: Recent anterograde and retrograde studies in the rat have provided detailed information on the origin and termination of the interconnections between the amygdaloid complex and the hippocampal formation and parahippocampal areas (including areas 35 and 36 of the perirhinal cortex and the postrhinal cortex). The most substantial inputs to the amygdala originate in the rostral half of the entorhinal cortex, the temporal end of the CA1 subfield and subiculum, and areas 35 and 36 of the perirhinal cortex. The amygdaloid nuclei receiving the heaviest inputs are the lateral, basal, accessory basal, and central nuclei as well as the amygdalohippocampal area. The heaviest projections from the amygdala to the hippocampal formation and the parahippocampal areas originate in the lateral, basal, accessory basal, and posterior cortical nuclei. These pathways terminate in the rostral half of the entorhinal cortex, the temporal end of the CA3 and CA1 subfields or the subiculum, the parasubiculum, areas 35 and 36 of the perirhinal cortex, and the postrhinal cortex. The connectional data are summarized and the underlying principles of organization of these projections are discussed.
TL;DR: Neurogenesis in the rat hippocampal region was examined with 3H‐thymidine autoradiography and the percentage of labelled cells and the proportion of cells added during each day of formation were determined at several anatomical levels.
Abstract: Neurogenesis in the rat hippocampal region was examined with 3H-thymidine autoradiography. The rats in the prenatal groups were the offspring of pregnant females given two injections of 3H-thymidine on consecutive days in an overlapping series: embryonic (E) day E13+E14, E14+E15,…, E21+E22. The rats in the postnatal (P) groups were injected in two nonoverlapping series: first, the day of birth (PO) and P1, P2+P3,…, P18+P19; second, P0–P3, P4–P7,…, P16–P19. On 60 days of age, the percentage of labelled cells and the proportion of cells added during each day of formation were determined at several anatomical levels within each structure of the hippocampal region (entorhinal cortex, parasubiculum, presubiculum, subiculum, Ammon's horn, and the dentate gyrus) and the hippocampal rudiment (tenia tecta, indusium griseum). The neurons in each structure arise in overlapping, but still significantly different, waves: the hippocampal rudiment between E16–E17; the entorhinal cortex between E15–E17; the para- and presubiculum between E16–E19; the subiculum between E16–E18; large cells in strata oriens, radiatum, lacunosum-moleculare of Ammon's horn between E15–E17; Ammon's horn pyramidal cells between E17–E19; large cells in the dentate hilus and molecular layer between E15–E19. Dentate granule cells begin to originate on E17, and 10% of the population forms after P18.
There are three characteristic gradients of formation within structures. First, deep cells are generated before superficial cells. Second, cells closer to the rhinal fissure are formed before those lying farther away (“rhinal to dentate” gradient). Third, later forming cells are flanked by earlier forming superficial and deep cells (“sandwich gradient”) in the entorhinal cortex (layer III cells originate after layers II and IV), Ammon's horn (pyramidal cells originate after large cells in strata oriens, radiatum, and lacunosum-moleculare), and the dentate gyrus (granule cells originate after large cells in the hilus and molecular layer). There is a “rhinal to dentate” gradient between structures. The entorhinal cortex starts first, next is the subiculum, then field CA3 of Ammon's horn, and finally, the dentate gyrus. Two structures are exceptions to this gradient. The para- and presubiculum form significantly later than the subiculum, and CA1 forms significantly later than adjacent CA3 cells; this late neurogenesis may be related to prominent thalamic input to both structures.
Neurogenetic gradients between the cells providing laminated afferent input to the Ammonic pyramidal and dentate granule cells correlate with their order of termination: afferents from progressively later-originating cells terminate progressively closer to the cell body. Topographic hippocampal projections along the dorsoventral axis correlate with formation patterns in target structures: dorsal hippocampal fibers project to zones occupied by earlier-forming cells in the lateral septal nucleus and pars posterior of the mammillary body; ventral hippocampal fibers project to zones occupied by later-forming cells in these structures.
TL;DR: The pathway from the entorhinal cortical region to the hippocampal formation has previously been shown to be comprised of two sub‐systems, one of which projects predominantly to the ipsilateral fascia dentata and regio inferior of the hippocampus proper, and a second which projects bilaterally to regio superior.
Abstract: The pathway from the entorhinal cortical region to the hippocampal formation has previously been shown to be comprised of two sub-systems, one of which projects predominantly to the ipsilateral fascia dentata and regio inferior of the hippocampus proper, and a second which projects bilaterally to regio superior. The goal of the present investigation was to determine if these two pathways might originate from different cell populations within the entorhinal area. The cells of origin of these entorhinal pathways were identified by retrograde labeling with horseradish peroxidase (HRP).
Injections which labeled the entorhinal terminal fields in both the fascia dentata and regio superior resulted in the retrograde labeling of two populations of cells in the entorhinal area. Ipsilateral to the injection, HRP reaction product was found in the cells of layer II (predominantly stellate cells) and the cells of layer III (predominantly pyramidal cells). Contralateral to the injections, however, the reaction product was found almost exclusively in the cells of layer III.
With selective injections of the entorhinal terminal field in regio superior, only the cells of layer III were labeled, but these were labeled bilaterally. Selective injection of the entorhinal terminal field in the fascia dentata, however, resulted in the labeling of cells of layer II, but not of layer III, and these cells of layer II were labeled almost exclusively ipsilaterally. A very small number of labeled cells in layer II were, however, found contralateral to the injection as well.
No labeled cells were found either in the presubiculum or parasubiculum following injections of the hippocampal formation. These cell populations were found capable of retrograde transport of HRP, however, since cells in both presubiculum and parasubiculum were labeled following HRP injections into the contralateral entorhinal area.
These results suggest that the projections to the fascia dentata originate from the cells of layer II, while the projections to regio superior originate from the cells of layer III of the entorhinal region proper. The very slight crossed projection from the entorhinal area to the contralateral area dentata probably originates from the small population of cells in layer II which are labeled following HRP injections in the contralateral area dentata.