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PubMed1986

The brain as a self-organizing system.

Clinical evidence and numerous results from animal experimentation indicate that cognitive functions have to be learned. Brain structures subserving these functions require sensory experience for their maturation. Genetic instructions are in principle not sufficient to specify neuronal connections with sufficient precision. Self-organization processes are implemented in addition which allow to optimize genetically determined blue prints of connectivity by making use of functional criteria. Thus, neuronal activity becomes an important shaping factor in the development of the structural and functional architecture of the forebrain. To the extent that this neuronal activity is modulated by sensory signals, environmental factors can influence the development of neuronal networks. Recent experiments indicate that these shaping processes are additionally controlled by modulatory systems. Both, the noradrenergic projection from the locus coeruleus and the cholinergic projection from the basal forebrain facilitate activity-dependent long-term changes of neuronal connections during development. The activity of these modulatory systems in turn depends on central states such as arousal, attention, and perhaps also motivation. It is inferred from this evidence that experience-dependent self-organization should not be considered as a passive imprinting process but rather as an active dialogue between the brain and its environment. The hypothesis is discussed that many developmental disturbances which are commonly attributed to deprivation are in fact due to defaults of the CNS which either lead to the formulation of wrong questions or to the reduction of exploratory drive.

Animals↗
PubMed2003

The recurrent mossy fiber pathway of the epileptic brain.

The dentate gyrus is believed to play a key role in the pathogenesis of temporal lobe epilepsy. In normal brain the dentate granule cells serve as a high-resistance gate or filter, inhibiting the propagation of seizures from the entorhinal cortex to the hippocampus. The filtering function of the dentate gyrus depends in part on the near absence of monosynaptic connections among granule cells. In humans with temporal lobe epilepsy and in animal models of temporal lobe epilepsy, dentate granule cells form an interconnected synaptic network associated with loss of hilar interneurons. This recurrent mossy fiber pathway mediates reverberating excitation that can reduce the threshold for granule cell synchronization. Factors that augment activity in this pathway include modest increases in [K+]o; loss of GABA inhibition; short-term, frequency-dependent facilitation (frequencies of 1-2 Hz); feedback activation of kainate autoreceptors; and release of zinc from recurrent mossy fiber boutons. Factors that diminish activity include short-term, frequency-dependent depression (frequencies < 1 Hz); feedback activation of type II metabotropic glutamate receptors; and the potential release of GABA, neuropeptide Y, adenosine, and dynorphin from recurrent mossy fiber boutons. The axon sprouting and reactive synaptogenesis that follow seizure-related brain damage can also create or strengthen recurrent excitation in other brain regions. These changes are expected to facilitate participation of these regions in seizures. Thus, reactive processes that are often considered important for recovery of function after most brain injuries probably contribute to neurological dysfunction in epilepsy.

Animals↗
PubMed2004

Effects of clonidine injections into the bed nucleus of the stria terminalis on fear and anxiety behavior in rats.

Emotions such as fear and anxiety are mediated by a neural network containing nuclei like the amygdala, the bed nucleus of the stria terminalis and the periaqueductal gray. Noradrenaline is a neurotransmitter closely connected with the processing of stimuli eliciting these emotions. The bed nucleus of the stria terminalis contains the highest density of noradrenaline within the brain. In the present study, we investigated effects of injections of the noradrenergic alpha2-adrenoceptor agonist clonidine into the bed nucleus of the stria terminalis on learned and unlearned fear (anxiety) in rats on different animal models of fear and anxiety: acquisition and expression of fear-potentiated startle, sensitization of the acoustic startle response by foot shocks and light-enhanced startle. Clonidine injections disrupted acquisition and expression of fear-potentiated startle, as well as light-enhanced startle, whereas sensitization was not affected. These results indicate that noradrenaline within the bed nucleus of the stria terminalis mediates both fear and anxiety. We suggest that there is rather a neurochemical than a neuroanatomical dissociation between learned fear and anxiety as hypothesized by Walker and Davis (Walker, D.L. and M. Davis, 1997b, Double dissociation between the involvement of the bed nucleus of the stria terminalis and the central nucleus of the amygdala in startle increases produced by conditioned versus unconditioned fear, J. Neurosci. 17, 9375-9383.).

Animals↗
PubMed2006

Impaired feedforward inhibition of the thalamocortical projection in epileptic Ca2+ channel mutant mice, tottering.

The tottering (tg) mice have a mutation in the CaV2.1 (P/Q-type) voltage-dependent Ca2+ channel alpha(1)2.1 subunit gene. tg mice show not only cerebellar ataxia but also absence epilepsy, which begins at approximately 3 weeks of age and persists throughout life. Similarities in EEG and sensitivity to antiepileptic drugs suggest that tg mice are a good model for human absence epilepsy. Although imbalance between excitatory and inhibitory activity in the thalamocortical network is thought to contribute to the pathogenesis of absence epilepsy, the effect of the mutation on thalamocortical synaptic responses remains unknown. Here we showed imbalanced impairment of inhibitory synaptic responses in tg mice using brain slice preparations. Somatosensory thalamocortical projection makes not only monosynaptic glutamatergic connections but also disynaptic GABAergic connections, which mediate feedforward inhibition, onto layer IV neurons. In tg mice, IPSC amplitudes recorded from layer IV pyramidal cells of the somatosensory cortex in response to thalamic stimulation became disproportionately reduced compared with EPSC amplitudes at later developmental stages (postnatal days 21-30). Similar results were obtained by local stimulation of layer IV pyramidal neurons. However, IPSC reduction was not seen in layer V pyramidal neurons of epileptic tg mice or in layer IV pyramidal neurons of younger tg mice before the onset of epilepsy (postnatal days 14-16). These results showed that the feedforward inhibition from the thalamus to layer IV neurons of the somatosensory cortex was severely impaired in tg mice and that the impairment of the inhibitory synaptic transmission was correlated to the onset of absence epilepsy.

6-Cyano-7-nitroquinoxaline-2,3-dione↗
PubMed1984

A quantitative analysis of the spatial organization of the vestibulo-ocular reflexes in lateral- and frontal-eyed animals--II. Neuronal networks underlying vestibulo-oculomotor coordination.

The neuronal connectivity underlying the vestibulo-ocular reflexes in cat and rabbit was evaluated in the light of quantitative data of the spatial orientation on semicircular canals and extraocular muscles. Neuronal connectivity was calculated using a matrix-analysis of the sensory and motor periphery, and of the brain stem pathways connecting semicircular canals and extraocular muscles. Two cases of vestibulo-ocular reflex compensation were considered. In the first case, vestibulo-oculor reflex compensation was assumed to be isotropic, i.e. the vestibulo-ocular reflex gain is the same for all directions of rotation. In the second case, the vestibulo-oculor reflex gain was assumed to be anisotropic with the "torsional" gain smaller than the "horizontal" and "vertical" gains. The theoretical calculation predicts that besides the principal vestibulo-ocular reflex pathways (classical three-neuron-arc connectivity), several accessory connections (other than principal connections, regardless of the synapses involved) exist which are characteristic for each species. These accessory connections were compared to physiological and anatomical data. In the cat theoretical connections for an isotropic vestibulo-ocular reflex gain agree with pathways observed experimentally, of which the most characteristic are excitatory connections to the superior rectus and inhibitory connections to the inferior rectus muscle from both of the anterior canals, and a mirror image pattern of connections from the posterior canals. In the rabbit experimentally obtained data and calculated connections rarely agree. However, for an anisotropic gain we find a higher rate of coincidence between experimental and theoretical connections. Our evaluation indicates, that accessory vestibulo-ocular reflex pathways serve to compensate for the incongruence between semicircular canal and extraocular muscle planes, at least in the cat. Available experimental data suggest an important role of a special subclass of accessory pathways via axon collaterals of principal projections (three-neuron-arc nature). With certain restrictions, the presented method of calculation promises to be a useful tool for a quantitative analysis of the vestibulo-ocular reflex.

Animals↗
PubMed2003

Aberrant neuronal connectivity in CHL1-deficient mice is associated with altered information processing-related immediate early gene expression.

In humans, loss or alteration of the CHL1/CALL gene may contribute to mental impairment associated with the 3p-syndrome, caused by distal deletions of the short (p) arm of chromosome 3, and schizophrenia. Mice deficient for the Close Homologue of L1 (CHL1) show aberrant connectivity of hippocampal mossy fibers and olfactory sensory axons, suggesting participation of CHL1 in the establishment of neuronal networks. Furthermore, behavioral studies showed that CHL1-deficient mice react differently towards novel experimental environments. These data raise the hypothesis that processing of information, possibly novel versus familiar, may be altered in the absence of CHL1. To test this hypothesis, brain activities were investigated after presentation of a novel, familiar, or neutral gustatory stimulus using metabolic mapping with ((14)C)-2-deoxyglucose (2-DG) and analysis of mRNA expression of the immediate early genes (IEGs) c-fos and arg 3.1/arc by in situ hybridization. 2-DG labeling revealed only small differences between CHL1-deficient and wild-type littermate mice. In contrast, while the specific novelty-induced increase in c-fos expression was maintained in most of the brain areas analyzed, c-fos mRNA expression was similar after the novel and familiar taste in several brain areas of the CHL1-deficient mice. Furthermore, in these mutants, arg 3.1/arc expression was slightly reduced after the novel taste and increased after the familiar taste, leading to a similar arg 3.1/arc mRNA expression after both stimuli. Our results indicate that, in contrast to controls, CHL1-deficient mice might process novel and familiar information similarly and suggest that the altered neuronal connectivity in these mutants disturbs information processing at the molecular level.

Animals↗
PubMed1996

Connectional networks within the orbital and medial prefrontal cortex of macaque monkeys.

The intrinsic cortico-cortical connections within the orbital and medial prefrontal cortex (OMPFC) were demonstrated with retrograde and anterograde tracers injected into each of the architectonic areas that constitute this region. Although many of the connections linked neighboring areas, others selectively connected relatively distant areas. Most, but not all, of the connections were reciprocal. Altogether, the connections formed at least two distinct networks within the OMPFC. The "orbital" prefrontal network linked most of the areas within the orbital cortex, with very few connections to medial prefrontal areas. Areas Iam, Iapm, Ial, 12l, 12m, and 12r in the caudal and lateral parts of the orbital cortex (which received inputs from several sensory modalities) had convergent connections with areas 13l, 13m, and 13b in the central orbital cortex, with further connections to the rostral orbital area 11l. For the connections between areas Iapm, Iam, Ial, 13m, 13l, and 11l, rostrally directed fibers arose mainly in layer V, whereas caudally directed fibers originated mainly in layer III. The "medial" prefrontal network selectively involved medial areas 14r, 14c, 24, 25, 32, and 10m, rostral orbital areas 10o and 11m, and agranular insular area Iai in the posterior orbital cortex. Two orbital areas, 13a and 12o, had substantial connections to both networks and may serve as points of interaction between them; otherwise there were relatively few interconnections. The two networks also had distinct connections with other cortical regions, with limbic structures, and with the mediodorsal thalamic nucleus. Their role in guidance of affective behaviour is discussed.

Animals↗
PubMed1981

Towards a theory of cell assemblies.

The term cell assembly, first introduced by D. O. Hebb, is defined in the framework of graph theory. This definition leads to some beautiful problems concerning the number and size of cell assemblies in large graphs. Some approaches to solve these problems are presented. In particular, the graphs Kn X Km are constructed that have n . m points, n + m - 2 connections per point, and at least 2n + 2m - 4 assemblies. Several new notions of connectivity in directed graphs are introduced and their relationships are investigated. The insight into these notions and their relationships will be helpful for further construction of graphs with many assemblies and/or high connectivity. The resulting graphs are not only important for the idea of cell assemblies in the content of neurodynamics, they may also find applications in the construction of communication networks and associative memories.

Brain↗
PubMed2005

General anesthesia and the neural correlates of consciousness.

The neural correlates of consciousness must be identified, but how? Anesthetics can be used as tools to dissect the nervous system. Anesthetics not only allow for the experimental investigation into the conscious-unconscious state transition, but they can also be titrated to subanesthetic doses in order to affect selected components of consciousness such as memory, attention, pain processing, or emotion. A number of basic neuroimaging examinations of various anesthetic agents have now been completed. A common pattern of regional activity suppression is emerging for which the thalamus is identified as a key target of anesthetic effects on consciousness. It has been proposed that a neuronal hyperpolarization block at the level of the thalamus, or thalamocortical and corticocortical reverberant loops, could contribute to anesthetic-induced unconsciousness. However, all anesthetics do not suppress global cerebral metabolism and cause a regionally specific effect on thalamic activity. Ketamine, a so-called dissociative anesthetic agent, increases global cerebral metabolism in humans at doses associated with a loss of consciousness. Nevertheless, it is proposed that those few anesthetics not associated with a global metabolic suppression effect might still have their effects on consciousness mediated at the level of thalamocortical interactions, if such agents scramble the signals associated with normal neuronal network reverberant activity. Functional and effective connectivity are analysis techniques that can be used with neuroimaging to investigate the signal scrambling effects of various anesthetics on network interactions. Whereas network interactions have yet to be investigated with ketamine, a thalamocortical and corticocortical disconnection effect during unconsciousness has been found for both suppressive anesthetic agents and for patients who are in the persistent vegetative state. Furthermore, recovery from a vegetative state is associated with a reconnection of functional connectivity. Taken together these intriguing observations offer strong empirical support that the thalamus and thalamocortical reverberant network loop interactions are at the heart of the neurobiology of consciousness.

Anesthesia, General↗
PubMed2001

Ion channels of importance for the locomotor pattern generation in the lamprey brainstem-spinal cord.

The intrinsic function of the spinal network that generates locomotion can be studied in the isolated brainstem-spinal cord of the lamprey, a lower vertebrate. The motor pattern underlying locomotion can be elicited in the isolated spinal cord. The network consists of excitatory glutamatergic and inhibitory glycinergic interneurones with known connectivity. The current review addresses the different subtypes of ion channels that are present in the cell types that constitute the network. In particular the roles of the different subtypes of Ca2+ channels and potassium channels that regulate integrated neuronal functions, like frequency regulation, spike frequency adaptation and properties that are important for generating features of the motor pattern (e.g. burst termination), are reviewed. By knowing the role of an ion channel at the cellular level, we also, based on previous knowledge of network connectivity, can understand which effect a given ion channel may exert at the different levels from molecule and cell to network and behaviour.

Animals↗
PubMed2000

Specific relationship between prefrontal neuronal N-acetylaspartate and activation of the working memory cortical network in schizophrenia.

OBJECTIVE: Abnormal activation of the dorsolateral prefrontal cortex and a related cortical network during working memory tasks has been demonstrated in patients with schizophrenia, but the responsible mechanism has not been identified. The present study was performed to determine whether neuronal pathology of the dorsolateral prefrontal cortex is linked to the activation of the working memory cortical network in patients with schizophrenia. METHOD: The brains of 13 patients with schizophrenia and 13 comparison subjects were studied with proton magnetic resonance spectroscopic ((1)H-MRS) imaging (to measure N-acetylaspartate as a marker of neuronal pathology) and with [(15)O]water positron emission tomography (PET) during performance of the Wisconsin Card Sorting Test (to measure activation of the working memory cortical network). An independent cohort of patients (N=7) was also studied in a post hoc experiment with (1)H-MRS imaging and with the same PET technique during performance of another working memory task (the "N-back" task). RESULTS: Measures of N-acetylaspartate in the dorsolateral prefrontal cortex strongly correlated with activation of the distributed working memory network, including the dorsolateral prefrontal, temporal, and inferior parietal cortices, during both working memory tasks in the two independent groups of patients with schizophrenia. In contrast, N-acetylaspartate in other cortical regions and in comparison subjects did not show these relationships. CONCLUSIONS: These findings directly implicate a population of dorsolateral prefrontal cortex neurons as selectively accounting for the activity of the distributed working memory cortical network in schizophrenia and complement other evidence that dorsolateral prefrontal cortex connectivity is fundamental to the pathophysiology of the disorder.

Adult↗
PubMed2005

Cortical abnormalities in bipolar disorder investigated with MRI and voxel-based morphometry.

Bipolar disorder (BD) has been associated with abnormalities of brain structure. Specifically, in vivo volumetric MRI and/or post mortem studies of BD have reported abnormalities of gray matter (GM) volume in the medial prefrontal cortex (PFC), amygdala, hippocampal subiculum and ventral striatum. These structures share anatomical connections with each other and form part of a "visceromotor" network modulating emotional behavior. Areas of the lateral orbital, superior temporal and posterior cingulate cortices project to this network, but morphometric abnormalities in these areas have not been established in BD. The current study assessed tissue volumes within these areas in BD using MRI and voxel-based morphometry (VBM). MRI images were obtained from 36 BD subjects and 65 healthy controls. To account for possible neurotrophic and neuroprotective effects of psychotropic medications, BD subjects were divided into medicated and unmedicated groups. Images were segmented into tissue compartments, which were examined on a voxel-wise basis to determine the location and extent of morphometric changes. The GM was reduced in the posterior cingulate/retrosplenial cortex and superior temporal gyrus of unmedicated BD subjects relative to medicated BD subjects and in the lateral orbital cortex of medicated BD subjects relative to controls. White matter (WM) was increased in the orbital and posterior cingulate cortices, which most likely reflected alterations in gyral morphology resulting from the reductions in the associated GM. The morphometric abnormalities in the posterior cingulate, superior temporal and lateral orbital cortices in BD support the hypothesis that the extended network of neuroanatomical structures subserving visceromotor regulation contains structural alterations in BD. Additionally, localization of morphometric abnormalities to areas known to exhibit increased metabolism in depression supports the hypothesis that repeated stress and elevated glucocorticoid secretion may result in neuroplastic changes in BD.

Adult↗
PubMed2003

Treatment of methamphetamine abuse: research findings and clinical directions.

Over the past few years, methamphetamine has appeared in mass quantities, in part, because of the ease and cost efficiency of manufacturing. With this increase in availability, the use of methamphetamine has increased significantly. The purpose of this article is to describe the existing treatment options for methamphetamine abuse and provide recommendations for practitioners and researchers. Methamphetamine abuse adversely impacts physical functioning, brain functioning and cognition, social support and social networks, and behavioral functioning. Negative consequences have also been documented to the environment and communities. In the studies reviewed on effective treatments, interventions consisted of aversion therapy, medication, psychosocial treatment, and case management. Each specific treatment is described as connected with an overall drug treatment program. If methamphetamine abuse continues to increase and the consequences continue to be so devastating, researchers and clinicians could advance the field by particular focus on the treatment of this type of drug use.

Adolescent↗
PubMed2003

Neural mechanisms of empathy in humans: a relay from neural systems for imitation to limbic areas.

How do we empathize with others? A mechanism according to which action representation modulates emotional activity may provide an essential functional architecture for empathy. The superior temporal and inferior frontal cortices are critical areas for action representation and are connected to the limbic system via the insula. Thus, the insula may be a critical relay from action representation to emotion. We used functional MRI while subjects were either imitating or simply observing emotional facial expressions. Imitation and observation of emotions activated a largely similar network of brain areas. Within this network, there was greater activity during imitation, compared with observation of emotions, in premotor areas including the inferior frontal cortex, as well as in the superior temporal cortex, insula, and amygdala. We understand what others feel by a mechanism of action representation that allows empathy and modulates our emotional content. The insula plays a fundamental role in this mechanism.

Adult↗
PubMed1998

What can monotremes tell us about brain evolution?

The present review outlines studies of electrophsyiological organization, cortical architecture and thalmocortical and corticocortical connections in monotremes. Results of these studies indicate that the neocortex of monotremes has many features in common with other mammals. In particular, monotremes have at least two, and in some instances three, sensory fields for each modality, as well as regions of bimodal cortex. The internal organization of cortical fields and thalamocortical projection patterns are also similar to those described for other mammals. However, unlike most mammals investigated, the monotreme neocortex has cortical connections between primary sensory fields, such as SI and VI. The results of this analysis lead us to pose the question of what monotremes can tell us about brain evolution. Monotremes alone can tell us very little about the evolutionary process, or the construction of complex neural networks, as an individual species represents only a single example of what the process is capable of generating. Perhaps a better question is: what can comparative studies tell us about brain evolution? Monotreme brains, when compared with the brains of other animals, can provide some answers to questions about the evolution of the neocortex, the historical precedence of some features over others, and how basic circuits were modified in different lineages. This, in turn, allows us to appreciate how normal circuits function, and to pose very specific questions regarding the development of the neocortex.

Animals↗
PubMed2002

Correlated random networks.

We develop a statistical theory of networks. A network is a set of vertices and links given by its adjacency matrix c, and the relevant statistical ensembles are defined in terms of a partition function Z= summation operator exp([-betaH(c)]. The simplest cases are uncorrelated random networks such as the well-known Erdös-Rényi graphs. Here we study more general interactions H(c) which lead to correlations, for example, between the connectivities of adjacent vertices. In particular, such correlations occur in optimized networks described by partition functions in the limit beta--> infinity. They are argued to be a crucial signature of evolutionary design in biological networks.

Brain↗
PubMed1999

Rehabilitation of brain damage: brain plasticity and principles of guided recovery.

Rehabilitation of the damaged brain can foster reconnection of damaged neural circuits; Hebbian learning mechanisms play an important part in this. The authors propose a triage of post-lesion states, depending on the loss of connectivity in particular circuits. A small loss of connectivity will tend to lead to autonomous recovery, whereas a major loss of connectivity will lead to permanent loss of function; for such individuals, a compensatory approach to recovery is required. The third group have potentially rescuable lesioned circuits, but guided recovery depends on providing precisely targeted bottom-up and top-down inputs, maintaining adequate levels of arousal, and avoiding activation of competitor circuits that may suppress activity in target circuits. Empirical data are implemented in a neural network model, and clinical recommendations for the practice of rehabilitation following brain damage are made.

Association Learning↗
PubMed1999

Forebrain connections of medial agranular cortex in the prairie vole, Microtus ochrogaster.

Fluorescent axonal tracers were used to investigate the connections of medial agranular cortex (frontal area 2, Fr2) in male prairie voles. The rostral and caudal portions of Fr2 (rFr2 and cFr2) have distinct but partially overlapping patterns of connections. Thalamic labeling after cFr2 injections was present in anteromedial nucleus (AM), ventrolateral nucleus (VL), lateral segment, mediodorsal nucleus (MDl), centrolateral nucleus (CL), ventromedial nucleus (VM), posterior nucleus (Po) and lateral posterior nucleus (LP). A band of labeled cells involving CL, central medial nucleus (CM) and rhomboid nucleus (Rh) formed a halo around the periphery of submedial (gelatinosus) nucleus (Sm). Within cFr2 there is a rostrocaudal gradient whereby projections from VL and MDl become progressively sparser caudally, whereas those from LP and Po become denser. Rostral Fr2 receives afferents from a similar group of thalamic nuclei, but has denser innervation from VL and MDl, lacks afferents from LP, and receives less input from nuclei around the periphery of Sm. Caudal Fr2 has extensive cortical connections including orbital cortex, rostral Fr2, Fr1, caudal parietal area 1 (Par1), parietal area 2 (Par2), and posterior parietal, retrosplenial and visual areas. Rostral Fr2 has similar connections with areas Fr1, Par1 and Par2; orbital connections focused in ventrolateral orbital cortex (VLO); connections with caudal Fr2; greatly reduced connections with posterior parietal cortex and the visual areas; and no connections with retrosplenial cortex. The axons linking rFr2 and cFr2 with each other and with other cortical areas travel predominately in the deep gray matter of layers VI and VII rather than in the white matter. Projections to the dorsal striatum from rFr2 are widespread in the head of the caudate, become progressively restricted to a dorsocentral focus more caudally, and disappear by the level of the anterior commissure. The projections from cFr2 are largely restricted to a focal dorsocentral region of the striatum and to the dorsolateral margin of the caudatoputamen. In comparison to area Fr2, the laterally adjacent area Fr1 has thalamic and cortical connections which are markedly restricted. Area Fr1 receives thalamic input from nuclei VL, anteroventral nucleus (AV), CL and Po, but none from mediodorsal nucleus (MD) or LP, and its input from VM is reduced. Cortical afferents to Fr1 originate from areas Fr2, caudal Par1 and Par2. Medial agranular cortex of prairie voles has a pattern of connections largely similar to that seen in rats, suggesting that area Fr2 in prairie voles is part of a cortical network that may mediate complex behaviors involving spatial orientation.

Animals↗