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PubMed1996

Interareal synchronization in the visual cortex.

The primary visual cortex (V1) is part of a highly interconnected network of cortical areas, hierarchically organized but operating concurrently across hierarchical levels. The high degree of reciprocal interconnection among visual cortical areas provides a framework for their interaction during the performance of visual scene analysis. The functional interdependency of visual cortical areas which develops during scene analysis can be investigated by techniques which measure interareal correlated activity. Evidence from monkeys performing a visual pattern discrimination suggests that synchronization of aperiodic activity from neuronal ensembles in cortical areas at different hierarchical levels is a relevant aspect of visual function. The near-periodic nature of the synchronized response to moving light bars in earlier studies may have been a result of the type of stimulus used. Various models of visual cortex are discussed in which interareal synchronization plays a functional role.

Animals↗
PubMed2000

The distribution of NADPH diaphorase and nitric oxide synthetase (NOS) in relation to the functional compartments of areas V1 and V2 of primate visual cortex.

The primary visual cortex (V1) of primates receives visual signals from cells in the koniocellular (K), magnocellular (M) and parvocellular (P) layers of the lateral geniculate nucleus (LGN). The functional role of the K pathway is unknown, but one proposal is that it modulates visual activity locally via release of nitric oxide (NO). One goal of this study was to examine the distribution of nitric oxide synthetase (NOS), the enzyme that produces NO, using immunocytochemistry for brain NOS (bNOS) or histochemistry for nicotinamide adenine dinucleotide phosphate (NADPH) diaphorase activity in the V1 target cells of the K pathway and within the LGN itself. A second goal was to examine bNOS and NADPH diaphorase activity within proposed functional compartments in the second visual area (V2). We examined the LGN, V1 and V2 in squirrel monkeys, owl monkeys and bushbabies. In V1 and V2, we found that dense neuropil staining for NADPH diaphorase mirrored the pattern of high metabolic activity shown with cytochrome oxidase (CO) staining but did not necessarily mirror the pattern of immunolabeling seen with antibodies against NOS. The smooth stellate cells stained for NADPH diaphorase or bNOS were sparse and did not colocalize with LGN recipient zones in V1 or with the CO compartments in V2. LGN cells projecting to V1, including K, M and P cells, were negative for bNOS and NADPH diaphorase. Therefore, high levels of NOS are not limited to the K pathway. Instead, dense NOS activity is present in interneurons and within the neuropil of V1 and V2 that exhibit high metabolic demand.

Animals↗
PubMed2001

Endogenous serotonin contributes to a developmental decrease in long-term potentiation in the rat visual cortex.

The primary visual cortex shows synaptic plasticity during a postnatal "critical period," and its plasticity declines with development. Indeed, we found a developmental decrease in the induction of long-term potentiation (LTP) in the rat visual cortex. In visual cortex slices obtained from 2- to 3-week-old rats, tetanic stimulation (100 Hz for 1 sec, twice at an interval of 30 sec) of the white matter reproducibly induced LTP of field potentials in layer II/III. However, in slices from 5-week-old rats, the same tetanic stimulation failed to induce LTP. We hypothesized that endogenous serotonin (5-HT) is responsible for the developmental decrease in visual cortex LTP, because the induction of visual cortex LTP was suppressed by the addition of exogenous 5-HT (10 microm) and because the amount of 5-HT in the visual cortex increased during development. To test this hypothesis, we investigated the effect of methysergide, a 5-HT receptor antagonist, on the induction of visual cortex LTP. When visual cortex slices from 5-week-old rats were perfused with 50 microm methysergide, tetanic stimulation of the white matter induced robust LTP in layer II/III. Furthermore, serotonergic neurons were lesioned by intracerebroventricular injection of 5,7-dihydroxytryptamine (5,7-DHT). LTP was induced in visual cortex slices from 5,7-DHT-treated, 5-week-old rats. These results suggest that the induction of visual cortex LTP in 5-week-old rats is suppressed by endogenous 5-HT. 5-HT may be a factor that determines a critical period for synaptic plasticity in the rat visual cortex.

5,7-Dihydroxytryptamine↗
PubMed1989

Nuclear terminations of corticoreticular fiber systems in rats.

Corticoreticular fiber systems were examined in adult albino and hooded rats using anterograde transport of wheat germ agglutinin-horseradish peroxidase (WGA-HRP) and anterograde degeneration. WGA-HRP injections were made stereotactically into the medial prefrontal cortex, the medial agranular cortex, the anterior cingulate cortex, the face motor cortex, the forelimb motor cortex, the trunk-hindlimb motor cortex, the face somatosensory cortex, the primary auditory cortex, the secondary visual cortex and the primary visual cortex. With exception of the cingulate cortex (which is relatively inaccessible to lesioning methods) and the primary visual cortex, electrocautery lesions were made into these same cortical areas. The precise locations of cortical injection/lesion sites were corroborated on the basis of cortical cytoarchitectonic criteria, patterns of retrograde and anterograde thalamic labeling, and patterns of anterograde labeling in non-reticular brainstem nuclei such as the red nucleus, trigeminal nuclei and dorsal column nuclei. The heaviest corticoreticular projections arise from the medial agranular cortex. The medial prefrontal cortex also gives rise to consistently strong corticoreticular projections. The anterior cingulate cortex sends robust corticoreticular projections to the upper brainstem but relatively weak projections to the lower brainstem. With respect to the primary motor cortex, the face area gives rise to the densest corticoreticular projections, rivaling those emanating from the medial agranular cortex. The trunk-hindlimb area gives rise to substantial corticoreticular projections, but those originating from the forelimb area are modest and directed chiefly to midbrain and medullary levels. The face area of the somatosensory cortex gives rise to rather weak corticoreticular projections, while those arising from the primary auditory cortex are fewer still. Descending projections from the secondary visual cortex are sparse, with labeled terminals occurring in a few pontine and medullary reticular nuclei. Only one brainstem reticular nucleus (nucleus cuneiformis) was found to receive projections from the primary visual cortex, and this input was extremely sparse. Corticoreticular projections to the upper brainstem terminate predominantly ipsilateral to the cortical injection site, whereas medullary corticoreticular projections distribute bilaterally. Corticoreticular fibers from the medial agranular, face motor and trunk-hindlimb motor cortex terminate heavily in somatomotor brainstem reticular nuclei such as the pontis oralis, the pontis caudalis and the gigantocellularis.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗
PubMed2001

MR imaging of the human brain at 1.5 T: regional variations in transverse relaxation rates in the cerebral cortex.

BACKGROUND AND PURPOSE: Heterogeneity in cortical signal intensity on T2-weighted MR images has been recently documented. Using a whole-brain, multiecho MR imaging technique, we sought to determine the T2 relaxation times of nine predefined regions in the cerebral cortex and one region in the deep gray matter. METHODS: Ten adult volunteers (nine men and one woman; age range, 18-40 y; average age, 30.8 y) underwent whole-brain imaging with an oblique coronal multiecho 3D Carr-Purcell-Meiboom-Gill MR sequence at 1000/25, 50, 75, 100, 125, and 150 (TR/TE). T2 measurements were obtained, with variably sized regions of interest, from the primary auditory cortex, primary visual cortex, caudate nucleus, superior frontal gyrus, inferior temporal gyrus, middle temporal gyrus, superior temporal gyrus, insula cortex, cingulate gyrus, and hippocampus. Repeated-measures analysis of variance was used to assess the existence of differences in T2 measurements among the anatomic locations. RESULTS: On the basis of T2 measurements, the gray matter structures examined could be divided into four statistically different groups. In ascending order of T2 measurements, the first group consisted of the primary auditory cortex and primary visual cortex; the second group, the caudate nucleus, superior frontal gyrus, inferior temporal gyrus, middle temporal gyrus, and superior temporal gyrus; the third group, the insula cortex; and the fourth group, the cingulate gyrus and hippocampus. CONCLUSION: Significant variation in T2 values among the cortical gray matter of the human brain exists.

Adolescent↗
PubMed2000

Induction of visual orientation modules in auditory cortex.

Modules of neurons sharing a common property are a basic organizational feature of mammalian sensory cortex. Primary visual cortex (V1) is characterized by orientation modules--groups of cells that share a preferred stimulus orientation--which are organized into a highly ordered orientation map. Here we show that in ferrets in which retinal projections are routed into the auditory pathway, visually responsive neurons in 'rewired' primary auditory cortex are also organized into orientation modules. The orientation tuning of neurons within these modules is comparable to the tuning of cells in V1 but the orientation map is less orderly. Horizontal connections in rewired cortex are more patchy and periodic than connections in normal auditory cortex, but less so than connections in V1. These data show that afferent activity has a profound influence on diverse components of cortical circuitry, including thalamocortical and local intracortical connections, which are involved in the generation of orientation tuning, and long-range horizontal connections, which are important in creating an orientation map.

Animals↗
PubMed1999

System-wide repercussions and adaptive plasticity: the sequelae of immature visual cortex damage.

Damage of primary visual cortex in mature mammals severely disrupts vision by disconnecting much of the cognitive processing machinery of extrastriate cortex from its source of visual signals in the retina. However, equivalent lesions incurred early in postnatal life unmask a substantial latent flexibility of the brain to minimize the disruption by specific and ordered pathway expansions that bypass the lesion. The expansions shape pathways from retina through thalamus to extrastriate cortex and onto the midbrain into new, useful forms that are retained into adult life. These useful modifications support relatively normal signal processing in a variety of structures and the sparing of certain visually guided behaviors, such as aspects of complex-pattern vision and localizing objects introduced into the visual field. Thus, both the brain and the individual are optimized, in the absence of primary visual cortex, by adaptations for useful interactions with the environment. So far, the repercussions of early visual cortex lesions have been most thoroughly documented in cats, although it is likely on the basis of known repercussions and similarity of visual system organization and developmental sequence, that broadly equivalent repercussions and adaptations occur in monkeys and humans following early lesions of primary visual cortex. The knowledge gained has implications for devising therapeutic strategies to attenuate defects in vision induced by cortical lesions.

Journal Article↗
PubMed1994

System-wide repercussions of damage to the immature visual cortex.

Damage of the primary visual cortex in mammals, including humans, severely disrupts vision by disconnecting much of the cognitive-processing machinery of extrastriate cortex from its source of visual signals in the retina. Studies of the anatomical consequences of damage to the immature primary visual cortex in cats reveal system-wide repercussions on neural circuitry that includes the retina, thalamus, midbrain and extrastriate cortex. The repercussions modify circuits that support relatively normal signal processing and the sparing of certain visually guided behaviors such as aspects of complex-pattern recognition and orienting to novel stimuli introduced into the visual field. These studies have implications for understanding the consequences of damage to the visual cortex in infant monkeys and humans, and for devising therapeutic strategies to attenuate defects in vision induced by cortical lesions.

Animals↗
PubMed2002

Human regional cerebral glucose metabolism during non-rapid eye movement sleep in relation to waking.

Sleep is an essential human function. Although the function of sleep has generally been regarded to be restorative, recent data indicate that it also plays an important role in cognition. The neurobiology of human sleep is most effectively analysed with functional imaging, and PET studies have contributed substantially to our understanding of both rapid eye movement (REM) and non-rapid eye movement (NREM) sleep. In this study, PET was used to determine patterns of regional glucose metabolism in NREM sleep compared with waking. We hypothesized that brain structures related to waking cognitive function would show a persistence of function into the NREM sleep state. Fourteen healthy subjects (age range 21-49 years; 10 women, 4 men) underwent concurrent EEG sleep studies and [(18)F]fluoro-2-deoxy-D-glucose PET scans during waking and NREM sleep. Whole-brain glucose metabolism declined significantly from waking to NREM sleep. Relative decreases in regional metabolism from waking to NREM sleep occurred in wide areas of frontal, parietal, temporal and occipital association cortex, primary visual cortex, and in anterior/dorsomedial thalamus. After controlling for the whole-brain declines in absolute metabolism, relative increases in regional metabolism from waking to NREM were found bilaterally in the dorsal pontine tegmentum, hypothalamus, basal forebrain, ventral striatum, anterior cingulate cortex and extensive regions of the mesial temporal lobe, including the amygdala and hippocampus, and in the right dorsal parietal association cortex and primary somatosensory and motor cortices. The reductions in relative metabolism in NREM sleep compared with waking are consistent with prior findings from blood flow studies. The relative increases in glucose utilization in the basal forebrain, hypothalamus, ventral striatum, amygdala, hippocampus and pontine reticular formation are new observations that are in accordance with the view that NREM sleep is important to brain plasticity in homeostatic regulation and mnemonic processing.

Adult↗
PubMed1997

Dopamine receptor transcript expression in striatum and prefrontal and occipital cortex. Focal abnormalities in orbitofrontal cortex in schizophrenia.

BACKGROUND: The identification of novel subtypes of the dopamine receptors has renewed interest in the involvement of dopaminergic mechanisms in schizophrenia. We determined the expression of transcripts encoding the dopamine receptors in the brains of schizophrenic patients. METHODS: The levels of the messenger RNA molecules encoding the 5 dopamine receptors were quantified in postmortem brain samples from 16 schizophrenic patients and 9 control subjects. Samples from multiple regions of the prefrontal cortex, primary visual cortex, and striatum were subjected to in situ hybridization followed by quantitative image analysis. RESULTS: Expression of dopamine receptor transcripts did not differ between schizophrenic patients and controls in striatum or visual cortex. Dramatic decreases of dopamine receptor transcripts were found in the prefrontal cortex, but these changes were restricted to the D3 and D4 receptors, and localized to Brodmann area 11 (orbitofrontal cortex). CONCLUSIONS: Cortical dopaminergic neurotransmission may be disrupted in schizophrenia at the level of receptor expression. There appears to be a focal abnormality of D3 and D4 messenger RNA expression in the prefrontal cortex, with down-regulation of both, consistent with prefrontal cortical hypodopaminergia in schizophrenia.

Aged↗
PubMed2001

Face processing occurs outside the fusiform 'face area' in autism: evidence from functional MRI.

Processing the human face is at the focal point of most social interactions, yet this simple perceptual task is difficult for individuals with autism, a population that spends limited amounts of time engaged in face-to-face eye contact or social interactions in general. Thus, the study of face processing in autism is not only important because it may be integral to understanding the social deficits of this disorder, but also, because it provides a unique opportunity to study experiential factors related to the functional specialization of normal face processing. In short, autism may be one of the only disorders where affected individuals spend reduced amounts of time engaged in face processing from birth. Using functional MRI, haemodynamic responses during a face perception task were compared between adults with autism and normal control subjects. Four regions of interest (ROIs), the fusiform gyrus (FG), inferior temporal gyrus, middle temporal gyrus and amygdala were manually traced on non-spatially normalized images and the percentage ROI active was calculated for each subject. Analyses in Talairach space were also performed. Overall results revealed either abnormally weak or no activation in FG in autistic patients, as well as significantly reduced activation in the inferior occipital gyrus, superior temporal sulcus and amygdala. Anatomical abnormalities, in contrast, were present only in the amygdala in autistic patients, whose mean volume was significantly reduced as compared with normals. Reaction time and accuracy measures were not different between groups. Thus, while autistic subjects could perform the face perception task, none of the regions supporting face processing in normals were found to be significantly active in the autistic subjects. Instead, in every autistic patient, faces maximally activated aberrant and individual-specific neural sites (e.g. frontal cortex, primary visual cortex, etc.), which was in contrast to the 100% consistency of maximal activation within the traditional fusiform face area (FFA) for every normal subject. It appears that, as compared with normal individuals, autistic individuals 'see' faces utilizing different neural systems, with each patient doing so via a unique neural circuitry. Such a pattern of individual-specific, scattered activation seen in autistic patients in contrast to the highly consistent FG activation seen in normals, suggests that experiential factors do indeed play a role in the normal development of the FFA.

Adult↗
PubMed2003

Neuronal classes in the isocortex of a monotreme, the Australian echidna (Tachyglossus aculeatus).

We have used Valverde-Golgi and Golgi-Colonnier techniques to analyze cortical neuronal morphology in four regions (frontal cortex, primary motor cortex, primary somatosensory cortex, primary visual cortex) of the isocortex of the echidna (Tachyglossus aculeatus). Eight classes of neurons could be identified--pyramidal, spinous bipolar, aspinous bipolar, spinous bitufted, aspinous bitufted, spinous multipolar, aspinous multipolar and neurogliaform. All except the pyramidal neurons were morphologically similar to neuronal classes seen in eutherian and metatherian isocortex. Pyramidal neurons made up a small proportion of all cortical neurons encountered in our preparations of echidna cortex (34% in visual cortex, 35% in somatosensory cortex, 41% in frontal cortex and 49% in motor cortex) compared to both reported values in eutherian cortex and values we found in rat cortex impregnations prepared in an identical fashion to the echidna material (75% in rat motor and 78% in rat somatosensory cortex). Many pyramidal neurons in the echidna isocortex were atypical (30-42% depending on region) with inverted somata, short or branching apical dendrites and/or few basal dendrites, very different from the usual pyramidal neuron morphology in eutherian cortex. Dendritic spine density on apical and basal dendrites of echidna pyramidal neurons in somatosensory cortex and apical dendrites of motor cortex pyramidal neurons was also lower than that found in the rat. The present findings are consistent with both pyramidal neurons and the many diverse types of non-pyramidal neurons having already emerged as discrete morphological entities very early in mammalian cortical evolution, at the time of divergence of the therian and prototherian lineage.

Animals↗
PubMed1999

Organization of intracortical circuits in relation to direction preference maps in ferret visual cortex.

Neurons in the primary visual cortex are selective for the direction of movement of a visual stimulus. Like other stimulus features, direction preference is mapped on the cortical surface in a systematic manner. Intracortical synaptic circuits, in particular inhibitory connections, have been implicated in the emergence of direction selectivity. Whether intracortical inhibition specifically suppresses responses to the nonpreferred direction or has a nonspecific "thresholding" effect is still controversial. To address these questions we investigated the relationship between patterns of intracortical synaptic connections and direction domains in ferret primary visual cortex (area 17) using a combined in vivo-in vitro approach. Excitatory synaptic inputs were iso-direction-tuned. The majority of local inhibitory inputs were also iso-direction-tuned. However, approximately 40% of inhibitory connections originated in regions preferring the opposite direction. These findings indicate that specific inhibitory interactions between cortical regions of opposite direction preference may contribute to the emergence and sharpening of direction selectivity.

Animals↗
PubMed1991

Relation of cortical cell orientation selectivity to alignment of receptive fields of the geniculocortical afferents that arborize within a single orientation column in ferret visual cortex.

Neurons in the primary visual cortex of higher mammals are arranged in columns, and the neurons in each column respond best to light-dark borders of particular orientations. The basis of cortical cell orientation selectivity is not known. One possible mechanism would be for cortical cells to receive input from several lateral geniculate nucleus (LGN) neurons with receptive fields that are aligned in the visual field (Hubel and Wiesel, 1962). We have investigated the relationship between the arrangement of the receptive fields of geniculocortical afferents and the orientation preferences of cortical cells in the orientation columns to which the afferents provide visual input. Radial microelectrode penetrations were made into primary visual cortex of anesthetized adult sable ferrets. Cortical cells were recorded throughout the depth of the cortex, and their orientation preferences were determined. Cortical cell responses were then eliminated by superfusion of the cortex with either kainic acid (Zahs and Stryker, 1988) or muscimol. After the drug treatment, responses from many single units with distinct receptive fields were recorded. These responses were presumed to be those of geniculocortical afferents, because they had the response properties characteristic of LGN neurons, and because they could be recorded only in cortical layers that receive geniculate input. In 16 of 18 cases, the afferent receptive fields recorded in a single penetration covered an elongated region of visual space. In these penetrations, the best-fit line through the centers of the afferent receptive fields generally paralleled the preferred orientation of cortical cells recorded at the same site in cortex. These results are consistent with the Hubel and Wiesel (1962) model for the construction of oriented visual cortical receptive fields from geniculate inputs with aligned receptive fields.

Afferent Pathways↗
PubMed1994

Differential localization of two glutamic acid decarboxylases (GAD65 and GAD67) in adult monkey visual cortex.

Adult monkey primary visual cortex contains a diverse population of stellate neurons that utilize the neurotransmitter gamma aminobutyric acid (GABA). Two glutamic acid decarboxylase (GAD) enzymes that synthesize GABA, GAD65 and GAD67, were localized within these stellate neurons by in situ hybridization of 35S or digoxigenin (DIG) labeled riboprobes. Double labels were done by using 35S GAD67 riboprobe and GABA immunocytochemistry on the same section to verify that the neuronal population identified by immunocytochemistry was the same one studied in the in situ hybridization experiments. We find that GAD65 mRNA and GAD67 mRNA are widely distributed in the cortex, with four bands of heavily labeled neurons in upper layer 2, lower 3, 4C, and 6. GAD67 labeled neurons were more obvious in layer 4C beta, while GAD65 containing neurons were common in layer 1 and white matter. Northern blots and in situ hybridization on sections with both 35S and DIG riboprobes indicate that cortical neurons typically contain more GAD67 mRNA. Cell counts show that 18% of all cortical neurons contain GAD67 mRNA and 13% contain GAD65 mRNA, suggesting that a small population of GABA neurons might lack GAD65. Cell bodies that contain high amounts of GAD65 mRNA are prominent in layers deep 3, 4B, 4C alpha, and 6 and often are the largest cells in their respective layers. Double labels demonstrate that 96% of all GABA+ neurons contain GAD67 mRNA. Neurons heavily labeled for GABA tend to have smaller cell bodies and contain less GAD67 mRNA, while lightly labeled GABA neurons are larger and contain more GAD67 mRNA. These data indicate that most GABA neurons in monkey striate cortex contain both GAD enzymes. Although the differences in GABA content, cell size, laminar distribution, and GAD mRNA concentration suggest different requirements for GAD67 and GAD65 in cortical circuits, our experiments do not reveal what different roles these two enzymes subserve within GABAergic stellate neurons.

Animals↗
PubMed2003

Limit of spared pattern vision following lesions of the immature visual cortex.

Lesions of primary visual cortex sustained early in life spare certain aspects of visual processing that can be linked to expansions of bypass pathways to extrastriate cortex. They also trigger, in an age-dependent way, partial or complete transneuronal retrograde degeneration of beta (X) retinal ganglion cells, which are implicated in visual processing under conditions of low contrast. We used two-dimensional geometric patterns whose saliency was reduced by gradually increasing levels of superimposed masking lines, and by reductions in spatial contrast. Normative data were collected from intact cats, and baseline lesion data were collected from cats with lesions sustained as young adults (postnatal day 180, P180). Experimental data were collected from cats that sustained lesions on P1-3 or P26-30. For high contrast patterns, the adult group was impaired at both acquisition (sequential progressive levels of masking) and concurrent (parallel high and low levels of masking) performance, whereas the early-lesioned groups were impaired only at concurrent performance. All lesion groups were equally impaired when contrast was reduced to modest or lower levels. These results show that sparing of masked-pattern learning is limited to the high end of the spatial contrast domain.

Animals↗
PubMed2002

Suppression without inhibition in visual cortex.

Neurons in primary visual cortex (V1) are thought to receive inhibition from other V1 neurons selective for a variety of orientations. Evidence for this inhibition is commonly found in cross-orientation suppression: responses of a V1 neuron to optimally oriented bars are suppressed by superimposed mask bars of different orientation. We show, however, that suppression is unlikely to result from intracortical inhibition. First, suppression can be obtained with masks drifting too rapidly to elicit much of a response in cortex. Second, suppression is immune to hyperpolarization (through visual adaptation) of cortical neurons responding to the mask. Signals mediating suppression might originate in thalamus, rather than in cortex. Thalamic neurons exhibit some suppression; additional suppression might arise from depression at thalamocortical synapses. The mechanisms of suppression are subcortical and possibly include the very first synapse into cortex.

Action Potentials↗
PubMed2002

The role of feedback in shaping neural representations in cat visual cortex.

In the primary visual cortex, neurons with similar response preferences are grouped into domains forming continuous maps of stimulus orientation and direction of movement. These properties are widely believed to result from the combination of ascending and lateral interactions in the visual system. We have tested this view by examining the influence of deactivating feedback signals descending from the visuoparietal cortex on the emergence of these response properties and representations in cat area 18. We thermally deactivated the dominant motion-processing region of the visuoparietal cortex and used optical and electrophysiological methods to assay neural activity evoked in area 18 by stimulation with moving gratings and fields of coherently moving randomly distributed dots. Feedback deactivation decreased signal strength in both orientation and direction maps and virtually abolished the global layout of direction maps, whereas the basic structure of the orientation maps was preserved. These findings could be accounted for by a selective silencing of highly direction-selective neurons and by the redirection of preferences of less selective neurons. Our data suggest that signals fed back from the visuoparietal cortex strongly contribute to the emergence of direction selectivity in early visual areas. Thus we propose that higher cortical areas have significant influence over fundamental neuronal properties as they emerge in lower areas.

Animals↗