On the computational architecture of the neocortex

This paper is a sequel to an earlier paper which proposed an active role for the thalamus, integrating multiple hypotheses formed in the cortex via the thalamo-cortical loop. In this paper, I put forward a hypothesis on the role of the reciprocal, topographic pathways between two cortical areas, one often a higher area dealing with more abstract information about the world, the other lower, dealing with more concrete data. The higher area attempts to fit its abstractions to the data it receives from lower areas by sending back to them from its deep pyramidal cells a template reconstruction best fitting the lower level view. The lower area attempts to reconcile the reconstruction of its view that it receives from higher areas with what it knows, sending back from its superficial pyramidal cells the features in its data which are not predicted by the higher area. The whole calculation is done with all areas working simultaneously, but with order imposed by synchronous activity in the various top-down, bottom-up loops. Evidence for this theory is reviewed and experimental tests are proposed. A third part of this paper will deal with extensions of these ideas to the frontal lobe.     

Comparison of the Connectional Properties of the Two Forelimb Areas of the Rat Sensorimotor Cortex: Support for the Presence of a Premotor or Supplementary Motor Cortical Area

The existence of multiple motor cortical areas that differ in some of their properties is well known in primates, but is less clear in the rat. The present study addressed this question from the point of view of connectional properties by comparing the afferent and efferent projections of the caudal forelimb area (CFA), considered to be the equivalent of the forelimb area of the primary motor cortex (MI), and a second forelimb motor representation, the rostral forelimb area (RFA). As a result of various tracing experiments (including double labeling), it was observed that CFA and RFA had reciprocal corticocortical connections characterized by preferential, asymmetrical, laminar distribution, indicating that RFA may occupy a different hierarchical level than CFA, according to criteria previously discussed in the visual cortex of primates. Furthermore, it was found that RFA, but not CFA, exhibited dense reciprocal connections with the insular cortex. With respect to their efferent projection to the basal ganglia, it was observed that CFA projected very densely to the lateral portion of the ipsilateral caudate putamen, whereas the contralateral projection was sparse and more restricted. The ipsilateral projection originating from RFA was slightly less dense than that from CFA, but it covered a larger portion of the caudate putamen (in the medial direction); the contralateral projection from RFA to the caudate putamen was of the same density and extent as the ipsilateral projection. The reciprocal thalamocortical and corticothalamic connections of RFA and CFA differed from each other in the sense that CFA was mainly interconnected with the ventrolateral thalamic nucleus, while RFA was mainly connected with the ventromedial thalamic nucleus. Altogether, these connectional differences, compared with the pattern of organization of the motor cortical areas in primates, suggest that RFA in the rat may well be an equivalent of the premotor or supplementary motor area. In contrast to the corticocortical, corticostriatal, and thalamocortical connections, RFA and CFA showed similar efferent projections to the subthalamic nucleus, substantia nigra, red nucleus, tectum, pontine nuclei, inferior olive, and spinal cord.     

Change in the functional significance of structures of the cat brain as a result of reorganization of its activity

A study was carried out on 8 adult cats of functional role of the frontal, parietal and occipital parts of the neocortex, and also of the dorsal hippocampus, mediodorsal thalamic nucleus and caudate nucleus head, in realization of a delayed spatial choice (DSCh) before and after compensatory reorganizations of the brain activity caused by multiple electrical stimulation of the frontal part of the cerebral cortex. Compensatory reorganization led to a change of functional significance of these structures. While before this change the frontal cortex, hippocampus and mediodorsal thalamic nucleus were critically necessary brain areas for the realization of the DSCh, after it parietal and occipital cortical areas acquired such significance. The obtained data are discussed proceeding from the principle of the integrity in the brain activity.     

Pharmacological evaluation of GABAergic and glutamatergic inputs to the nucleus basalis--cortical and the septal-hippocampal cholinergic projections

The presence of different receptor populations within a given brain area can be effectively evaluated via the local injections of defined receptor agonists and antagonists. Using this approach, it has become evident that the nucleus basalis - cortical cholinergic pathway possesses an inhibitory GABAergic input to the nucleus basalis from the nucleus accumbens as well as a positive glutamatergic feedback from the cortex. The septal-hippocampal cholinergic pathway also possesses an inhibitory GABAergic regulation which consists of a large GABAergic interneuron population in the septum. A glutamatergic feedback from the hippocampus is also present. These regulatory inputs to cholinergic cells are not tonically active but appear to function as phasic modulators of cholinergic transmission in both pathways.     

Properties of the pyramidal tract neuron system within the precentral wrist and hand area of primate motor cortex.

1. To obtain basic anatomical data that will be useful in interpreting the results of studies of primate pyramidal tract neurons (PTNs), extracellular, single-unit recording techniques were used to determine a number of the properties of the PTN population within the electrically defined, precentral wrist zone of the monkey's motor cortex. 2. Recordings were obtained from a total of 1,375 antidromically identified PT and corticospinal tract (CST) cells. A mathematical model was then used to correct the statistics of the sample for variations in the probability of unit detection, which arise from variations in neuronal size and extracellular field dimensions. 3. Both the experimentally observed and theoretically corrected results suggest that the PT projection from this cortical zone is derived principally from slowly conducting, and presumably small to medium-sized cells (an estimated 85% of the resident PTN population). 4. Both the fast and slow cell subpopulations were found to be concentrated within cortical layer V, where they tend to congregate in small, mixed clusters of 2 to 5 neurons. Estimates of the total packing density of PTNs within layer V of this cortical zone suggest that they account for only 10-20% of the neurons within this major efferent layer. 5. 70% of the slow and 82% of the fast PT neurons within this cortical area were found to send their axons into the contralateral, lateral corticospinal tract. Thus, in futur functional studies of PTNs in this cortical area, it can be assumed that three of every four neurons will in fact influence segmental cells of one category or another directly. 6. Extensive data are also presented on the incidence of axon collateral branching from PT and CST cells to the red nucleus, the medial medullary reticular formation and the cuneate nucleus. 7. Some general implications of these findings for the design of future functional studies of anatomically identified motor cortex cell systems are then discussed.     

Connections between the anterior thalamic nuclei in turtles (data of the peroxidase method)

It turtles, Testudo horsfieldi (Gray) connections of anterior dorsomedial and dorsolateral thalamic nuclei have been investigated by means of horseradish peroxidase, injected ionophoretically. Retrogradely labelled neurons are predominantly revealed ipsilaterally in the cerebral structures belonging to the limbic system: in the forebrain--basal parts of the hemisphere, septum, adjoining nucleus, nuclei of the anterior and hippocampal commissures, hippocampal cortex, preoptic area; in the diencephalon--in the subthalamus (suprapeduncular nucleus), in some hypothalamic structures (para- and periventricular nuclei, posterior nucleus, lateral hypothalamic area, mamillary complex); in the brain stem--ventral tegmental area, superior nucleus of the suture. Less vast connections are with nonlimbic cerebral formations: projections to the striatum, afferents from the laminar nucleus of the acoustic torus, nuclei of the posterior commissure. Similarity and difference of the nuclei investigated in the turtles with the thalamic anterior nuclei in lizards, with the anterior and intralaminar nuclei in Mammalia are discussed. An idea is suggested on functional heterogeneity of the anterior nuclei in reptiles and on their role for ensuring limbic functions at the thalamic level.     

Unit responses in area 5b of the suprasylvian gyrus to stimulation of the posterior lateral thalmamic nucleus

In response to stimulation of the posterior lateral nucleus in unanesthetized cats immobilized with D-tubocurarine an evoked potential consisting of three components with a latent period of 3–5 msec appeared in area 5b of the suprasylvian gyrus. All three components were reversed at about the same depth in the cortex (1500–1600 µ). Reversal of the potential shows that it is generated in that area by neurons evidently located in deeper layers of the cortex and is not conducted to it physically from other regions. Responses of 53 spontaneously active neurons in the same area of the cortex to stimulation of the posterior lateral nucleus were investigated. A characteristic feature of these reponses was that inhibition occurred nearly all of them. In 22 neurons the responses began with inhibition, which lasted from 30 to 400 msec. In 30 neurons inhibition appeared immediately after excitation while one neuron responded by excitation alone. The latent periods of the excitatory responses varied from 3 to 28 msec. The short latent period of the evoked potentials and of some single units responses (3–6 msec) confirms morphological evidence of direct connections between the posterior lateral nucleus and area 5b of the suprasylvian gyrus. Repetitive stimulation of that nucleus led to strengthening of both excitation and inhibition. Influences of the posterior lateral nucleus were opposite to those of the specific nuclei: the posterior ventrolateral nucleus and the lateral and medial geniculate bodies. Stimulation of the nonspecific reticular nucleus, however, evoked discharges from neurons like those produced by stimulation of the posterior lateral nucleus.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 5, pp. 502–509, September–October, 1973.     

Coding of the reach vector in parietal area 5d

Competing models of sensorimotor computation predict different topological constraints in the brain. Some models propose population coding of particular reference frames in anatomically distinct nodes, whereas others require no such dedicated subpopulations and instead predict that regions will simultaneously code in multiple, intermediate, reference frames. Current empirical evidence is conflicting, partly due to difficulties involved in identifying underlying reference frames. Here, we independently varied the locations of hand, gaze, and target over many positions while recording from the dorsal aspect of parietal area 5. We find that the target is?represented in a predominantly hand-centered reference frame here, contrasting with the relative code seen in dorsal premotor cortex and the mostly gaze-centered reference frame in the parietal reach region. This supports the hypothesis that different nodes of the sensorimotor circuit contain distinct and systematic representations, and this constrains the types of computational model that are neurobiologically relevant.     

Study of the ancient tecto-thalamo-cortical pathway of the visual system in rats

Recording the evoked potentials and neuronal activity, electrophysiological studies have been made on tecto-thalamo-cortical tract in rats. The existence of a system of efferent projections in the superficial, visual layers of the superior colliculi was shown which are diffusely present in the nucleus lateralis posterior (n. LP), indicating low level of morpho-functional organization of this region of the dorsal thalamus in rats. In response to electrical stimulation of the n. LP, in laterocaudal parts of the visual system (fields 17 and 18a of the cortex) the evoked potentials of primary-negative polarity were observed which are associated mainly with the superficial (I--IV) cortical layers. Predominant representation of tecto-thalamo-cortical system in the laterocaudal visual area of the cortex indicates the tendency to separate representation (with respect to cortical areas and cortical layers) of retino-geniculate and retino-tecal visual systems in rats.     

Diagnosis of complex coastal ecological systems: Environmental GIS analysis of a highly stressed Mediterranean lagoon through spatiotemporal indicators

This article explores the need for implementing spatiotemporal criteria in environmental planning of special protected areas, applying the analysis of reaction patterns in the case study of the Mar Menor, a Mediterranean coastal lagoon in South-Eastern Spain. To perform this, the Mar Menor environmental patterns suffering traditionally from intense coastal urbanization, and in recent years from important eutrophication problems, will be analyzed from an integrated approach and retrospectively assessed through territorial indicators. The evaluation will provide a clear example of a protected area regulated by multiple European Natura 2000 figures of protection that nevertheless is suffering from a strong and varied anthropic process over time. The results will show that paradoxically this process sometimes has little territorial linkage origin with the area concerned. This aspect will be addressed by using GIS indicators, to present the real correlation between coastal urbanization and agricultural land transformation with Mar Menor evolution over time (in which it is pointed out that the weight of the latter is much greater than the former) and how far it is necessary to establish an area of influence of the environmental impacts by proposing a new scope of application for the resolution of the current situation of eutrophication in its waters.     

大鼠电惊厥后c－fos在中枢神经系统中的表达

本实验应用Ｆｏｓ蛋白免疫组织化方法对大鼠电惊厥后不同时程中枢神经系统内的ｃ－ｆｏｓ表达进行了观察,结果表明;①ｃ－ｆｏｓ表达于电惊厥后３０ｍｉｎ开始,２ｈ达高峰,４ｈ后逐渐下降,１２ｈ基本恢复正常。②Ｆｏｓ阳性神经元呈对称性分布,分布于颞叶听皮质、梨状皮质、内嗅皮质、ｓｔｒ１８区、杏仁体、海马、齿状回、斜角带核、弓状核、下丘脑腹内侧核、．视前区、丘脑、上丘灰质、中央灰质和脑桥腹侧部等结构。③ｃ－ｆｏｓ在颞叶听皮质、梨状皮质和内嗅皮质最先表达,表达程度最强,恢复也最慢,提示这些结构可能是电惊厥发生的起源结构,并呈现皮层结构向皮层下结构播散的过程,主要播及边缘结构。     

The role of the cell nucleus in mechanotransduction

Mechanical forces are known to influence cellular processes with consequences at the cellular and physiological level. The cell nucleus is the largest and stiffest organelle, and it is connected to the cytoskeleton for proper cellular function. The connection between the nucleus and the cytoskeleton is in most cases mediated by the linker of nucleoskeleton and cytoskeleton (LINC) complex. Not surprisingly, the nucleus and the associated cytoskeleton are implicated in multiple mechanotransduction pathways important for cellular activities. Herein, we review recent advances describing how the LINC complex, the nuclear lamina, and nuclear pore complexes are involved in nuclear mechanotransduction. We will also discuss how the perinuclear actin cytoskeleton is important for the regulation of nuclear mechanotransduction. Additionally, we discuss the relevance of nuclear mechanotransduction for cell migration, development, and how nuclear mechanotransduction impairment leads to multiple disorders.     

Cell death and the creation of regional differences in neuronal numbers.

Regional variations in cell death are ubiquitous in the nervous system. In the retina, cell death in retinal ganglion cells is elevated in the retinal periphery and may be important in setting up the initial conditions that produce central retinal specializations such as an area centralis or visual streak. In central visual system structures, pronounced spatial and spatiotemporal inhomogeneities in cell death are seen both in layers and regions of the lateral geniculate nucleus and superior colliculus; similar indications of inhomogeneities are seen in those nonvisual structures that have been examined. Cell death in the cortex is highly nonuniform, by layer and by cortical area. A variety of possible functions for these regional losses are proposed, in the context of a uniform mechanism for cell death that allows it to assume multiple functions.     

Electrophysiological analysis of corticofugal connections with the medial dorsal thalamic nucleus

In acute experiments on cats anesthetized with pentobarbital and chloralose, focal responses were recorded to study projections of various parts of the orbitofrontal cortex and cortex of the temporal pole in the region of the medial dorsal nucleus of the thalamus and interaction in this nucleus between stimuli arriving from the medio-basal portions of the neocortex. Different parts of the orbitofrontal cortex were found to have local projections in the medial dorsal nucleus so arranged that the rostral zones of the cortex send stimuli to the medio-dorsal portions of the nucleus, whereas regions of the cortex radiating fanwise from the pole in dorsal and caudal directions are arranged in the lateral and basal portions of the nucleus. The cortex of the temporal pole has relatively diffuse projections in the medial part of the medial dorsal nucleus. Stimuli reaching the medial dorsal nucleus from the basal structures of the neocortex (temporal pole) were shown to facilitate response to stimulation of the orbitofrontal cortex. Meanwhile, stimulation of this region of the cortex depresses the receptive capacity of the nucleus for impulses arriving from the temporal cortex.     

Emergence in the central nervous system

“Emergence” is an idea that has received much attention in consciousness literature, but it is difficult to find characterizations of that concept which are both specific and useful. I will precisely define and characterize a type of epistemic (“weak”) emergence and show that it is a property of some neural circuits throughout the CNS, on micro-, meso- and macroscopic levels. I will argue that possession of this property can result in profoundly altered neural dynamics on multiple levels in cortex and other systems. I will first describe emergent neural entities (ENEs) abstractly. I will then show how ENEs function specifically and concretely, and demonstrate some implications of this type of emergence for the CNS.     

The distribution of cholinergic neurons in the human central nervous system

Choline acetyltransferase (ChAT), the enzyme responsible for the biosynthesis of acetylcholine, is presently the most specific marker for identifying cholinergic neurons in the central and peripheral nervous systems. The present article reviews immunohistochemical and in situ hybridization studies on the distribution of neurons expressing ChAT in the human central nervous system. Neurons with both immunoreactivity and in situ hybridization signals of ChAT are observed in the basal forebrain (diagonal band of Broca and nucleus basalis of Meynert), striatum (caudate nucleus, putamen and nucleus accumbens), cerebral cortex, mesopontine tegmental nuclei (pedunculopontine tegmental nucleus, laterodorsal tegmental nucleus and parabigeminal nucleus), cranial motor nuclei and spinal motor neurons. The cerebral cortex displays regional and laminal differences in the distribution of neurons with ChAT. The medial septal nucleus and medial habenular nucleus contain immunoreactive neurons for ChAT, which are devoid of ChAT mRNA signals. This is probably because there is a small number of cholinergic neurons with a low level of ChAT gene expression in these nuclei of human. Possible connections and speculated functions of these neurons are briefly summarized.     

Thoughts on the cerebral cortex

The cortex is often described as a network processing information in the direction from sensory to motor areas. However, the structure of the cortex is asymmetrical only in the vertical direction, suggesting an input-output transformation between layers rather than between areas. This operation must be a very generally applicable one, since the plan of the cortex is basically the same everywhere. In an attempt to understand it, a skeleton cortex of only pyramidal cells is considered. They are characterized by a double dendritic expansion, an apical one in the first layer, which is considered as the input layer, and a basal one which receives excitation from the axon collaterals of other pyramidal cells. If pyramidal cells learn (perhaps by growing dendritic spines) to respond to frequent constellations of activity in their afferents, each will learn a property of the input (through its apical dendrites) provided that it was preceded by other properties sensed by neighbouring pyramidal cells (which influences it through its basal dendrites). Thus the pyramidal cells will code the input in terms of properties which have a tendency to follow each other. This will be a coding which reflects the causal structure of the world. Various uses of a network embodying the conditional probabilities of events in the input are described, including recognition of familiar sequences and prediction. The local variation of fiber patterns in the cerebral cortex of man, described as myeloarchitectonics, is interpreted as a macroscopical expression of the different statistics of the set of conditional probabilities linking the events represented by individual pyramidal cells in different areas (in different functional contexts).     

An examination of somatosensory area SIII in ferret cortex

A somatotopically organized region on the suprasylvian gyrus of the ferret was examined using multiunit recordings and anatomical tracer injections. This area, which contains a representation of the face, was bordered by the primary somatosensory area (SI), anteriorly, and by the visually responsive rostral posterior parietal cortex (PPr), posteriorly. Anatomical tracers revealed connections to this region from cortical areas MI, SI, MRSS, PPr, and the thalamic posterior nucleus. These results are consistent with previous work in ferrets as well as with the location, physiology, and connectivity of area SIII in cats. Given its associations, functional properties, location, and homology, it is proposed that this region represents the third cortical somatosensory area (SIII) in ferrets.