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PubMed1986

Factors modulating the effects of teleost telencephalon ablation on retention, relearning, and extinction of instrumental avoidance behavior.

Two experiments on the nature of avoidance behavior in the goldfish are reported. In the first experiment, a master/yoked design was used to determine the degree to which Pavlovian conditioning would generate acquisition of a shuttle response by goldfish. Results indicated that the mere sequence of signal-shock pairings experienced by the avoidance animals is not in this situation sufficient to explain the development of the instrumental avoidance responding. The second experiment was conducted to study the effects of three variables on the retention and extinction of the avoidance response. First, unilateral lesions of the telencephalon produced a transient deficit from which the subjects recovered to normal levels. Retention of the avoidance response after bilateral telencephalon ablation was impaired and never fully recovered with additional training. Second, this deficit was reduced under higher intensities of electric shock. Third, overtraining on the task before the ablation had only a nonsignificant enhancing effect on retention levels. Extinction was always poorer following ablation, independent of the type of ablation, intensity of shock, degree of overtraining, and terminal retention level of performance. The results are discussed in terms of various theories of telencephalic function in fish.

Animals↗
PubMed1985

Serial ablations of the telencephalon and avoidance learning by goldfish (Carassius auratus).

Simultaneous bilateral ablation of the teleost telencephalon was confirmed to dramatically impair the retention, extinction, and "reversal learning" of a preoperatively learned instrumental avoidance response in a shuttle box. The focus of the experiment was on whether two-stage serial unilateral ablations of the teleost telencephalon would ameliorate the effects seen following one-stage bilateral ablation and whether such amelioration would be a function of the fish's experiences between the staged ablations. The degree of retention, relearning, and extinction of the avoidance response was not modulated by the serial ablation procedure whether or not there was interoperation retraining experience. However, reversal learning, in which the previous warning signal became the safety signal and the previous safety signal became the new warning signal, was less impaired following serial ablations than following single-stage bilateral ablation. These results are contrasted with those from experiments with mammalian subjects.

Animals↗
PubMedDate not supplied

Transient cells of the developing mammalian telencephalon are peptide-immunoreactive neurons.

In the development of the mammalian telencephalon, the genesis of neurons destined for the various layers of the cerebral cortex is preceded by the generation of a population of cells that comes to reside in the subplate and marginal zones (see ref. 2 for nomenclature). In the cat, these cells are present in large numbers during development, when their location is correlated with the arrival and accumulation of ingrowing axonal systems and with synapses. However, as the brain matures, the cells disappear and the white matter and layer 1 of the adult emerge. Their disappearance occurs in concert with the invasion of the cortical plate by the axonal systems and with the elimination of the synapses from the subplate. Here we report that the subplate cells have properties typical of mature neurons. They have the ultrastructural appearance of neurons and receive synaptic contacts. They also have long projections and are immunoreactive for MAP2 (microtubule associated protein 2). Further, subpopulations are immunoreactive for one of several neuropeptides. These observations suggest that during the fetal and early postnatal development of the mammalian telencephalon the subplate cells function as neurons in synaptic circuitry that disappears by adulthood.

Animals↗
PubMed2006

The Sox2 regulatory region 2 functions as a neural stem cell-specific enhancer in the telencephalon.

Sox2 is expressed at high levels in neuroepithelial stem cells and persists in neural stem/progenitor cells throughout adulthood. We showed previously that the Sox2 regulatory region 2 (SRR2) drives strong expression in these cells. Here we generated transgenic mouse strains with the beta-geo reporter gene under the control of the SRR2 in order to examine the spatiotemporal function of this regulatory region. We show that the SRR2 functions specifically in neural stem/progenitor cells. However, unlike Nestin 2nd intronic enhancer, the SRR2 shows strong regional specificity functioning only in restricted areas of the telencephalon but not in any other portions of the central nervous system such as the spinal cord. We also show by in vitro clonogenic assay that at least some of these SRR2-functioning cells possess the hallmark properties of neural stem cells. In adult brains, we could detect strong beta-geo expression in the subventricular zone of the lateral ventricle and along the rostral migrating stream where actively dividing cells reside. Chromatin immunoprecipitation assays reveal interactions of POU and Sox factors with SRR2 in neural stem/progenitor cells. Our data also suggest that the specific recruitment of these proteins to the SRR2 in the telencephalon defines the spatiotemporal activity of the enhancer in the developing nervous system.

Animals↗
PubMed1999

Comparison of the mammalian and avian telencephalon from the perspective of gene expression data.

Pallial and subpallial morphological subdivisions of the mouse and chicken telencephalon were examined from the new perspective given by gene markers expressed in these territories during development. The rationale of this approach is that common gene expression patterns may underlie similar histogenetic specification and, consequently, comparable morphological nature. The nested expression domains of the genes Dlx-2 and Nkx-2.1 are characteristic for the subpallium (lateral and medial ganglionic eminences). Similar expression of these markers in parts of the mouse septum and amygdala suggests that such parts may be considered subpallial. The genes Pax-6, Tbr-1 and Emx-1 are expressed in the pallium. Complementary areas of the septum and amygdala shared expression of these genes, suggesting these are the pallial parts of these units. Differences in the relative topography of pallial marker genes also define different regions of the pallium, which can be partially traced into the amygdala. Importantly, there is evidence of a novel "ventral pallium" subdivision, which is a molecularly distinct pallial territory intercalated between the striatum and the lateral pallium. Its derivatives in the mouse apparently belong to the claustroamygdaloid complex. Chicken genes homologous sequence-wise to these mouse developmental genes are expressed in topologically comparable patterns during development. The avian subpallium -the paleostriatum- expresses Dlx-2 and Nkx-2.1; expression extends as well into the septum and anterior and medial parts of the archistriatum. The avian pallium expresses Pax-6, Tbr-1 and Emx-1 and also contains a distinct ventral pallium, formed by the neostriatum and ventral intermediate parts of the archistriatum. The lateral pallium comprises the hyperstriatum ventrale, overlying temporo-parieto-occipital corticoid layer and piriform cortex, plus dorsal intermediate and posterior archistriatum. The dorsal pallium includes the dorsal, intercalated and accessory hyperstriatum, plus the dorsolateral corticoid area. The medial pallium contains the hippocampus and parahippocampal area. A dorsal part of the septum shares pallial molecular markers. Gene markers thus suggest common sets of molecular developmental determinants in either pallial or subpallial domains of the mouse and chicken telencephalon, extending all the way from the posterior pole (amygdala) to the septum. Ventral pallial derivatives identified as claustroamygdaloid in the mouse correlate with avian neostriatum and parts of the archistriatum.

Animals↗
PubMed2003

Dlx2 progenitor migration in wild type and Nkx2.1 mutant telencephalon.

The transcription factor Dlx2 is expressed widely throughout the ventral telencephalon. We have examined the in vitro and in vivo migration of Dlx2 progenitors originating from the different ganglionic eminences of both wild type and Nkx2.1 mutant animals. By examining the expression of tauLacZ targeted into the Dlx2 locus we were able to visualize the distribution of cells expressing this gene at both embryonic and postnatal stages. This analysis suggested that Dlx2-expressing cells traverse a number of characteristic migratory routes to populate both cortical and subcortical regions. We also examined how these patterns of migration were affected in Nkx2.1 mutant animals. In these mutants, the early but not late populations of Dlx2-expressing cells originating in the ventral telencephalon that migrate to the cortex are lost. This recovery may be, at least in part, a result of the late migration of Dlx2 progenitors from the caudal ganglionic eminences (CGE), which, based on our previous work, does not appear to require Nkx2.1 gene function.

Animals↗
PubMed2002

Determination of cell fate within the telencephalon.

The telencephalon (basal ganglia, septum, cerebral cortex and olfactory bulb) contains two general classes of neurons: those that project axons to distant targets and those that make only local connections. While projection neurons can be either excitatory (such as those in the olfactory bulb and cortex) or inhibitory (such as those in the striatum), local circuit neurons (interneurons) are usually inhibitory. Within these two general classes of neurons there are a myriad of cell subtypes based upon axonal and dendritic morphology, chemical markers, neurotransmitters, connectivity and physiology. A crucial issue regarding the development of the telencephalon is the molecular determination of neuronal subtypes. Since important aspects of neuronal fate determination occur within the proliferative zone, the consideration of determinants of a mature neuron's fate requires consideration of that cell's origin.

Animals↗
PubMed2002

Telencephalon-specific Rb knockouts reveal enhanced neurogenesis, survival and abnormal cortical development.

Correct cell cycle regulation and terminal mitosis are critical for nervous system development. The retinoblastoma (Rb) protein is a key regulator of these processes, as Rb-/- embryos die by E15.5, exhibiting gross hematopoietic and neurological defects. The extensive apoptosis in Rb-/- embryos has been attributed to aberrant S phase entry resulting in conflicting growth control signals in differentiating cells. To assess the role of Rb in cortical development in the absence of other embryonic defects, we examined mice with telencephalon-specific Rb deletions. Animals carrying a floxed Rb allele were interbred with mice in which cre was knocked into the Foxg1 locus. Unlike germline knockouts, mice specifically deleted for Rb in the developing telencephalon survived until birth. In these mutants, Rb-/- progenitor cells divided ectopically, but were able to survive and differentiate. Mutant brains exhibited enhanced cellularity due to increased proliferation of neuroblasts. These studies demonstrate that: (i) cell cycle deregulation during differentiation does not necessitate apoptosis; (ii) Rb-deficient mutants exhibit enhanced neuroblast proliferation; and (iii) terminal mitosis may not be required to initiate differentiation.

Alleles↗
PubMed1995

5 beta-reductase and other androgen-metabolizing enzymes in primary cultures of developing zebra finch telencephalon.

Enzymes in the avian brain irreversibly catalyze the conversion of androgens into either active metabolites (aromatase and 5 alpha-reductase) or inactive metabolites (5 beta-reductase), 5 beta-reductase is thought to influence the formation of active metabolites by reducing the concentration of androgenic substrate available for these reactions. However, because these enzymes have different regional, cellular and subcellular distributions in brain, the traditional method to measure enzyme activity in brain homogenates may be inaccurate because of artificial mixing of enzymes. Recently, we have prepared primary cell cultures from the telencephalons of developing zebra finches. Cell cultures offer the advantage that enzyme activity can be measured in live cells in which the relative distribution of enzymes may more closely reflect that found in vivo. We have previously reported that aromatase is expressed at high levels in these cultures, and that it is active in both neurons and in glia. Here we report on the activities of 5 alpha- and 5 beta-reductase in these cell cultures. Along with aromatase, 5 beta-reductase was expressed at high levels in these mixed cell cultures, including cultures highly enriched in glia. This suggests that 5 beta-reductase is present in non-neuronal cells in brain, possibly co-localized with aromatase. However, despite the presence of 5 beta-reductase, aromatase was detected reliably in vitro even when the concentration of substrate was low. Thus, 5 beta-reductase does not prevent the synthesis of estrogen in the telencephalon of developing zebra finches. By contrast, 5 alpha-reductase activity was very low or absent in these cultures.(ABSTRACT TRUNCATED AT 250 WORDS)

Androgens↗
PubMed2005

GABAergic modulation of the expression of genes involved in GABA synaptic transmission and stress in the hypothalamus and telencephalon of the female goldfish (Carassius auratus).

GABA is one of the most abundant neurotransmitters in the vertebrate central nervous system and is involved in neuroendocrine processes such as development, reproduction, feeding and stress. To examine the effect of GABA on gene expression in the brain, we used a cDNA macroarray containing 26 genes involved in GABA synaptic transmission (GABA receptor subunits, GABA transporters), reproduction (gonadotrophin-releasing hormone isoforms and oestrogen receptor alpha), feeding (neuropeptide Y and cholecystokinin), and stress [corticotrophin-releasing factor (CRF)]. To elevate GABA levels in the brain, we injected female goldfish with gamma-vinyl GABA (300 microg/g of body weight) (24 h), an irreversible inhibitor of the enzyme GABA transaminase (GABA-T). We found that increased levels of GABA in the hypothalamus resulted in a 2.2-fold down-regulation of GABA(A) receptor beta4 subunit mRNA. In the telencephalon, we found that increased GABA levels resulted in a 1.5-fold increase of CRF mRNA and a 1.8-fold decrease of GABA(A) receptor beta2 subunit mRNA. Increasing GABA in the hypothalamus and telencephalon of the goldfish did not significantly affect the mRNA abundance of genes involved in GABA synthesis (glutamic acid decarboxylase isoforms) and degradation (GABA-T), feeding, or reproduction. Our preliminary study suggests that the regulation of GABA receptor subunit mRNA expression by GABA may be a conserved evolutionary mechanism in vertebrates to modulate GABAergic synaptic transmission.

Animals↗
PubMed2005

RARbeta isoform-specific regulation of DARPP-32 gene expression: an ectopic expression study in the developing rat telencephalon.

Dopamine and adenosine 3':5'-monophosphate-regulated phosphoprotein (DARPP-32) is a key molecule for dopamine neurotransmission. The molecular mechanisms underlying the regulation of DARPP-32 in the developing brain remains elusive. Previous studies have shown that retinoids are capable of inducing DARPP-32 in striatal cell culture, suggesting that retinoids are candidate molecules for controlling DARPP-32 expression. In the present study, we first studied the expression profiles of retinoid receptors and their associated co-factors in the developing rat telencephalon by RT-PCR. The results showed that among the retinoid receptors, RARbeta and RXRgamma were nearly selectively expressed in the developing striatum. By contrast, the retinoid receptors associated transcriptional co-factors, including the co-repressors of N-CoR and SMRT, and the co-activators of SRC-1 and P/CAF, were ubiquitously expressed in the developing telencephalon. In light of the previous findings that DARPP-32 was inducible by retinoids in striatal culture, but not in cortical culture, we hypothesized that the striatum-selective RARbeta and RXRgamma may mediate DARPP-32 induction by retinoids. To test this hypothesis, we used the gain-of-function approach to ectopically express RARbeta and RXRgamma in the developing cerebral cortex that lacked these two retinoid receptors. Ectopic expression of RARbeta1, but not RXRgamma1, up-regulated DARPP-32 in the cortical explant culture. Notably, DARPP-32 was up-regulated only by the RARbeta1 isoform, but not by other RARbeta isoforms. Our study suggests that RARbeta signaling may regulate DARPP-32 gene expression by an isoform-specific mechanism in developing telencephalic neurons. The molecular diversity of RARbeta isoforms may underlie parts of the complex gene regulation by retinoids during neural development.

Animals↗
PubMed1989

Ontogenesis of alpha 2-adrenoceptor coupling with GTP-binding proteins in the rat telencephalon.

The ontogenesis of alpha 2-adrenoceptors and GTP-binding proteins and their coupling activity were investigated in telencephalon membranes of developing rats. The manganese-induced elevation of [3H]clonidine binding was increased in an age-dependent manner but the guanosine 5'-O-(3-thio)triphosphate-induced decrease in binding did not change. The extent of the binding of [3H]clonidine at 15 nM (saturable concentration) increased in an age-dependent manner and reached the adult level at 4 days after birth. Cholera toxin and pertussis toxin catalyzed ADP-ribosylation of proteins of 46 and 41/39 kilodaltons (kDa) in solubilized cholate extracts of the membranes. The 41/39-kDa proteins ADP-ribosylated by pertussis toxin (Gi alpha + Go alpha) were increased with age and reached the adult level at day 12, whereas the 46-kDa protein (Gs alpha) reached its peak on day 12 and then decreased to the fetal level at the adult stage. The immunoblot experiments of the homogenates with antiserum (specific antibody against alpha- and beta-subunit of GTP-binding proteins) demonstrated that the 39-kDa alpha-subunit of (Go alpha) and the 36-kDa beta-subunit of GTP-binding protein (beta 36) increased with postnatal age. In contrast, 35-kDa beta-subunit (beta 35) did not change. From these results, it is suggested that the coupling activity of alpha 2-adrenoceptor with GTP-binding protein gradually develops in a manner parallel with the increase of alpha 2-adrenoceptor and pertussis toxin sensitive GTP-binding proteins, Gi, and that alpha 39 beta 36 gamma may be related to the differentiation and/or growth of nerve cells in rat telencephalon.

Adenosine Diphosphate Ribose↗
PubMed1980

Nonolfactory sensory pathway to the telencephalon in a teleost fish.

Pathways conveying lateral-line sensory information within the brain of a bullhead catfish terminate in a localized zone within the telencephalon. Thus, the telencephalon in teleosts, as in amniote species, contains regions that receive specific sensory input. Therefore, this lemniscal organization is not restricted to mammalian or amniote species but is a feature common to most, if not all, vertebrates.

Afferent Pathways↗
PubMed1984

Response of cerebellum to stimulation of telencephalon in the catfish (Ictalurus nebulosus).

In the siluroid teleost Ictalurus nebulosus, averaged evoked potentials and unit responses were recorded from various parts of the cerebellum following electrical stimulation applied to the telencephalon. Responses are most readily evoked in the cerebellum when area dorsalis pars centralis (Dc), a group of scattered large cells in the middle of the telencephalon, is stimulated. The configuration of the evoked field potentials differs according to the rate of stimulation. At a relatively low repetition rate, 0.5/s or lower, the response takes the shape of a positive wave, Pmol2, peaking at about 58 ms in the molecular layer. In the granular layer, the response is primarily a negative wave, Ngr1, followed by a slow positive deflection, Pgr3, with peak latencies of 52 and 130 ms, respectively. Ngr1 appears to be the result of summed mossy fiber-granule cell synaptic activations, with Pmol2 its concomitant passive source. No neuronal basis can be proposed for Pgr3. At higher rates of stimulation (greater than 2/S), the response to each shock is of longer latency and more complex. The stabilized waveform (i.e., after 5-9 cycles) is dominated by a surface-negative wave, Nmol, peaking at about 80 ms. The negative wave disappears at about 300 micron below the surface (slightly above the level where Purkinje cells are located) where it is replaced by a positive and negative sequence, P-Npur, peaking at about 80 and 160 ms, respectively. Both responses are bilaterally distributed but the ipsilateral responses are usually larger in amplitude and shorter in latency. Rostrocaudally, a latency gradient with longer rostral latencies is also found in both responses. To facilitate further comparative studies, the changes of peak latency and amplitude with stimulus strength of these two patterns of response were examined. Other dynamic characteristics of these two patterns of response were also examined and compared by stimulating with pairs and short trains of electric shocks. In discussing the functional implications of these findings, this telencephalocerebellar pathway appears to be a striocerebellar pathway.

Action Potentials↗
PubMed2003

Connections of the pallial telencephalon in the Senegal bichir, Polypterus.

The connections of the dorsal telencephalon (pallium) of the Senegal bichir (Polypterus senegalus) were investigated in order to test the hypothesis that the simple everted pallium of polypterid fishes is directly comparable to the evaginated pallium of most vertebrates. Neuroanatomical tracers (DiI or biotinylinated dextran amines) were injected into the three traditionally recognized divisions of the bichir pallium to determine their afferent and efferent connections. The connections were corroborated with injections into regions identified as projecting to, or receiving input from, one or more pallial divisions. The results suggest that the bichir pallium consists not of three, but only two primary divisions: a dorsomedial one and a dorsolateral one. This reinterpretation refutes the hypothesis of a direct one-to-one comparison with the tripartite pallium of most vertebrates. Homologues of the bichir dorsomedial and dorsolateral pallium are recognized in the lateral and medial pallium of anuran amphibians, respectively. The pallium of bichirs is compared to that of derived ray-finned fishes as a link between derived ray-finned fishes and other vertebrates. The available information on the connections of the pallium of teleosts suggests that only the olfactory recipient (pars posterior) of the pallium can be directly compared to bichirs and amphibians and that the remaining divisions of the pallium in teleosts are uniquely derived features of the teleost telencephalon.

Animals↗
PubMed1990

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↗
PubMed2003

Loss of Emx2 function leads to ectopic expression of Wnt1 in the developing telencephalon and cortical dysplasia.

Leptomeningeal glioneuronal heterotopias are a focal type of cortical dysplasia in which neural cells migrate aberrantly into superficial layers of the cerebral cortex and meninges. These heterotopias are frequently observed as microscopic abnormalities in the brains of individuals with central nervous system (CNS) malformations and epilepsy. Previous work has demonstrated that the function of Emx2, which encodes a homeodomain transcription factor, is essential for development of the cortical preplate, which gives rise to the marginal zone and subplate. However, transcriptional targets of EMX2 during CNS development are unknown. We report that leptomeningeal glioneuronal heterotopias form in Emx2(-/-) mice that are equivalent to human lesions. Additionally, we observed ectopic expression of Wnt1 in the embryonic roofplate organizer region and dorsal telencephalon. To determine the phenotypic consequences of such Wnt1 misexpression, we deleted a putative EMX2 DNA-binding site from the Wnt1 enhancer and used this to misexpress Wnt1 in the developing murine CNS. Heterotopias were detected in transgenic mice as early as 13.5 days postcoitum, consistent with a defect of preplate development during early phases of radial neuronal migration. Furthermore, we observed diffuse abnormalities of reelin- and calretinin-positive cell populations in the marginal zone and subplate similar to those observed in Emx2-null animals. Taken together, these findings indicate that EMX2 is a direct repressor of Wnt1 expression in the developing mammalian telencephalon. They further suggest that EMX2-Wnt1 interactions are essential for normal development of preplate derivatives in the mammalian cerebral cortex.

Animals↗
PubMed2004

The homeobox gene Gsh2 is required for retinoid production in the embryonic mouse telencephalon.

We have examined the role of the homeobox gene Gsh2 in retinoid production and signaling within the ventral telencephalon of mouse embryos. Gsh2 mutants exhibit altered ventral telencephalic development, including a smaller striatum with fewer DARPP-32 neurons than wild types. We show that the expression of the retinoic acid (RA) synthesis enzyme, retinaldehyde dehydrogenase 3 (Raldh3, also known as Aldh1a3), is reduced in the lateral ganglionic eminence (LGE) of Gsh2 mutants. Moreover, using a retinoid reporter cell assay, we found that retinoid production in the Gsh2 mutants is markedly reduced. The striatal defects in Gsh2 mutants are thought to result from ectopic expression of Pax6 in the LGE. Previously, we had shown that removal of Pax6 from the Gsh2 mutant background improves the molecular identity of the LGE in these double mutants; however, Raldh3 expression is not improved. The Pax6;Gsh2 double mutants possess a larger striatum than the Gsh2 mutants, but the disproportionate reduction in DARPP-32 neurons is not improved. These findings suggest that reduced retinoid production in the Gsh2 mutant contributes to the striatal differentiation defects. As RA promotes the expression of DARPP-32 in differentiating LGE cells in vitro, we examined whether exogenous RA can improve striatal neuron differentiation in the Gsh2 mutants. Indeed, RA supplementation of Gsh2 mutants, during the period of striatal neurogenesis, results in a significant increase in DARPP-32 expression. Thus, in addition to the previously described role for Gsh2 to maintain correct molecular identity in the LGE, our results demonstrate a novel requirement of this gene for retinoid production within the ventral telencephalon.

Animals↗