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An immunohistochemical study of the telencephalon of the senegal bichir (Polypterus senegalus).

The telencephalon in ray-finned fish (actinopterygians) is everted, in contrast to the evaginated telencephalic hemispheres in all other vertebrates. In the more derived ray-finned fish, the teleosts, proliferation of neurons and their migration from the ependymal zone of the pallium renders comparisons between telencephalic cell groups of the teleosts and members of other vertebrate groups extremely difficult. The telencephalon of Polypterus (a primitive living ray-finned fish), although everted, is cytoarchitecturally much simpler than that of teleosts. We have thus applied immunohistochemical techniques to the study of the telencephalon of Polypterus to help clarify the evolution of the telencephalon in teleosts and facilitate comparisons between the telencephalon in ray-finned fish and other vertebrates. Antisera against the following neuroactive substances were used: 1) serotonin (5HT), 2) tyrosine hydroxylase (TH), 3) substance P (SP), 4) leucine-enkephalin (ENK), 5) neuropeptide Y (NPY), and 6) the neurotensin-related hexapeptide LANT6. Several features of the labeling patterns obtained suggested that the dorsal and ventral subdivisions of the area ventralis are homologous as a field to the basal ganglia and septum plus other basal telencephalic regions of land vertebrates, sharks and lungfish: 1) an abundance of SP+, NPY+, and ENK+ fibers; 2) an abundance of TH+ fibers, possibly of posterior tubercle/tegmental origin; 3) the presence of an SP+ fiber bundle that appeared to descend from basal telencephalic levels and terminate in the posterior tubercle/tegmentum, which contain TH+ (possibly dopaminergic) neurons; and 4) an abundance of 5HT+ fibers, presumably of posterior tubercle/tegmental origin. It was not possible, however, to recognize distinct pallidal and striatal subdivisions within the area ventralis of Polypterus. The olfactory pallium (P1) was generally poor in most of the substances examined, except for the presence of LANT6+ fibers. The P3 pallial field was conspicuously rich in SP+ and ENK+ fibers throughout its extent, and the caudal and lateral parts of the P2 field were rich in SP+ fibers and ENK+ fibers. Since this is characteristic of the medial pallial and/or dorsomedial pallial walls of the telencephalon in lungfish, sharks, frogs, and reptiles, the P3 field and caudolateral part of the P2 field may be homologous to these portions of the telencephalon in other vertebrates. More rostromedial parts of P2 may correspond to those parts of the pallium in land vertebrates that are in receipt of specific sensory input from the thalamus, since low neuropeptide levels are characteristic of these regions.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Isolation and characterization of a library of cDNA clones that are preferentially expressed in the embryonic telencephalon.

In order to isolate genes involved in development of the mammalian telencephalon we employed an efficient cDNA library procedure. By subtracting an adult mouse telencephalic cDNA library from an embryonic day 15 (E15) mouse telencephalic cDNA library we generated two subtracted libraries (ES1 and ES2). We estimate that ES1 contains between 200 and 600 different cDNA clones, which approximates the number of genes that are preferentially expressed in the E15 telencephalon, compared to the adult telencephalon. Northern analysis of 20 different cDNA clones shows that 14 of these are expressed at least 5-fold more in the E15 telencephalon than the adult telencephalon. Limited sequencing of the 14 differentially expressed clones reveals that 10 have no significant identity to sequences in GenBank and EMBL databases, whereas the other 4 have significant sequence identity to vimentin, histone 3.3, topoisomerase I and the B2 repeat element. In situ hybridization using one of the differentially expressed cDNAs, TES-1, demonstrates that it is transiently expressed in the anlage of the basal ganglia. In situ hybridization with another differentially expressed cDNA clone, TES-4, shows that it is specifically expressed in differentiating cells of the neural axis with a distinctive rostral-caudal temporal pattern. These findings, and the methods that we have developed, provide a framework for future investigations of the genetic control of telencephalon development.

Animals

[The activities of the transmitter metabolizing enzymes monoamineoxidase and acetylcholinesterase and of succinic dehydrogenase were investigated in the telencephalon of Salmo irideus (teleostei) (author's transl)].

The activities of the transmitter metabolizing enzymes monoamineoxidase and acetylcholinesterase and of succinic dehydrogenase were investigated in the telencephalon of Salmo irideus (teleostei). There is a high reaction intensity of the MAO in the whole telencephalon with graded differences between the various nuclear areas. It is highest in the area dorsalis telencephali; the larger fiber tracts (Tr. olfactorius medialis et lateralis; medial and lateral forebrain bundle) are strong MAO-positive, too. There is an extremely weak AchE-reaction throughout the telencephalon. The only nuclear area which is clearly histochemically demonstrable is the regio Dm2 in the area dorsalis telencephali. The SDH-activity is demonstrable in the whole telencephalon. The activity is higher in the area dorsalis as compared with the area ventralis telencephali. In the ependym there are only a few traces of MAO- and SDH-reaction products. The recent findings may support, from a histochemical view, the hypothesis that the telencephalon of Salmo irideus represents brain structures which show structural and functional similarities with limbic telencephalic structures of higher vertebrates.

Acetylcholinesterase

Characterisation of an allosteric modulatory protein associated with alpha-[3H]amino-3-hydroxy-5-methylisoxazolepropionate binding sites in chick telencephalon: effects of high-energy radiation and detergent solubilisation.

alpha-[3H]Amino-3-hydroxy-5-methylisoxazolepropionate ([3H]AMPA) binds to 1-day-old chick telencephalon membranes with KD and Bmax values of 138 nM and 2.56 pmol/mg of protein, respectively. High-energy radiation bombardment of intact frozen telencephalon resulted in a biphasic inactivation curve for [3H]AMPA binding. At a 5.8-Mrad radiation dose, the affinity of [3H]AMPA binding was increased (54 nM), but there was no apparent alteration in the Bmax value (2.76 pmol/mg of protein). We attribute this phenomenon to the inactivation of a high molecular weight modulatory protein that down-regulates the affinity of [3H]AMPA binding. The estimated molecular masses of the AMPA binding site and of the modulatory component were 59 and 108 kDa, respectively. Solubilisation with n-octyl-beta-glucopyranoside resulted in an increase in the Bmax (4.7 pmol/mg of protein) with no pronounced alteration in the affinity (109 nM) of [3H]AMPA binding. However, the solubilisation-induced increase in Bmax did not occur in telencephalon irradiated before solubilisation. In contrast, the increase in affinity induced by radiation treatment was still detected in solubilised extracts. These results suggest that the number and affinity of [3H]AMPA sites in chick telencephalon are closely regulated and that the modulatory systems involved are affected by both irradiation and solubilisation.

Allosteric Regulation

Histochemical distribution of zinc in the brain of the rainbow trout, Oncorhynchos myciss. I. The telencephalon.

The present paper which describes the distribution of zinc in the telencephalon of the rainbow trout, Oncorhynchos myciss, is the first report on the distribution of a heavy metal in the fish brain. Zinc was demonstrated histochemically by silver enhancement using the Neo-Timm method. The staining was mainly confined to the neuropil, but both moderately and intensely stained nerve cell bodies were of common occurrence. Stained fibers were never observed. The staining revealed a specific distribution pattern which could easily be correlated with the telencephalic nuclei defined on the basis of cytoarchitectural features. However, the telencephalon stained much more weakly than the rest of the brain, in striking contrast to the situation in the reptilian, mammalian, and avian brain. In these classes, high staining intensities are observed almost exclusively in the telencephalon. The staining was essentially restricted to the nuclei of the ventral telencephalic area. In the dorsal telencephalic area, only the medial and central zones and medial part of the posterior zone showed comparable staining intensities. The Neo-Timm staining pattern lends support to the view that the pallio-subpallial boundary is between the medial and dorsal zones of the dorsal telencephalic area. The distribution of zinc has been compared with the terminal field of afferent projections, known from experimental mapping, and also with the distribution of substance P and vasoactive intestinal peptide. Finally, the possible functional implications of zinc in synaptic vesicles are considered.

Animals

Immunohistochemical localization of glutamine synthetase in mesencephalon and telencephalon of the lizard Gallotia galloti during ontogeny.

The immunohistochemical localization of glutamine synthetase, an astrocyte marker in mammals, was determined in the telencephalon and mesencephalon of the lizard Gallotia galloti during development by using an antiserum raised against chicken brain glutamine synthetase. Ependymal glial cells and their radial processes were glutamine synthetase immunoreactive, and they were present also in the adult. Immunoreactivity was also detected in two populations of scattered cell bodies, each preferentially localized in different zones: star-shaped cells morphologically similar to mammalian astrocytes, and ovoid or pear-shaped cell bodies, the processes of which were aligned with radial fibers and formed perivascular end-feet. Both populations displayed ultrastructural characteristics of astrocytes even though a comparison with our previous results (Monzon-Mayor et al., 1989; Yanes et al., 1989) indicated that many of these cells did not react with antibodies directed against the astrocyte-specific glial fibrillary acidic protein. During ontogeny, glutamine synthetase immunoreactivity appeared in radial glial processes and in ependymal glial cells of midbrain at embryonic stage 35 (E35) and of telencephalon at E37; in both regions, immunoreactivity in the radial glia increased until hatching and then decreased until adulthood, but it did not disappear. Labelled scattered cells became progressively more numerous and more immunoreactive. A comparative analysis of the distribution of these cells at different ages tends to suggest that some of the "ovoid" astrocytes originate in, and migrate out from, the proliferative zone of the different sulci, whereas the star-shaped cells appear directly in situ, probably because they begin to express glutamine synthetase after they have reached their final location.

Aging

Myelin and myelinization in the telencephalon and mesencephalon of the lizard Gallotia galloti as revealed by the immunohistochemical localization of myelin basic protein.

We have studied in the telencephalon and mesencephalon of the lizard Gallotia galloti the localization and the chronology of appearance of the immunoreactivity due to the presence of a myelin-specific protein: the Myelin Basic Protein (MBP). MBP-like immunoreactivity was present with different degrees of intensity in many nerve fibers (isolated, in tracts and in commissurae) and it was apparently more abundant in mesencephalon. During ontogeny the earliest MBP-like immunoreactivity was detected at E.36 in few tracts in mesencephalon and appeared at E.40 in telencephalon, proceeding caudo-rostrally and from the ventral (basal) to the dorsal (alar) regions. Accumulation of MBP continued after hatching. Oligodendrocyte cell bodies were not immunopositive, not even at the youngest ages studied.

Animals

Reciprocal connections between the 'nucleus rotundus' and the dorsal lateral telencephalon in the weakly electric fish Gnathonemus petersii.

Injections of horseradish peroxidase into the dorsal lateral telencephalon of the weakly electric mormyrid fish Gnathonemus petersii resulted in retrogradely filled cells and anterogradely labelled terminals in the 'nucleus rotundus' of the ipsilateral rostral diencephalon. This connection courses via the lateral part of the lateral forebrain bundle. The present results suggest a particularly close relationship between the Dla cell group of the lateral telencephalon and the 'nucleus rotundus'.

Animals

Mouse telencephalon exhibits an age-related decrease in glutamate (AMPA) receptors but no change in nerve terminal markers.

The central excitatory amino acid receptor selective for alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid (AMPA) was examined in brain tissue from mice at 3 and 25 months after birth. Antibodies against the rat GluR-A glutamate receptor subunit (selective for kainate and AMPA) labeled a mouse brain component of about M(r) 100,000. Telencephalic tissue from the older group of mice exhibited 31% less immunoreactivity towards this component as compared with that from the young group. Binding of [3H]AMPA also decreased with age in the telencephalon to an extent which was similar to the loss of receptor immunoreactivity. Scatchard analysis revealed that this reduction is due to a decrease in receptor density and not to a change in binding affinity. In contrast, there were only small age-related changes in AMPA receptor immunoreactivity and binding levels in the brain stem and cerebellum. Binding to dopamine, serotonin, or GABA receptors was not significantly reduced in the older mice. Since the nerve terminal markers synaptophysin and the SV2 glycoprotein were not detectably different in the two groups of mice, the age-related reduction in AMPA receptors is not likely to be due to a general decrease in synaptic density. These data suggest that glutamatergic neurotransmission mediated by AMPA-type receptors is selectively impaired with aging in the telencephalon.

Aging

Telencephalon-restricted expression of BF-1, a new member of the HNF-3/fork head gene family, in the developing rat brain.

We have previously characterized a novel transcription factor family in mammals, the HNF-3 family, by the members' homology to one another and to the Drosophila homeotic gene fork head. The expression of fork head is restricted to the anterior and posterior termini of the early fly embryo. Brain factor 1 (BF-1) is a new member of this family isolated from rat brain with an expression pattern and DNA binding specificity distinct from the HNF-3 genes. Expression is highly restricted in the developing neural tube to its rostral end, which gives rise to the telencephalon. These results suggest that BF-1 plays an important role in the establishment of the regional subdivision of the developing brain and in the development of the telencephalon.

Amino Acid Sequence

Distribution of choline acetyltransferase immunoreactivity in the telencephalon of the lizard Gekko gecko.

The presumptive cholinergic elements in the telencephalon of the lizard Gekko gecko were demonstrated with the AB8 anti-choline acetyltransferase (ChAT) antibody. Somata positive for ChAT were observed in the striatum, nucleus accumbens, the dorsal ventricular ridge, nucleus olfactorius anterior, tuberculum olfactorium, diagonal band of Broca, septum, bed nucleus of the medial forebrain bundle and lateral preoptic area. Staining of the neuropil was most conspicuous in the striatum but also occurred in the medial cortex and nucleus septi impar. The results indicate that the distribution of ChAT-positive somata in the telencephalon in reptiles is comparable to that in mammals; however, the relation of cholinergic somata and ChAT-positive neuropil seen in the striatum of G. gecko is different from that in mammals.

Animals

Transient calbindin-D28k-positive systems in the telencephalon: ganglionic eminence, developing striatum and cerebral cortex.

Calbindin-D28k (calbindin) is a member of the superfamily of calcium-binding proteins implicated in the regulation of intracellular calcium. In the mature brain, calbindin is widely expressed in neurons of the forebrain and the hindbrain, and in the telencephalon calbindin-like immunoreactivity is particularly strongly expressed by medium-sized neurons of the striatum and by certain other neurons in the cortex and subcortex. We have traced the development of calbindin expression in the forebrain of the rat, and report here that in addition to the steady development of these calbindin-positive neuronal systems, transient waves of calbindin expression occur in cells of the ventricular zones of the basal ganglia and cortex and in cells of the telencephalic regions derived from these ventricular zones including radial glia of the developing striatum. In the striatum and its ventricular zone (the ganglionic eminence, or GE) we identified four transient calbindin-positive systems in the perinatal period. First, calbindin-immunoreactive cells began to appear in the GE by embryonic day (E)18, and by E20 an extensive dorsal and lateral part of the GE was marked by dense calbindin-like immunoreactivity in the ventricular zone. This calbindin system peaked at postnatal day (P)0-P3 and disappeared by P15. Its presence suggests that the GE is divisible on a molecular basis into lateral and medial districts that may correspond to derivatives of the lateral and medial ventricular ridges. Second, a system of calbindin-positive processes appeared in the dorsal and lateral caudoputamen with temporal and spatial distributions matching the germinal zone system. Many of these processes could be traced from calbindin-positive cells in the ventricular zone of the GE, including processes stretching across the full width of the dorsal caudoputamen. Double-staining experiments demonstrated that these radial processes were Rat.401-positive, suggesting that they form a subset of radial glia in the developing telencephalon. These findings demonstrate that during development calbindin is expressed in glial as well as neural cells. They further suggest that the radial glia associated with the GE form heterogeneous populations, the transient calbindin-positive radial glia being associated with the lateral ridge of the GE and its derivatives. Third, a scattered population of calbindin-positive cells with morphologies different from the common medium-sized calbindin-immunoreactive neurons of the striatum appeared in the dorsal and lateral striatum from about E20 to P15. Some of these cells were close to the transient calbindin-positive radial processes in the same region, but others were not.(ABSTRACT TRUNCATED AT 400 WORDS)

Aging

Neuronal migration and contact guidance in the primate telencephalon.

Over the last decade, evidence from experimental studies on neuronal migration in non-human primates has accumulated to the point where it can significantly amplify our understanding of the normal and pathological development of the human telencephalon. Systematic analysis of neuron genesis by the method of H3-thymidine autoradiography shows that in rhesus monkeys all neurons destined for the neocortex are generated near the surface of the lateral ventricle during a two-month period in the middle of gestation. Following their last cell division, young neurons migrate outwards across the cerebral wall to the developing cortical mantle, a journey that requires one to three days at early stages of neurogenesis, or more than two weeks towards the end of cortical development. From the very beginning, the basic columnar organization of the neuroepithelium favours radial migration. During later stages, when the primate telencephalic wall expands unevenly in thickness and surface area and begins to form primary fissures and cerebral promontoria, young neurons migrate to their cortical destinations in apposition to fascicles of radial glial fibres which span the full distance between the ventricular and pial surface. Furthermore, it appears that several generations of neurons all originate in the same restricted location at the ventricular surface, migrate along the same glial fascicles and consequently accumulate in the same radial cortical 'columns' in which, as a rule, somas of later generated neurons take positions external to the somas of their predecessors. It is proposed that fascicles of radial fibres (a) facilitate neuronal migration to the distant cortical plate through a complex assembly of closely-packed cells and processes that compose the developing primate telencephalon; (b) provide constraints which preserve a radial alignment of clonally related neurons in cortical columns; and (c) reproduce the mosaicism of the germinal ventricular zone at the expanded and curved cerebral surface.

Animals

Formaldehyde-induced fluorescence in the telencephalon and diencephalon of the eel (Anguilla anguilla l.). A fluorescence-microscopic and microspectrofluorometric investigation with special reference to the innervation of the pituitary.

In the telencephalon and diencephalon of the eel (Anguilla anguilla L.) formaldehyde-induced fluorescence was studied microscopically and microfluorometrically with special emphasis on the innervation of the pituitary. In the telencephalon fluorescent fibers contained predominantly noradrenaline fluorophores. Fluorescent nuclei could not be established. In the diencephalon fluorescent perikarya were found in: (1) the paraventricular organ (PVO), possessing either dopamine or, to a lesser extent, serotonin fluorophores; (2) the PVO-accompanying group, exhibiting spectral data resembling those of noradrenaline fluorophores; (3) the nucleus hypothalami anterior (NHA), a small paired group of catecholamine-containing cells posterior to the commissura transversa.--The nucleus lobi inferioris exhibited a high density of delicate, most probably dopamine-containing terminals, while fibers surrounding this nucleus contained noradrenaline fluorophores. A high density of fluorescent terminals containing dopamine and/or noradrenaline was found in the habenular complex. Fluorescent terminals in the pituitary contained fluorophores resembling either dopamine or noradrenaline. Fluorescent tracts entered the pituitary from different directions. A rostral, unpaired tract enters the neurointermediate lobe, as also verified experimentally. The rostral pars distalis receives two paired tracts, one from a rostral and one from a dorsal direction. The proximal pars distalis also receives two paired tracts, one from a dorsal and one from a posterior direction.

Anguilla

Ascending afferents to the telencephalon of ranid frogs: an anterograde degeneration study.

Sources of telencephalic afferents were examined in two species of frogs, Rana catesbeiana and Rana pipiens, by study of anterograde degeneration resulting from hemisection of the brain at the isthmus or at the caudal border of the thalamus and from electrolytic lesions in various portions of the diencephalon. The results indicate that some telencephalic afferents arise from levels caudal to the isthmus, that some striatal afferents arise from a level between the isthmus and the caudal border of the thalamus, and that a projection to the ipsilateral striatum arises from the anterior and/or middle portions of the dorsal thalamus. Diencephalic projections to pallial portions of the hemisphere were also observed. These results demonstrate substantial non-olfactory afferent projections to the telencephalon in ranid frogs.

Animals

The distribution of neurotransmitters and neurotransmitter-related enzymes in the dorsomedial telencephalon of the pigeon (Columba livia).

Immunoreactivity to four neurotransmitters/transmitter-related enzymes was found in the dorsomedial telencephalon (hippocampal region) of the pigeon. Putative afferent fibers containing choline acetyltransferase-like, serotonin-like, and tyrosine hydroxylase-like immunoreactivity were seen in a fiber tract passing through the septo-hippocampal junction and along the medial wall of the hippocampal region. The most intensive labeling of neuropil and terminals of all four substances was found in the dorsomedial area of the hippocampal region. Glutamic acid decarboxylase-like immunoreactivity was seen in sparsely scattered cells throughout the region. These results are discussed in relation to hypotheses about the boundaries and subdivisions of the hippocampal region of the pigeon.

Afferent Pathways

The distribution of neuropeptides in the dorsomedial telencephalon of the pigeon (Columba livia): a basis for regional subdivisions.

The distribution of six neuropeptides [substance P (SP), leucine (leu5-) enkephalin (LENK), vasoactive intestinal polypeptide (VIP), cholecystokinin (CCK), neuropeptide Y (NPY), and somatostatin (SS)] in the dorsomedial telencephalon (hippocampal region) of the pigeon was studied by immunohistochemistry. All six peptides were found in fibers passing through the septo-hippocampal junction and along the medial wall of the hippocampal region. NPY-, SS-, and VIP-like staining of fibers was seen in the hippocampal commissure. NPY and SS had similar distributions within the hippocampal region, both being most conspicuous in cell bodies, terminals, and fibers of the medial hippocampal region. VIP-positive cells were found in an area dorsal to the SS/NPY cell region. CCK-like immunoreactivity was found in terminal baskets surrounding large cells of a v-shaped structure in the ventromedial hippocampal region. SP- and LENK-like immunoreactivity was found in neuropils in a lateral-dorsal region, the two substances showing similar distributions. This region is thought to lie lateral to the limit of the hippocampal region. Parallels with the distribution of immunoreactivity in the mammalian hippocampus are used to suggest possible equivalent subdivisions of the avian and mammalian hippocampal regions.

Animals

An aberrant nucleus in the telencephalon following administration of ENU during neuroembryogenesis.

Administration of ENU to rats during embryonic development caused the emergence of aberrant nuclei in the dorsal telencephalon. They were located in the corpus callosum, and were composed of pyramidal and stellate neurons. This suggested that the aberrant nuclei arose out of the neurons of cerebral cortex that had failed to migrate during embryogenesis. The aberrant nuclei were found predominantly in the animals receiving ENU on day 18 or earlier of gestation.

Animals