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Somatosensory areas in the telencephalon of the pigeon. II. Spinal pathways and afferent connections.

There are two somatosensory areas in the telencephalon of the pigeon which receive an input from the spinal somatosensory system: one in the rostral Wulst which consists of the three hyperstriatal layers (h. accessorium (HA), h. intercalatus superior (HIS) and h. dorsale (HD] and one in the caudal telencephalon (neostriatum caudale (NC), neostriatum intermedium (NI) and hyperstriatum ventrale (HV]. Recordings of evoked single unit or multi unit activity and of field potentials before and after lesions of spinal pathways at a high cervical level (C4) were made to determine the contribution of these pathways to the transmission of somatosensory signals to these telencephalic areas. The rostral Wulst area receives somatic signals only through dorsal tracts contralateral to the recording site. Inputs from the wing arise mainly through the dorsal columns (DC) and those from the leg largely through the dorsolateral funiculus (DLF). The spinal projection pathway to the caudal neostriatal area includes the dorsal tracts and parts of the lateral funiculi on both sides. There was no difference in response form between the wing and leg responses. Signals transmitted through the lateral pathways were found to elicit the earliest responses (6-13 ms, electrical stimulation) in the caudal forebrain, while signals travelling through the DC arrive later in the caudal area (about 14 ms for wing stimulation) than in the rostral Wulst area (about 9 ms). The afferent thalamic and intratelencephalic connections of the two somatosensory areas in the telencephalon of the pigeon were investigated with retrograde transport of the neuronal tracers horseradish-peroxidase (HRP) or wheatgerm agglutinated HRP (WGA-HRP), Fast Blue (FB) and Rhodamine-isothiocyanat (RITC). Small tracer-injections were made under electrophysiological control at somatosensory responsive locations. These investigations confirm the projection of the caudal part of the nucleus dorsolateralis posterior (DLPc) to the caudal area and of the nucleus dorsalis intermedius ventralis anterior (DIVA) to the rostral area. In addition, it could be shown that the NI/NC projects to the HV thus confirming the electrophysiological results reported in a companion paper (Funke 1989) that the HV is a secondary area. The integrative function of HV is supported by connections to other sensory and motor telencephalic areas. Combined injections of FB and RITC revealed a topographic projection from the DIVA to the anterior Wulst.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Somatostatin and neuropeptide Y are almost exclusively found in the same neurons in the telencephalon of turtles.

In mammals, somatostatin and neuropeptide Y (NPY) are largely found in the same neurons of the telencephalon. To determine if this is a phylogenetically ancient feature of telencephalic organization, the brain of red-eared turtles was examined using immunofluorescence double-labeling procedures. The results showed that somatostatin and NPY are found almost exclusively in the same neurons in the telencephalon of turtles, but these neuropeptides rarely co-occur in neurons outside the telencephalon. Thus, the extensive co-occurrence of NPY and somatostatin appears to be a feature of telencephalic organization that was present in the reptilian common ancestors of mammals and modern reptiles.

Animals↗

Cholinergic neurons of fetal rat telencephalon in aggregating cell culture respond to NGF as well as to protein kinase C-activating tumor promoters.

Serum-free aggregating cell cultures of fetal rat telencephalon treated with the potent tumor promoter phorbol 12-myristate 13-acetate (PMA) showed a dose-dependent, persistent stimulation of the enzymes choline acetyltransferase (ChAT), glutamic acid decarboxylase and glutamine synthetase. After elimination of the proliferating cells by treatment of the cultures with Ara-C (0.4 microM) only the cholinergic marker enzyme, ChAT, could be stimulated by tumor promoters. The non-promoting phorbol ester, 4 alpha-phorbol 12,13-didecanoate proved to be inactive in these cultures, whereas the potent non-phorbol tumor promoter, mezerein, produced an even greater stimulatory effect than PMA. Since PMA and mezerein are potent and specific activators of protein kinase C, the present results suggest a role for this second messenger in the development of cholinergic telencephalon neurons. Stimulation of ChAT required prolonged exposure (48 h) of the cultures to PMA and the responsiveness of the cholinergic neurons to the tumor promoters decreased with progressive cellular maturation. The cholinergic telencephalon neurons showed the same pattern of responsiveness for tumor promoters as for nerve growth factor (NGF). However, the combined treatment with NGF and either PMA or mezerein produced an additive stimulatory effect, suggesting somewhat different mechanisms of action.

Animals↗

Unilateral ablation of telencephalon induces appearance of contralateral cortical and subcortical projections to thalamic nuclei.

This study sought to investigate the afferent connections of the thalamus in the rat following massive telencephalic lesions. After unilateral removal of the telencephalon, horseradish peroxidase (HRP) was injected into the thalamic nuclei ipsilateral or, for control purposes, contralateral to the lesioned side. Injection of HRP into the ventral or posterior thalamus ipsilateral to the lesioned telencephalon led to retrograde transport and HRP-labeling of cells in unilateral projection areas in the cortex, diencephalon and mesencephalon and, in addition, unexpectedly, in their mirror-image sites in the hemisphere contralateral to the injected thalamus. HRP-positive cells in the contralateral hemisphere were found whenever HRP was injected 7 days postlesion, when the animals had ceased to exhibit spontaneous turning behavior, but not when it was injected immediately after the lesion. We favor the hypothesis that the development of contralateral afferents to the thalamic nuclei represents a morphological substrate of the behavioral reorganization necessitated by the asymmetrical removal of the telencephalon.

Afferent Pathways↗

Visual-evoked responses in the salmon telencephalon change during smolt transformation.

During smolt transformation, salmon behaviour changes dramatically and the fish are imprinted to their natal stream. This brief episode is accompanied by changes in neurochemistry and connectivity of the brain. Comparing visual-evoked potentials, recorded from optic tectum and telencephalon by glass microcapillaries, we found pronounced differences before and after smolt transformation in the telencephalon. In the forebrain of presmolts, only small short-latency responses are present whereas postsmolts display large long-latency waves which characteristic dynamic properties in addition. These findings suggest changes in the functional role of the telencephalon during smolt transformation.

Animals↗

Neurons of the olfactory organ projecting to the caudal telencephalon and hypothalamus: a carbocyanine-dye labelling study in the brown trout (Teleostei).

The caudal extrabulbar projections and their neurons of origin in the trout were studied after carbocyanine-dye (DiI) labelling in either the olfactory organ or the caudal telencephalon. DiI application to the caudal telencephalon labelled bipolar neurons in the olfactory epithelium, where they were sparsely distributed throughout the olfactory lamellae. Labelled fibres ran scattered in the olfactory nerve without forming bundles. DiI application to the olfactory organ labelled extrabulbar projections to the ventral telencephalon, preoptic region and tuberal hypothalamus. These results confirm that primary sensory fibres running in the medial olfactory tract of trout have an olfactory origin.

Animals↗

Development of neuronal markers in aggregating fetal rat telencephalon cells cultured in the presence of triiodothyronine.

The concentrations of the general neuronal markers D2-protein (N-CAM), D3-protein and neuron specific enolase (NSE) in reaggregating cultures of fetal rat telencephalon cells were affected by the presence of 30 nM triiodothyronine in the defined culture medium. The extent of normal developmental changes were enhanced by triiodothyronine, as demonstrated by crossed immunoelectrophoresis. From 13 to 19 days in culture, the concentration of D2-protein decreased, and the concentrations of both D3-protein and NSE increased. Nerve growth factor (NGF) was without effect on the development of these general neuronal markers. However, as shown previously both triiodothyronine and NGF increased the activity of choline acetyltransferase, a marker for cholinergic neurons. The results suggest an enhanced overall differentiation of several types of telencephalon neurons in the presence of triiodothyronine, and a specific stimulation of cholinergic telencephalon neurons by NGF.

Animals↗

Morphological and GABA-immunoreactive development of the embryonic chick telencephalon.

The development of neurons utilizing gamma-aminobutyric acid (GABAergic neurons) in prosencephalon and telencephalon from chicken embryonic days 4-14 (E4-E14) was studied by means of immunohistochemistry. Furthermore, routine histology and transmission electron microscopy. respectively, were performed in order to study the morphological development in the designated area. The main finding is that development of GABAergic neurons in the chick telencephalon is rapid; the GABA neurons are appearing in bulk at day 8, being "overexpressed" at days 10-11, decreasing in numbers thereafter and achieving mature morphology on day 14, which is considerably faster than in the rodent. Morphological analysis revealed that the prosencephalon mainly consisted of a thin layer of undifferentiated neuroblasts in the E4 embryo. By E6, the prosencephalon had increased in thickness and occasional cells outside the neuroepithelium showed a more mature morphology with a few cells weakly staining positive for GABA. At E8, the prospective granular and subventricular layers had developed. At E14, the appearance of the telencephalon is approximating that of the adult since both ependymal cells and morphologically mature neurons can be seen.

Animals↗

Evolution of neurotransmitter-related markers in the vertebrate telencephalon. Comparative microchemical study in discrete brain regions of a frog and a turtle.

1. Neurochemical markers related to cholinergic, GABAergic and glutamatergic/aspartatergic neurotransmission have been measured in telencephalic areas obtained by microdissection from frog (Rana esculenta) and turtle (Pseudemys scripta elegans) brain. 2. In both species, pallial areas showed remarkably higher levels of synaptosomal D-3H-aspartate high affinity uptake than basal regions. Conversely, striatal and septal areas possessed higher levels of the GABAergic marker glutamate decarboxylase (GAD) than the pallium. 3. A differential distribution of GAD was noticed in striatal regions, highest levels of the enzyme being present in the ventral striatum, followed by the nucleus accumbens and the dorsal striatum. 4. Cholinergic markers choline acetyltransferase (ChAT) and acetylcholinesterase (AChE) were rather uniformly distributed in the frog telencephalon, while, in the turtle, cholinergic markers were several-fold higher in the basal telencephalon, particularly in the striatum, than in the pallium. 5. The turtle dorsal ventricular ridge possessed ChAT levels more similar to the striatal than to the cortical ones. On the contrary, D-3H-aspartate uptake in the dorsal ventricular ridge was close to the highest levels found in cortical areas. 6. The quantitative neurochemical approach adopted for the present study appears to be a useful tool to investigate the problem of homologies and to gain new information on the evolution of neuron populations and neuronal connections in the vertebrate telencephalon.

Acetylcholinesterase↗

Expression patterns of developmental regulatory genes show comparable divisions in the telencephalon of Xenopus and mouse: insights into the evolution of the forebrain.

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.

Animals↗

Semaphorin5A expression in the developing chick telencephalon.

In the present study, we analyzed the expression of Semaphorin5A (Sema5A), a gene implicated in axon guidance and many other processes of neuronal development, in the developing chick telencephalon. By using a heterologous mouse probe and in situ hybridization techniques, we showed distinct patterns of Sema5A expression within the chick telencephalon. In early development, Sema5A was present in pallial regions, mainly in the neuroepithelium and in the deep mantle of ventral and lateral pallia, and in the subpallium. As development proceeds, some ventral pallial derivatives maintained a moderate to strong Sema5A expression, whereas other lateral or dorsal pallial derivatives showed low to moderate expression of Sema5A. The overall expression of Sema5A during development in the chick telencephalon was similar to that reported in mouse. Moreover, the expression of Sema5A in mesencephalic, diencephalic, and telencephalic centers related to the tectofugal system suggests an important role of this gene in the development.

Age Factors↗

Involvement of the telencephalon in spaced-trial avoidance learning in the goldfish (Carassius auratus).

Goldfish (Carassius auratus) received escape-avoidance training in a shuttle-response situation at a rate of a single trial per day. Widely spaced training evaluates the ability of a discriminative stimulus to control an avoidance response in the absence of stimulus carry-over effects from prior recent trials. In Experiment 1, master goldfish exhibited significantly faster avoidance learning than yoked controls. The results suggest that the shuttle response was instrumentally acquired. Experiment 2 demonstrated a significant deficit in the acquisition of avoidance behavior following ablation of the telencephalon. The implications of spaced-trial, telencephalon-dependent avoidance learning, as demonstrated in these experiments for the first time, are discussed in the context of comparative research on instrumental learning in goldfish. These results provide further support for the hypothesis that the fish telencephalon contains an emotional system that is critical for fear conditioning.

Animals↗

The dopaminergic innervation of the avian telencephalon.

The present review provides an overview of the distribution of dopaminergic fibers and dopaminoceptive elements within the avian telencephalon, the possible interactions of dopamine (DA) with other biochemically identified systems as revealed by immunocytochemistry, and the involvement of DA in behavioral processes in birds. Primary sensory structures are largely devoid of dopaminergic fibers, DA receptors and the D1-related phosphoprotein DARPP-32, while all these dopaminergic markers gradually increase in density from the secondary sensory to the multimodal association and the limbic and motor output areas. Structures of the avian basal ganglia are most densely innervated but, in contrast to mammals, show a higher D2 than D1 receptor density. In most of the remaining telencephalon D1 receptors clearly outnumber D2 receptors. Dopaminergic fibers in the avian telencephalon often show a peculiar arrangement where fibers coil around the somata and proximal dendrites of neurons like baskets, probably providing them with a massive dopaminergic input. Basket-like innervation of DARPP-32-positive neurons seems to be most prominent in the multimodal association areas. Taken together, these anatomical findings indicate a specific role of DA in higher order learning and sensory-motor processes, while primary sensory processes are less affected. This conclusion is supported by behavioral findings which show that in birds, as in mammals, DA is specifically involved in sensory-motor integration, attention and arousal, learning and working memory. Thus, despite considerable differences in the anatomical organization of the avian and mammalian forebrain, the organization of the dopaminergic system and its behavioral functions are very similar in birds and mammals.

Afferent Pathways↗

Expression of Ngn1, Ngn2, Cash1, Gsh2 and Sfrp1 in the developing chick telencephalon.

Patterning of the chick telencephalon has been debated, especially in regard to a ventral (subpallial) or dorsal (pallial) nature of the dorsal ventricular ridge (DVR). Here we report the expression patterns of chick homologues of molecules known to be involved in telencephalic patterning in other vertebrate species. We show here that the transcription factors Ngn1, Ngn2, Cash1, Gsh2 and the secreted frizzled related protein 1 (sfrp1), a wnt receptor, are expressed in characteristic telencephalic domains during chick development. At embryonic day 7 (E7) Ngn1 and Ngn2 are localized in the dorsal pallium and the DVR, similar to the region of Pax6 expression. In contrast, Gsh2 and Cash1 are restricted to the subpallium and some DVR cells. Interestingly Sfrp, a dorsoventral boundary marker in mouse telencephalon, is expressed between the DVR and subpallium. Gsh2 and Pax6 double-positive cells further characterize this boundary region in the developing chick telencephalon.

Animals↗

Dorsal-ventral patterning in the mammalian telencephalon.

The telencephalon is the most diverse region of the brain with respect to both morphology and neuronal subtypes. This fact makes the task of unraveling the mechanisms underlying the development of this brain region rather daunting. Recent attempts to subdivide the embryonic telencephalon into distinct progenitor domains along the dorsal-ventral axis have provided an important framework on which to begin this process. These progenitor domains are defined by the restricted expression of transcriptional regulators and are proposed to give rise to specific subtypes of neurons. Work over recent years has provided important insights into the establishment and maintenance of these progenitor domains in the developing telencephalon.

Animals↗

A subset of clones in the chick telencephalon arranged in rostrocaudal arrays.

BACKGROUND: Different areas of the vertebrate central nervous system appear to follow different rules during development for determining the position of sibling cells. For example, in the chick hindbrain, clones are frequently confined to a single functional unit that derives from a single rhombomere. In contrast, clones in the mammalian cerebral cortex often cross functional boundaries because of the extensive migration of sibling cells in orthogonal directions. We have investigated whether the pattern of clonal distribution in the chick telencephalon is similar to that of the hindbrain or to the more functionally analogous mammalian cerebral cortex. Progenitor cells in the chick telencephalon were marked using a retroviral library encoding alkaline phosphatase and over 10(5) distinct molecular tags. Patterns of dispersion were detected using alkaline phosphatase histochemistry, followed by the recovery and sequencing of the molecular tag. We also analyzed the phenotypes of cells that occurred within the clones. RESULTS: A subset of progenitors gave rise to clones that were found in rostrocaudal arrays resembling tubes. Arrays were restricted in the mediolateral and dorsoventral planes but could span up to 4 mm in the rostrocaudal direction. They were found throughout the telencephalon and a single clone often spanned more than one telencephalic nucleus. Rostrocaudal clones comprised 60% of clones containing five or more cells and contained many different types of neurons, astrocytes, oligodendrocytes, or various combinations of these cell types. CONCLUSIONS: Telencephalic progenitors are multipotent, producing progeny that become distinct cell types. Clonally related cells can migrate rostrocaudally within domains that are restrained in the mediolateral and dorsoventral directions. A subset of rostrocaudal clones resemble those seen in the mammalian cerebral cortex, with respect to the crossing of functional boundaries, but all rostrocaudal clones differ from the cerebral cortical clones in the pattern of spread of sibling cells, with the rostrocaudal clones being more constrained in the mediolateral and dorsoventral directions. A role for lineage in the patterning of the chick forebrain is supported by these observations. In addition, these data suggest a role for cues within the telencephalic marginal zone that serve to guide clones in their rostrocaudal migration.

Animals↗

Seasonal variation in androgen-metabolizing enzymes in the diencephalon and telencephalon of the male European starling (Sturnus vulgaris).

In seasonally breeding songbirds, seasonal fluctuations occur in serum testosterone (T) concentrations and reproductive behaviours. Many T-dependent behaviours are regulated by the activity of androgenic and oestrogenic metabolites within specific brain regions. Male European starlings breed in spring when circulating T concentrations peak. T and its metabolites act within portions of the diencephalon to regulate the pituitary-gonadal axis and to activate courtship and copulation. Song in male starlings is critical for mate attraction during the breeding season and is regulated by steroid-sensitive nuclei in the telencephalon and diencephalon. Outside the breeding season, T is undetectable, however, males continue to sing at high levels. This suggests that singing outside of the breeding season might not be T-dependent as it appears to be in the spring. Alternatively, singing when T is low might continue to be regulated by T due to increased sensitivity of the brain to the action of the steroid. This increased sensitivity could be mediated by changes in intracellular T metabolism leading to increased production of active or decreased production of inactive metabolites. To explore the relationship between T-metabolism and reproductive behaviour, we analysed seasonal changes in the activity of four brain T-metabolizing enzymes: aromatase, 17beta-hydroxysteroid dehydrogenase (17beta-HSDH), 5alpha-reductase (all three convert T into active metabolites) and 5beta-reductase (converts T into an inactive metabolite) in the diencephalon and telencephalon. In the anterior and posterior diencephalon, the highest aromatase was observed in spring when this region is critical for courtship and copulation. In the telencephalon, aromatase was highest and 5beta-reductase was lowest throughout the winter months well prior to the reproductive season and these enzymes presumably maximize T-activity within this region. Although these data do not indicate whether the metabolic changes occur specifically within song nuclei, these findings are compatible with the idea that singing in male starlings outside the breeding season may be regulated by steroids despite the presence of low serum T concentrations. Overall, seasonal changes in T-metabolizing enzymes appear to play a significant role in seasonal changes in behaviour and reproductive physiology.

17-Hydroxysteroid Dehydrogenases↗

Telencephalon-specific antigen identified by monoclonal antibody.

Immunohistochemical screening of monoclonal antibodies (mAbs) raised against fractions rich in the dendrodendritic synaptosomes of the rabbit olfactory bulb revealed that one of the mAbs (mAb 271A6) recognized a telencephalon-specific antigen or antigens. Thus, the stain with mAb 271A6 was observed throughout the gray matter of all regions of the neocortex, piriform cortex, hippocampus, striatum, septum, and the amygdaloid nucleus, in addition to the main and accessory olfactory bulbs. The mAb 271A6, however, labeled neither nontelencephalic regions of the central nervous system nor the peripheral nervous system so far examined. Dot-immunobinding assays of homogenates of various brain regions also showed the telencephalon-specific distribution of the antigen designated as 271A6. Antigen 271A6 is developmentally regulated. At birth, the antigen was expressed in a small quantity only in phylogenetically older telencephalic regions such as the olfactory bulb, piriform cortex, striatum, cingulate cortex, and hippocampus. It was hardly detectable in most areas of the neocortex. The densities and areas of 271A6-positive structures increased during the early postnatal period. These results demonstrate a molecular specificity of the most rostral brain segment, the telencephalon. mAb 271A6 may be a good tool for obtaining a better understanding of the molecular basis of the segmental organization or the segment-specific functions of the brain.

Aging↗