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PubMed1999

Entorhinal cortex of aged subjects with Down's syndrome shows severe neuronal loss caused by neurofibrillary pathology.

In Alzheimer's disease (AD), neurofibrillary degeneration of neurons starts in the transentorhinal cortex and spreads in a time-dependent manner to the entorhinal cortex, which provides a major input to the hippocampus--a key structure of the memory system. People with Down's syndrome (DS) develop neurofibrillary changes more than 30 years earlier than those with sporadic AD. To characterize AD-related pathology in the entorhinal cortex in DS, we examined seven subjects with DS of 60-74 years of age who died in the end stage of AD, and four age-matched control subjects. The volume of the entorhinal cortex in brains of subjects with DS was 42% less than that in control cases; however, the total number of neurons free of neurofibrillary changes was reduced in DS by 90%: from 9,619,000 +/- 914,000 (mean +/- standard deviation) to 932,000 +/- 504,000. The presence of 2,488,000 +/- 544,000 neurofibrillary tangles in the entorhinal cortex of people with DS, the prevalence of end-stage tangles, and the significant negative correlation between the total number of intact neurons and the percentage of neurons with neurofibrillary changes indicate that neurofibrillary degeneration is a major cause of neuronal loss in the entorhinal cortex of people with DS. The relatively low amyloid load (7 +/- 1%) and lack of correlation between the amyloid load and the volumetric or neuronal loss suggest that the contribution of beta-amyloid to neuronal loss in the entorhinal cortex is unsubstantial.

Aged↗

PubMed1978

Afferent and efferent subcortical projections of behaviorally defined sectors of prefrontal granular cortex.

Although the functional significance of the midprincipalis region is well known, the afferent and efferent connections of this zone, in comparison to the anterior and posterior portions of the cortex lining the principal sulcus, are poorly understood. In 3 animals the retrograde tracer HRP and the anterograde tracers, tritiated proline, lysine and leucine, were injected into the sulcal cortex lining the principal sulcus. The cortex forming the banks of the principal sulcus was divided into anterior, middle and posterior sectors with one animal used for each zone. As expected from previous studies, the heaviest afferents to the cortex forming the principal sulcus were from the parvocellular portions of the medical dorsal nucleus. The medial pulvinar nucleus and the nucleus limitans projected to only the anterior and posterior portions of the cortex lining the principal sulcus. Projections were seen to all 3 sectors from the anterior, midline, intralaminar and lateral thalamic nuclei. Although cells were seen in the hypothalamus following injections in all 3 sectors of the cortex lining the principal sulcus, the heaviest hypothalamic projections were noted after injections into the mid-sector of the cortex. These HRP-positive cells were in the dorsal and lateral hypothalamic area, dorsal medial nucleus and in the lateral mammillary nucleus. These findings link the midprincipalis region with the prefrontolimbic circuit, and suggest that the midprincipalis region, n. medialis dorsalis, the mammillary bodies and perhaps the cingulate gyrus constitute part of an anatomical circuit concerned with memory processes.

Afferent Pathways↗

PubMed1981

Postnatal development of presynaptic alpha-adrenoceptors in rat cerebral cortex: studies with brain slices and synaptosomes.

The development of alpha-adrenoceptor mediated presynaptic modulation of noradrenaline (NA) release in the rat cerebral cortex was studied during the first postnatal month by examining the effects of exogenous NA and phentolamine in K+-induced [3H]NA release from superfused slices or synaptosomes. Exogenous NA strongly inhibited [3H]NA release from cortex slices of newborn rats, indicating that presynaptic alpha-receptors exist already on the day of birth. In fact, at least up to and including the 17th day after birth, Na caused a stronger inhibition of [3H]NA release from neonatal than from adult cortex slices. However, with synaptosomal preparations the inhibitory effect of NA was found to be similar in neonatal and adult rats. Further analysis of the data obtained with brain slices suggested that the differences between neonatal and adult rats observed with regard to the presynaptic modulation of [3H]NA release could be ascribed to differences in the concentrations of released endogenous NA in the extracellular spaces of brain slices. This hypothesis was strengthened by the data obtained in experiments using cortex slices of adult rats in which the endogenous NA level was lowered by pretreatment with alpha-methylparatyrosine. In cortex slices of neonatal (7-day-old) rats the concentration of endogenous NA released upon stimulation with 26 mM K+ in the vicinity of presynaptic alpha-receptors was estimated to be about 10 nM, as compared to approximately 25 nM in adult cortex slices. In both cases NA (i.e. exogenous + endogenous) appeared to cause maximally about 80% inhibition of [3H]NA release from cortex slices.

Aging↗

PubMed1984

Heterotopic and homotopic callosal connections in rat visual cortex.

Heterotopic and homotopic callosal projections of rat visual cortex are evaluated. Callosal termination zones in visual cortex are identified with a degeneration technique following complete section of the corpus callosum. The zones which receive callosal afferents are the lateral one-third of area 17, an anteroposterior strip in dorsal and in ventral areas 18a, and 4 patches in area 18b. Following a large injection of lectin-bound horseradish peroxidase (WGA-HRP) into visual cortex, many retrogradely labeled neurons are found in the medial two-thirds of area 17 which does not receive callosal afferents, as well as in the lateral, callosal-recipient zone. These data suggest that heterotopic callosal pathways exist in visual cortex. Injections of tritiated amino acids into restricted parts of visual cortex show the following heterotopic connections: lateral area 17 projects to dorsal area 18a; medial area 17 projects to lateral area 17 and dorsal area 18a; area 18a projects to lateral area 17 and anteromedial area 18b; area 18b projects to lateral area 17, dorsal area 18a, heterotopic sites in area 18b, and to area 29d . Heterotopic connections are generally less dense than homotopic ones. In addition, heterotopic connections are generally less dense than homotopic ones. In addition, heterotopic projections terminate in the supragranular layers. This contrasts with the homotopic afferents of areas 17 and 18a which have additional strong projections to layer V. The distribution of label through the depth of the cortex in some of the callosal recipient zones has been quantified. Injections of WGA-HRP restricted to areas 17, 18a or 18b corroborate the presence of each of the heterotopic connections described above. Heterotopic afferents originate mostly from layer V neurons, whereas homotopic afferents arise from neurons primarily in layers II-V. Like the afferents, the numbers of callosal projection cells in heterotopic regions are substantially less than that in homotopic sites. heterotopic callosal connections may be one factor responsible for binocular vision and also may provide the basis for large, nonoriented receptive fields of units in layer V of rodent visual cortex.

Animals↗

PubMed1985

The involvement of the anterior cingulate cortex in blood pressure control.

The visceromotor nature of the rat anterior cingulate cortex was investigated by electrically stimulating this area in both anesthetized and awake animals. Initial studies demonstrated that electrical stimulation of any division of the anterior cingulate cortex elicits a significant fall in blood pressure in the sodium pentobarbital-anesthetized rat. Depressor responses were greatest (up to 50% decreases) following stimulation of the ventral third of the anterior cingulate cortex. Heart rate was not altered by cingulate cortex stimulation. In the awake animal, stimulation of previously identified depressor sites in the rostral third of the pregenual cingulate cortex elicited pressor responses. In contrast, stimulation of the caudal third of this cortex elicited depressor responses, and stimulation of the middle third elicited biphasic (pressor followed by depressor) responses. These results indicate that the anterior cingulate cortex is a visceromotor region which may provide a cortical output for the regulation of blood pressure responses associated with learning and or stress.

Anesthesia↗

PubMed1986

Neuropeptides in cerebral cortex of macaque monkey (Macaca fuscata fuscata): regional distribution and ontogeny.

The concentrations of vasoactive intestinal polypeptide, somatostatin and substance P were determined in various cerebral subdivisions of adult and foetal Japanese monkeys (Macaca fuscata fuscata) by specific radioimmunoassays. In adult tissues, the highest level of vasoactive intestinal polypeptide was found in the somatosensory cortex and the lowest level in the occipital cortex. A high level of somatostatin was found in the association cortex (prefrontal, parietal and temporal cortex); the lowest level was noted in the occipital cortex. Substance P was found to be high in prefrontal and temporal cortex. The highest levels of substance P and somatostatin were obtained in the amygdala. Between embryonic 4 and 5.5 months, concentrations of peptides increased dramatically, and in the adult, all neuropeptides in cortical subdivisions significantly decreased. By the gel filtration method, only one immunoreactivity which coeluted with substance P and vasoactive intestinal polypeptide was demonstrated in extracts of 4-, 5.5-month-old and adult monkey cerebral cortex. In contrast, somatostatin immunoreactivity eluted as 3 peaks. Almost 80% of the immunoreactivity co-eluted with synthetic somatostatin, regardless of the age of the tissue. The molecular weights of two larger molecules were determined to be 13 and 3 kdaltons.

Age Factors↗

PubMed1987

Blockade of cholinergic receptors by an irreversible antagonist, propylbenzilylcholine mustard (PrBCM), in the rat cerebral cortex causes deficits in passive avoidance learning.

Studies were made on the effects of blockade of muscarinic acetylcholine (mACh) receptors in the rat cerebral cortex on learning and memory assessed by performance of a step-through passive avoidance task. Bilateral injection of propylbenzilylcholine mustard (PrBCM) into both the frontal and parietal cortex at doses of 2.25 X 4 to 22.5 X 4 micrograms decreased mACh receptors dose-dependently, as assessed by [3H]quinuclidinyl benzilate binding studies. When the training trial of a step-through passive avoidance task was performed 24 h after injection of 7.5 X 4 to 22.5 X 4 micrograms PrBCM into the frontal and parietal cortex, and then a retention test was made 24 h after the training trial, the treated rats showed shorter latencies than controls. In contrast, injection of PrBCM into the occipital cortex had no significant effect on performance in the test. These results confirm the notion that cholinergic neurotransmission in the cerebral cortex, especially the frontoparietal cortex, is important in learning and memory. The effects of injection of PrBCM (22.5 X 4 micrograms) into the frontoparietal cortex on 3 postulated phases of the learning and memory process (i.e. registration, retention and recall) were also examined. When PrBCM was injected 24 h before the training trial, no retention of the task was observed 14 days after the training trial. However, when PrBCM was injected 24 h after the training trial, retention of the task 14 days after the training trial was not affected. When PrBCM was injected 3-24 h after the initial training trial, the latencies in the retention test examined 24 h later were shorter than those of control rats.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1988

Immunocytochemical localization of somatostatin in the cerebral cortex of lizards.

The distribution of somatostatin-like immunoreactivity in the cerebral cortex of two lizards has been studied. Results are similar in both species. Somatostatin-positive neurons show variable morphology; they are bipolar, multipolar or pyramidal cells. Their distribution within the cerebral cortex is not homogeneous: they tend to be found in the innermost cortical layer, the deep plexiform layer, where they constitute a constant population in the region of the dorsomedial cortex. All immunoreactive processes observed in the cerebral cortex belong to somatostatin cortical neurons since no immunoreactive fiber is found to reach or leave the telencephalic cortex. Most of the dendrites are smooth or sparsely spinous. Axons are abundant in the deep plexiform layer; in the dorsomedial cortex a prominent terminal field appears in the outermost region of the superficial plexiform layer, which may arise from the neuronal somatostatin immunoreactive population found deeper in the same cortex.

Animals↗

PubMed1989

Serotonin innervation of the cerebral cortex in lizards.

The serotonin (5-HT) innervation of the cerebral cortex in two species of lizards has been studied. Results show no differences between both species. Most of the cerebral cortex of these lizards is innervated by serotoninergic fibers, which are fine and varicose. Their density varies greatly from one cortical region to another: the areas with higher density of serotoninergic fibers and terminals are parts of the medial and dorsal cortices. There is a laminar pattern of distribution of serotoninergic fibers. In the medial cortex, 5-HT fibers are found preferentially in both plexiform layers just above and below the cellular layer. In the dorsomedial cortex, there is an immunoreactive plexus in the outermost third of the superficial plexiform layer and another in the depth of the layer, whereas 5-HT fibers are distributed evenly in the deep plexiform layer of this cortex. In the pars medialis of the dorsal cortex, serotoninergic fibers are abundant in all layers, whereas in the pars lateralis, fibers are found predominantly in the external third of the superficial plexiform layer. The lateral cortex is almost devoid of immunoreactive fibers. These results show a different organization of the cortical serotonin innervation between lizards and turtles.

Animals↗

PubMed1991

Peptide heterogeneity of GABAA/benzodiazepine receptors in bovine cerebral cortex and cerebellum.

The GABAA/benzodiazepine receptor complex has been purified from both bovine cerebral cortex and cerebellum by immunoaffinity chromatography on immobilized monoclonal antibody 62-3G1. Sodium dodecyl sulfate-polyacrylamide gel electrophoresis of the purified receptor from either cerebral cortex or cerebellum revealed 3 main bands corresponding to 51,000, 55,000 and 57,000 Mr silver-stained peptides. In addition, a minor band corresponding to a 53,000 Mr peptide was also found. The differences between the two receptor preparations were: (1) that the main silver-stained 55,000 Mr subunit was present in a relative smaller quantity in cerebellum than in cerebral cortex, and (2) when the membrane-bound receptor was photoaffinity-labeled with [3H]flunitrazepam and subsequently immunoaffinity-purified, two photolabeled peptide bands of 51,000 and 57,000 Mr were found in cerebral cortex while only the 51,000 Mr photolabeled peptide was detected in cerebellum following one-dimension sodium dodecyl sulfate-polyacrylamide gel electrophoresis. Peptide maps of the 57,000 Mr [3H]flunitrazepam photoaffinity-labeled peptide indicated that it was composed of two closely migrating photolabeled peptides of 55,000 Mr and 57,000 Mr. Peptide mapping and deglycosylation experiments using the [3H]flunitrazepam photolabeled receptor suggested that the photolabeled peptides commonly present in cerebellum and cerebral cortex are qualitatively similar if not identical. The results suggest that there are subunits of some type(s) of GABAA R/BZDR complex(es) which are more abundant in cerebral cortex than in cerebellum. Photoaffinity labeling with [3H]muscimol showed similar photolabeled peptides in both cerebral cortex and cerebellum: two main peptides of 54,000 and 57,000 Mr were photolabeled with [3H]muscimol to a similar extent in both receptor preparations.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1996

Enhanced acetylcholine release in the hippocampus and cortex during acquisition of an operant behavior.

The activity of the septo-hippocampal and nucleus basalis-cortical cholinergic pathways was investigated by measuring changes in the extracellular acetylcholine levels in the hippocampus and parietal cortex, by means of transversal microdialysis, during the acquisition and recall of a positively reinforced operant behavior. Adult male Wistar rats were trained in a sound-isolated operant chamber equipped with a single lever. The positive reinforcement was represented by food pellets and the number of cumulative reinforced responses was recorded every 30 min. Five groups of rats were used. Unoperated animals were used as controls. In two groups of untrained animals, the microdialysis tubes were transversally implanted in the parietal cortex, and hippocampus and the training in the operant behavior chamber began 24 h after surgery. In two further groups the microdialysis tubes were implanted in the parietal cortex, and hippocampus after training for 15 days in the operant chamber. Food was removed 12 h before training. The time needed by the control rats to reach a stable baseline of reinforced responses was 83 +/- 12 min, while in the untrained rats implanted with dialysis probes in the cerebral cortex and in the hippocampus was 621 +/- 129 and 521 +/- 126 min, respectively, and in those pretrained and implanted in cerebral cortex and in the hippocampus was 116 +/- 38 and 217 +/- 59 min, respectively. In the untrained operated rats, both cortical and hippocampal extracellular acetylcholine levels remained constant until the number of reinforced responses was low but increased significantly (+156% in the cortex and +183% in the hippocampus) in the first 30 min period in which there was a sharp rise in the reinforced responses. In the pretrained operated rats, neither in the cortex nor in the hippocampus was the increase in response rate accompanied by a statistically significant increase in extracellular acetylcholine levels. Our findings demonstrate that activation of the forebrain cholinergic pathways occurs during the acquisition of a rewarded operant responses, while recall of the same behavior is not associated with the activation of the cholinergic system.

Acetylcholine↗

PubMed1984

Cadmium/zinc relationships in kidney cortex and metallothionein of horse and red deer: histopathological observations on horse kidneys.

Cadmium and zinc were determined in kidney cortex of 63 horses and 51 red deer (Cervus elaphus). Cadmium and zinc were also determined in protein fractions obtained by Sephadex chromatography of kidney cortex from 10 horses and 4 red deer. Histopathological parameters in kidney cortex of horses were compared to cadmium content. The metal contents (on wet weight basis) in kidney cortex of the horses were 0.31 +/- 0.22 mmole Cd/kg (range 0.03-1.21) and 0.63 +/- 0.17 mmole Zn/kg (range 0.36-1.23). The Zn content increased with the Cd content, the Zn increase being less at higher concentrations. No significant increase of the Cd content with age of the horses (range 2-19 years) was found. The metal contents in kidney cortex of the red deer were 0.030 +/- 0.031 mmole Cd/kg (range 0.002-0.13) and 0.51 +/- 0.37 mmole Zn/kg (range 0.30-2.82). Here the Zn content did not increase significantly with Cd content, and in this case a positive linear relation of Cd content with age of the deer (range 0-11 years) was observed. The molar Zn/Cd ratio in the metallothionein fractions of horse kidney cortex was less variable at higher Cd contents. All Cd and the increase of Zn above an average basal level of 0.36 mmole Zn/kg was recovered in the metallothionein fractions. No gross histopathological changes could be observed in kidney cortex of the horses. Nevertheless a slight increase of the thickness of the basement membrane of Bowman's Capsule and also of the diameter of Bowman's Capsule in relation to Cd content up to 0.3 mmole/kg could be observed.

Animals↗

PubMed1984

Transition from spindles to generalized spike and wave discharges in the cat: simultaneous single-cell recordings in cortex and thalamus.

The relationships between the activity of the cortex and that of a "specific" (n. lateralis posterior, LP) and an intralaminar thalamic nucleus (n. centralis medialis, NCM) were studied in the cat during the transition from spontaneous spindles to generalized spike and wave (SW) discharge following i.m. penicillin injection. The EEG and extracellular single-unit activity were recorded in cortex and thalamus during the spindle stage and at different intervals after penicillin until well developed SW discharges were present. Computer-generated EEG averages and histograms of single-unit activity were triggered by either peaks of cortical or thalamic EEG transients or by cortical or thalamic action potentials. In agreement with previous observations, cortical neurons increasingly fired during the spindle wave as it was transformed into the "spike" of the SW complex, while a period of neuronal silence gradually developed as the "wave" of the SW complex emerged. Similar changes developed in the thalamus, particularly in LP, either concurrently with or more often after the onset of the changes in the cortex. Most neurons in NCM, continued to fire randomly even after well developed SWs and rhythmic neuronal discharges had developed in cortex and LP. Only 4/11 NCM neurons did ultimately exhibit a rhythmic firing pattern similar to that seen in the cortex and LP. The correlation between cortical and thalamic unit activity was low during spindles, but gradually increased during the development of SW discharges. These data confirm that the cortex is the leading element in the transition from spindles to SWs. Increasingly, in the course of this transition, cortical and thalamic neuronal firing becomes more intimately phase-locked. This mutual interrelationship appears to be more pronounced between cortex and "specific" than intralaminar thalamic nuclei.

Action Potentials↗

PubMed1986

Maternal ethanol consumption: lack of effect on synaptogenesis in layer I of the motor cortex in 19-day-old rat offspring.

Previous studies showed that in utero exposure to ethanol has an adverse effect on development of the central nervous system. Included among those abnormalities is the evidence that in utero exposure to ethanol alters the development of the cerebral cortex in terms of the arborization of pyramidal dendrites and the concentration of neurotransmitters and neurotransmitter receptors. However, limited information is available about the influence of in utero exposure to ethanol on synaptogenesis in the cortex. We examined the influence of chronic maternal ethanol consumption, prior to parturition, on synaptogenesis in layer I of the motor cortex of 19-day-old-rat offspring. Layer I (the molecular layer) was chosen because of the early timing of its development. The motor cortex was chosen because of the evidence of motor problems in children afflicted with the fetal alcohol syndrome. The results of these studies demonstrated that the 19-day-old offspring of rats that were pair-fed, using control or 6.6% (v/v) ethanol-containing liquid diets on a chronic basis prior to parturition, had a comparable density of synapses and distribution of paramembranous density lengths in layer I of the motor cortex as well as a comparable thickness of cortical layer I. Thus, evidence by this and other laboratories, of cortical neurotransmitter and dendritic abnormalities are not reflected in synaptic changes in layer I of the motor cortex at 19 days. It is possible that synaptogenesis is affected at another age or in other regions of the cerebral cortex.

Animals↗

PubMed1993

Antagonists of N-methyl-D-aspartate receptors block seizures induced by putrescine in the deep prepiriform cortex.

The role of excitatory amino acid receptors in the genesis of motor and electrocortical seizures, elicited by administration of the polyamine putrescine into the deep prepiriform cortex, has been evaluated in rats. Motor and electrocortical seizures occurred in rats receiving unilateral local injections into the deep prepiriform cortex, of putrescine (10 or 20 nmol). The selective N-methyl-D-aspartate receptor antagonist, 2-amino-7-phosphonoheptanoate (AP7), injected previously (15 min) into the deep prepiriform cortex, prevented the development of seizures induced by putrescine, injected at the same site. In addition, dizocilpine (MK-801), a non-selective NMDA antagonist or ifenprodil, a specific inhibitor of the polyamine site at the NMDA receptor, when injected into the deep prepiriform cortex, 15 min prior to putrescine, significantly protected against seizures elicited by this polyamine. A subconvulsant dose of putrescine (5 nmol) potentiated the convulsant effects of NMDA, when injected into the deep prepiriform cortex. These data indicate a potential role of polyamines in the genesis of seizures, elicited from the deep prepiriform cortex. They further suggest that activation of the polyamine site, located at excitatory amino acid NMDA receptors, within the deep prepiriform cortex, may contribute to the genesis of seizure activity in this area.

2-Amino-5-phosphonovalerate↗

PubMed1993

Effects of bilateral lesions of auditory cortex in mice on the acoustic startle response.

The acoustic startle response (ASR) was used to investigate the effects of auditory cortical lesions on a brain stem-mediated auditory behavior. The ASRs were obtained longitudinally from young adult C57BL/6J mice before bilateral ablation of auditory cortex, 1 day after ablation, and 1 month later. Control mice received lesions of nonauditory cortex. For some mice, averaged brain stem-evoked responses (ABR) were obtained, and these indicated no effects of lesions on auditory sensitivity. One month after surgery, mice with auditory cortex ablations were statistically indistinguishable from controls on all suprathreshold measures of ASR. However, 1 day after ablation of auditory cortex, experimental animals (but not controls) exhibited a change in ASR amplitude (but not threshold or latency). When a noise burst of 80 dB SPL was used to elicit the ASR, the amplitude was diminished, but with a 110 dB stimulus, amplitude was enhanced. The findings can be interpreted in one of two ways: temporary interference with modulation of the ASR normally performed by auditory cortex; or a general effect of auditory cortex ablation on brain stem auditory circuits not specific to the ASR. In any event, if auditory cortex plays a modulatory role with regard to the ASR, it is apparently nonessential and/or readily compensated for after ablation.

Acoustic Stimulation↗

PubMed1991

Resection of intrinsic tumors from nondominant face motor cortex using stimulation mapping: report of two cases.

We report two right-handed patients who underwent resection of intrinsic glial tumors from the nondominant hemisphere, face motor cortex. Both patients underwent preoperative assessment with computed tomography and magnetic resonance imaging localizing the tumor in the inferior region of the Rolandic cortex. With the patients under general anesthesia and without muscular paralysis, the tumor volume was determined by intraoperative ultrasound and resective surgery accomplished with the aid of cortical and subcortical stimulation mapping techniques. Radical resection of the tumor from the face motor cortex was achieved in both patients. A transient contralateral facial weakness and apraxia were noted in each patient, and this resolved within 6 to 8 weeks following surgery. Removal of intrinsic tumors involving the nondominant face motor cortex may be safely achieved using brain mapping techniques to localize inferior Rolandic cortex and avoid resection of the hand motor cortex and descending subcortical motor pathways. Permanent disability will be prevented due to the bilateral representation of face motor function at the neocortical level. However, due to language localization in cortical zones contiguous with the dominant hemisphere, face motor cortex, we do not recommend resection of this region.

Adolescent↗

PubMed1982

Development of the dopaminergic innervation of the rat cerebral cortex. A light microscopic immunocytochemical study using anti-tyrosine hydroxylase antibodies.

A precise topographical analysis of the distribution of tyrosine hydroxylase-like immunoreactive processes was performed in the frontal, cingular and parietal cortex of the rat during late embryonic and early postnatal life. Until birth, labeled processes were only observed in the restricted cortical areas known to receive a dopaminergic innervation in the adult brain. Their distribution differed markedly from that of noradrenergic fibers as identified by their dopamine-beta-hydroxylase-like immunoreactivity. Thus we considered TH-like immunoreactivity to be a selective marker of the cortical dopaminergic innervation during late fetal life, at least with the antibody we used. With this marker, dopaminergic fibers were first detected in the anterior frontal cortex at day 16 of embryonic life (E16). They developed as two bundles passing medially and laterally to the ventricular layer without penetrating it. From E20 on, the terminal fields extended to the cingular and rhinal cortex, still being restricted to the intermediate zone. No fibers were visible in the lateral and dorsal frontal cortex at this time, nor in the cortical plate and molecular layer in any cortical area. At E21, rare labeled fibers were seen in the molecular layer of the medial frontal and cingular cortex. After birth, the terminal fields of the TH-containing fibers extended further caudally in the cingular cortex and also superficially in the cortical plate. Moreover, labeled axons now also appeared in the lateral frontal and parietal cortex where their density gradually increased. At P14, two different patterns of distribution were observed: a high density of TH-positive fibers in the cortical areas known to receive a dopaminergic innervation; a low density of fibers in the other cortical areas which represented noradrenergic fibers. Indeed these TH-containing presumed noradrenergic fibers were absent at P14 following a bilateral destruction of the locus coeruleus in 4-day-old pups.

Animals↗