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PubMed1996

Evidence for the Hebbian hypothesis in experience-dependent physiological plasticity of neocortex: a critical review.

Over the past decade, the number of experimental papers reporting physiological plasticity in primary neocortical regions, following certain types of controlled sensory experience, have increased greatly. These reports have been characterized by specific changes in receptive fields of individual neurons and/or the distributions of receptive fields across cortical maps. There is a widespread belief these types of plasticities have underlying Hebbian/covariance induction mechanisms. This belief appears to be based mainly on: (a) indirect evidence, largely from experiments on the kitten visual cortex, indicating that Hebbian induction mechanisms could be involved in neocortical plasticity; (b) the observation that some types of plasticity in systems other than neocortex follow Hebbian rules of induction; and (c) the adaptability of Hebbian induction mechanisms to models of neural plasticity. In addition, some experiments have directly tested the role of Hebbian induction mechanisms in experience-dependent neocortical plasticity. The present review critically analyzes these (and related) experiments, in order to evaluate the evidence for the Hebbian Hypothesis in experience-dependent physiological plasticity of neocortex. First, we present a set of criteria to show the involvement of a Hebbian process in any form of plasticity. Next, we compare evidence from each primary neocortical region to these criteria. Finally, we examine unresolved issues. While selected developmental studies are included, emphasis is placed on plasticity in the adult neocortex. It is concluded that there is some evidence meeting the criteria for the Hebbian hypothesis in neocortical plasticity. However, this evidence is quite limited considering the popular belief in the validity of the Hebbian hypothesis.

Animals↗

PubMed1997

Empirical assessment of synapse numbers in primate neocortex.

Reliable methods are needed to assess the impact of synaptic loss on brain function. In this empirical study we demonstrate a novel and efficient method using immunocytochemistry (ICC) and modern stereological techniques to quantify synapses in neocortex of adult primates (Macaca fascicularis). Systematic-uniform-random sections through forebrain from two 10-year-old monkeys were immunostained for estimation of synaptophysin-immunoreactive (synaptophysin-IR) presynaptic boutons (synapses). Adjacent sections were stained with cresyl violet for estimation of total number of neuronal cell bodies. The unbiased Cavalieri method was used to estimate total forebrain and neocortical volumes to a high level of precision (coefficient of error (CE) < or = 0.10)). Synapse-to-neuron ratios varied from 860 in frontal cortex to 2300 in parietal-temporal cortex. The combination of Cavalieri and optical disector methods provided a direct means of estimating approximately 1.25 trillion (x 10(12)) total synaptophysin-immunopositive boutons and approximately 1.01 billion (x 10(9)) cell bodies in neocortex, with low CEs (0.12). Time required to make precise estimates of total neocortical and forebrain volumes and total numbers of synapses and neurons in neocortex was approximately 2-3 h per case from stained sections. The approach is a direct and efficient technique to quantify total synapse and neuron numbers within a defined brain structure.

Animals↗

PubMed1989

Tongue protrusion mediated by spared anterior ventrolateral neocortex in neonatally decorticate rats: behavioral support for the neurogenetic hypothesis.

Many changes in anatomical organization and behavior follow circumscribed, neonatal cortical ablations. These include functional sparing and compensatory anatomical changes. An objective of this study was to examine the generality of such changes by reversing the usual experimental procedure, removing all cortex but a circumscribed area and examining whether the remnant made new anatomical connections, adopted new functions and whether it continued to subserve the typical functions of that area. All neocortex and cingulate cortex, except a small portion of anterior-lateral neocortex, which is normally involved in tongue and mouth use, was removed from one-day-old or adult rats. Fluorescent labelling and behavioral tests were used to evaluate its function. The results showed: (1) The remnant cortical tissue maintained similar connections in neonatal and adult groups and similar connections to those found in rats that had received no lesions. (2) The rats still displayed behaviors normally supported by this cortex, including tongue protrusion to obtain food and picking up and eating hard food efficiently. (3) Impairments were obtained on tests normally mediated by the ablated cortex, including skilled reaching and grooming. (4) When the cortical remnant was removed, tongue protrusion and efficiency of food consumption were similarly impaired in both neonate and adult groups. An additional serendipitous finding was a dissociation between two types of tongue movement: licking from a ventrally-located surface survived cortical removal but tongue protrusion did not. The results show that bilaterally spared small remnants of neocortex maintain normal functions and do not assume new functions or make new connections despite neonatal decortication. The results provide behavioral support for the neurogenetic hypothesis, which postulates that cortical circuitry is specified during early embryonic development. This suggests that there are constraints on neural remodelling and behavioral recovery following neonatal lesions. The implications of these results are discussed with respect to the documented remodelling and sparing that occur following partial or unilateral lesions within functional systems.

Animals↗

PubMed1976

Preservation of cross-modal transfer of a rate discrimination in the bushbaby (Galago senegalensis) with lesions of posterior neocortex.

The effect of lesions of posterior neocortex was assessed, using a test method that permits the demonstration of cross-modal transfer in intact bushbabies. Eight bushbabies were trained to discriminate light flashes of 18/sec and 3/sec in a go-no-go shock-avoidance task. On completion of training, four bushbabies received lesions of posterior neocortex by aspiration. After 6 wk both lesion and intact animals were returned to training in the visual discrimination. On the day following criterion performance on the visual tests, auditory clicks off the same rate and contingencies were substituted and maintained to criterion. All eight bushbabies demonstrated rapid transfer and the lesion animals were not retarded as compared with intact subjects. The cross-modal transfer of a specific rate discrimination was thus preserved in the absence of posterior intersensory neocortex. The results are discussed in terms of a hypothetical subcortical system capable of the amodal coding of simple stimulus dimensions.

Animals↗

PubMed1978

Behavior of the rat after removal of the neocortex and hippocampal formation.

After surgical removal of the neocortex and hippocampal formation, rats retained most of the movement patterns of locomotion, climbing, grooming, feeding, and fighting. However, forepaw immobility during swimming was abolished. Feeding behavior was suppressed temporarily but recovered partially. The distinctive postures of sleep and walking and a circadian rhythm of motor activity were retained. However, behaviors were often not performed at the appropriate time and place. The normal sequence of grooming behavior was disrupted; food hoarding and social behavior were essentially abolished. Removal of the neocortex alone had much the same effect as removal of neocortex and hippocampus together. Removal of hippocampus alone produced only a mild disruption of behavior. It is suggested that ascending nonspecific projections to the cerebral cortex play an important role in the moment-to-moment control of behavior but are not essential for the sleep-waking cycle.

Aggression↗

PubMed2000

Cellular and network mechanisms of rhythmic recurrent activity in neocortex.

The neocortex generates periods of recurrent activity, such as the slow (0.1-0.5 Hz) oscillation during slow-wave sleep. Here we demonstrate that slices of ferret neocortex maintained in vitro generate this slow (< 1 Hz) rhythm when placed in a bathing medium that mimics the extracellular ionic composition in situ. This slow oscillation seems to be initiated in layer 5 as an excitatory interaction between pyramidal neurons and propagates through the neocortex. Our results demonstrate that the cerebral cortex generates an 'up' or depolarized state through recurrent excitation that is regulated by inhibitory networks, thereby allowing local cortical circuits to enter into temporarily activated and self-maintained excitatory states. The spontaneous generation and failure of this self-excited state may account for the generation of a subset of cortical rhythms during sleep.

Action Potentials↗

PubMed1980

The basic uniformity in structure of the neocortex.

The number of neuronal cell bodies has been counted in a narrow strip (30 micrometers) through the depth of the neocortex in several different functional areas (motor, somatic sensory, area 17, frontal, parietal and temporal and in many species (mouse, rat, cat, monkey and man). With the exception of area 17 of the visual cortex in a number of primates the same absolute number (congruent to 110) of neurons has been found in all areas and in all species. In the binocular part of area 17 of the primates there are approximately 2.5 times more neurons. Thus in mammalian evolution the area of the neocortex increases in larger brains but the number of neurons through the depth remains constant, except in area 17 of primates. From these and other findings it is suggested that the intrinsic structure of the neocortex is basically more uniform than has been thought and that differences in cytoarchitecture and function reflect differences in connections.

Adult↗

PubMed1997

The columnar organization of the neocortex.

The modular organization of nervous systems is a widely documented principle of design for both vertebrate and invertebrate brains of which the columnar organization of the neocortex is an example. The classical cytoarchitectural areas of the neocortex are composed of smaller units, local neural circuits repeated iteratively within each area. Modules may vary in cell type and number, in internal and external connectivity, and in mode of neuronal processing between different large entities; within any single large entity they have a basic similarity of internal design and operation. Modules are most commonly grouped into entities by sets of dominating external connections. This unifying factor is most obvious for the heterotypical sensory and motor areas of the neocortex. Columnar defining factors in homotypical areas are generated, in part, within the cortex itself. The set of all modules composing such an entity may be fractionated into different modular subsets by different extrinsic connections. Linkages between them and subsets in other large entities form distributed systems. The neighborhood relations between connected subsets of modules in different entities result in nested distributed systems that serve distributed functions. A cortical area defined in classical cytoarchitectural terms may belong to more than one and sometimes to several distributed systems. Columns in cytoarchitectural areas located at some distance from one another, but with some common properties, may be linked by long-range, intracortical connections.

Animals↗

PubMedDate not supplied

Gap junctional communication and the development of local circuits in neocortex.

In the neocortex, as well as in many other brain regions, neurons responding to similar stimulus features are usually found close to one another. Here we examine the possible role of gap junctional communication in forming and defining these local neuronal groupings, examples of which may be the columns found in the neocortex of virtually all mammalian species. We have approached this question experimentally in cortical brain slices using calcium imaging to visualize multicellular activity patterns, and tracer injections to identify the anatomical pattern of gap junction coupling in the developing neocortex. Our results suggest that dendrodendritic gap junctional communication may be involved in the formation of local connectivity, most likely by synchronizing electrical or biochemical activity among neighboring neurons.

Animals↗

PubMed1994

NADPH diaphorase neurones are evenly distributed throughout cat neocortex irrespective of functional specialization of each region.

Using NADPH diaphorase histochemistry as a marker for nitric oxide synthase we investigated the possible sites of nitric oxide synthesis in cat cerebral neocortex. Intensely stained neurones were found mainly in the deep layers of the neocortex and underlying medulla. Virtually all neurones in the cerebral medulla were NADPH diaphorase positive. The density of diaphorase neurones was estimated in the cortex/medulla border zones of each neocortical gyrus. Diaphorase neurones were evenly distributed throughout the neocortex and no significant statistical difference between gyri was observed. These findings indicate that the density of diaphorase neurones is irrespective of functional specialization of each region and are more in line with the hypothesis that NADPH diaphorase neurones are involved in the control of local cortical blood flow.

Amino Acid Oxidoreductases↗

PubMed1990

The total number of neurons in the human neocortex unbiasedly estimated using optical disectors.

An efficient method is presented for obtaining, in under 4 h, an unbiased estimate of the total number of neurons in the human neocortex, with a coefficient of error on the estimate of approximately 5%. The novel sampling scheme used in this study is unbiased and was designed so that only a small amount of neocortical grey matter had to be removed. Hence, the majority of the cerebral grey matter and all the internal grey matter was left intact for further resampling and analysis. Each cerebral hemisphere was subdivided into the four major neocortical regions, sliced coronally at 7-mm intervals and the volume of the neocortex determined using Cavalieri's principle. Uniform sampling of neocortex was performed in the hemisphere followed by regional subsampling with a varying sampling fraction being taken from each region. Neuronal density estimates were made in thick plastic sections using optical disectors. Shrinkage estimates were made in parallel with the number estimates and found to be negligible. The total number of neocortical neurons in the right hemisphere of five normal 80-year-old men was found to be 13.7 x 10(9) with an inter-individual coefficient of variation of 12%.

Aged↗

PubMed1985

Cyclic AMP concentrations in rat neocortex and hippocampus during and following incomplete ischemia: effects of central noradrenergic neurons, prostaglandins, and adenosine.

The concentrations of cyclic AMP, noradrenaline, glycogen, glucose, lactate, pyruvate, labile phosphate compounds, and free fatty acids were investigated in the rat neocortex and hippocampus during and following cerebral ischemia. An incomplete ischemia of 5 and 15 min duration was induced by bilateral carotid clamping combined with hypotension. The postischemic events were studied after 5, 15, and 60 min of recirculation. Five minutes of ischemia did not significantly alter the neocortical or hippocampal concentrations of cyclic AMP. After 15 min of ischemia the neocortical levels decreased significantly below control values. In the recirculation period following ischemia a significant elevation of the cyclic AMP concentrations was observed. Following 5 min of recirculation after 5 min of ischemia the levels increased from 2.53 +/- 0.21 nmol X g-1 to 5.18 +/- 0.09 nmol X g-1 in the neocortex and from 2.14 +/- 0.16 nmol X g-1 to 3.52 +/- 0.35 nmol X g-1 in the hippocampus. Five minutes of recirculation following 15 min of ischemia led to a significant increase in the levels of cyclic AMP, to 12.86 +/- 1.43 nmol X g-1 in the neocortex to 5.58 +/- 0.57 nmol X g-1 in the hippocampus. With longer recirculation periods the cyclic AMP levels progressively decreased and were similar to control values after 60 min. Depletion of cortical noradrenaline by at least 95% was performed by injections of 6-hydroxydopamine into the ascending axon bundles from the locus ceruleus. The lesion did not significantly change the ischemic or post-ischemic neocortical and hippocampal levels of cyclic AMP, glycogen, or free fatty acids including arachidonic acid.(ABSTRACT TRUNCATED AT 250 WORDS)

Adenosine↗

PubMed1989

Noradrenaline metabolism in neocortex and hippocampus following transient forebrain ischemia in rats: relation to development of selective neuronal necrosis.

Noradrenaline (NA) metabolism in the neocortex and hippocampus was examined in rats at 1, 24, and 48 h following 15 min of reversible forebrain ischemia. As assessed by the ratio of accumulated 3,4-dihydroxyphenylalanine (DOPA) to the tissue NA level after inhibition of DOPA decarboxylase, the NA turnover rates were markedly increased (120-148% above the control) at 1 h postischemia in both the neocortex and hippocampal formation (CA1 and CA3 plus dentate gyrus). The DOPA:NA ratio went back to control levels after longer postischemic survival times. The ratio between levels of the deaminated NA metabolite, 3,4-dihydroxyphenylethyleneglycol (DOPEG), and NA, which gives another measure of NA turnover rate, showed similar changes. In the neocortex and the CA3 plus dentate gyrus, the DOPEG:NA ratio was markedly increased (89-118%) 1 h after the ischemia, but this change had disappeared at 24 and 48 h. Thus, both the DOPA accumulation experiments and the NA and DOPEG measurements indicate that following transient forebrain ischemia, there is an increased NA turnover in the hippocampus and cortex only in the early recirculation period and not after longer postischemic survival times. The degree of neuronal necrosis in the CA1 region was examined light microscopically on celestine blue-acid fuchsin-stained sections at 24, 48, and 96 h following the ischemic insult. The neuronal damage in CA1 was sparse after 24 h of recovery, had increased markedly after 48 h, and was very pronounced at 96 h.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1996

Coevolution of the mammalian middle ear and neocortex.

Phylogenetic analysis with x-ray computed tomography of fossilized and recent crania implicates differential growth of the neocortex in the evolution and development of the mammalian middle ear. In premammalian tetrapods, the middle ear evolved as a chain of bones attached to the mandible and cranium, but in adult mammals the chain is detached from the mandible and lies behind it. The neocortex evolved concurrently with detachment of the chain. In mammalian development the auditory chain arises connected to the mandible but later detaches, recapitulating the phylogenetic transformation. In modern didelphid development, the auditory chain reaches mature size by the third week after birth and is then separated from the jaw and displaced caudally as the neocortex grows for another 9 weeks.

Animals↗

PubMedDate not supplied

Intracortical dentate fascia grafts: mossy fiber synapses in the host neocortex.

Embryonic dentate fascia was grafted into a cavity in the area of the adult rat neocortex which represents the vibrissae (barrel field). We wished to test the possibility of development of connections between the two brain areas which do not have synaptic or tissue contacts in situ. The unique characteristics of the giant synaptic boutons of the dentate mossy fibers were used for detection of the dentate synaptic contacts with neocortical neurons at the electron microscopic level. Ultrastructural analysis nine months postgrafting has shown that the bundles of mossy axons enter the host neocortex and develop multiple terminal and en passant contacts with typical characteristics. Neuronal perikarya, large dendritic trunks and fine caliber terminal dendritic branches were used by the mossy fibers as postsynaptic targets, as well as spines of various complexity and configurations. The subsynaptic dendrites seemed to be modified by synapsing giant boutons. Accumulation of cytoplasmic organelles was observed at these sites. Various bumps and protuberances were formed by the subsynaptic dendrite surface. The contents of these appendages were variable; some of them contained organelles typical of dendroplasm, while others were more spine-like, often with inclusion of ribosomes. It is concluded that mossy fibers growing into the host neocortex can develop typical contacts with inappropriate targets with all the ultrastructural features of functional synapses.

Animals↗

PubMed1981

An analysis of some thalamic projections to parietofrontal neocortex in the marsupial native cat, Dasyurus viverrinus (Dasyuridae).

Thalamocortical projections to parietofrontal neocortex in the marsupial native cat, Dasyurus viverrinus, were examined using the retrograde transport of horseradish peroxidase. The results show that the organisation in Dasyurus is similar to that reported in another Australian marsupial, the brush-tailed possum, Trichosurus vulpecula, and is different to that found in the primitive American form, Didelphis virginiana. In Daysurus, as in Trichosurus, the areas of neocortex receiving projections from the trigeminothalamic system do not appreciably coincide with those areas receiving input from the cerebellothalamic system. In Didelphis there appears to be complete overlap of these areas of motor and somaesthetic neocortex. These results suggest that the motor--somaesthetic organisation in Australian marsupials is divergent from the presumed stem pattern seen in the American forms. If the Australian marsupials, as is now believed, were isolated from the stem Polyprotodonta in the upper Cretaceous, it would appear that the Australian marsupials may have begun to develop certain aspects of their characteristic CNS configuration very early, as the oldest known marsupial fossils are not much older than this.

Animals↗

PubMed1989

Comparative distribution of acid phosphatase and simple esterase in the mouse neocortex and hippocampal formation.

The present study deals with the detailed distribution of acid phosphatase (AcP) and simple esterase (SE) in different layers of the neocortex and hippocampal formation of the mouse brain. The neurons, in general, had moderate to intense enzyme activity for AcP and mild to moderate activity for SE. The AcP activity dominated in the neuronal population as compared to the neuropil; the neuropil stained mildly for SE. The large pyramidal cells in the neocortex and cornu ammonis, and the granular cell layer of the gyrus dentatus, demonstrated strong enzyme activity both in AcP and SE preparations. The role of AcP and SE has been discussed in relation to various structures of the neocortex and hippocampal formation.

Acid Phosphatase↗

PubMed1997

Marr's theory of the neocortex as a self-organizing neural network.

Marr's proposal for the functioning of the neocortex (Marr, 1970) is the least known of his various theories for specific neural circuitries. He suggested that the neocortex learns by self-organization to extract the structure from the patterns of activity incident upon it. He proposed a feedforward neural network in which the connections to the output cells (identified with the pyramidal cells of the neocortex) are modified by a mechanism of competitive learning. It was intended that each output cell comes to be selective for the input patterns from a different class and is able to respond to new patterns from the same class that have not been seen before. The learning rule that Marr proposed was underspecified, but a logical extension of the basic idea results in a synaptic learning rule in which the total amount of synaptic strength of the connections from each input ("presynaptic") cell is kept at a constant level. In contrast, conventional competitive learning involves rules of the "postsynaptic" type. The network learns by exploiting the structure that Marr assumed to exist within the ensemble of input patterns. For this case, analysis is possible that extends that carried out by Marr, which was restricted to the binary classification task. This analysis is presented here, together with results from computer simulations of different types of competitive learning mechanisms. The presynaptic mechanism is best known in the computational neuroscience literature. In neural network applications, it may be a more suitable mechanism of competitive learning than those normally considered.

Algorithms↗