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Generation of temporal patterns in the olivo-cerebellar system
Lecture
Thursday, January 22, 2009
Hour: 12:30
Location:
Jacob Ziskind Building
Generation of temporal patterns in the olivo-cerebellar system
Dr. Gilad Jacobson
Dept of Neurobiology
Hebrew University, Jerusalem
The olivo-cerebellar system plays a crucial role in timing of both motor and non-motor tasks. The mechanisms underlying this timing capability are still unclear. Here I propose a plausible mechanism in which a temporal pattern reflects accurate phase relationships between the oscillatory activity of olivary neurons. I provide evidence from chronic multi-electrode recordings in awake rats that inferior olive oscillations possess hitherto unknown properties that: (1) Oscillations in different parts of the inferior olive can maintain constant, non-zero phase differences; (2) The oscillation frequency of olivary neurons is co-modulated; and (3) Phase differences are well maintained despite frequency changes. Thus, the inferior olive can generate not only “clock ticks” at the oscillation cycle duration, but more importantly shorter intervals that emerge by combining different parts of the olivary circuitry. This enables the olivo-cerebellar circuit to support timing in the range implicated by behavioural studies.
Personal theories and self-images: Critical tools in the rehabilitation from a severe brain injury
Lecture
Sunday, January 18, 2009
Hour: 14:45
Location:
Arthur and Rochelle Belfer Building for Biomedical Research
Personal theories and self-images: Critical tools in the rehabilitation from a severe brain injury
Prof. Yoram Eshet
Dept of Psychology & Education
The Open University of Israel
The lecture is given by a person who suffers from a severe (right-parietal) brain injury from the Yom Kippur War (1973). It discusses the injury as it is perceived by the injured person. The lecture focuses on self-images of the injury and emphasizes the pivotal role of higher cognitive processes, such as personal theories and narratives, as critical tools for a successful; rehabilitation.
Learning to smell: Cortical plasticity and odor perception
Lecture
Wednesday, January 14, 2009
Hour: 10:30
Location:
Arthur and Rochelle Belfer Building for Biomedical Research
Learning to smell: Cortical plasticity and odor perception
Prof. Donald Wilson
New York University School of Medicine
& Emotional Brain Institute
Nathan Kline Institute for Psychiatric Research
Odor perception - discrimination and recognition of volatile chemicals in the environment – is critical for wide ranging behaviors including kin recognition, mate selection, predator avoidance, homing, and feeding. Most naturally occurring odors are complex mixtures, often containing hundreds of different components. Furthermore, natural odors invariably occur against odorous backgrounds. Thus, olfaction and odor perception involves far more than simple odor ligands binding to receptors in the nose. I will describe recent work
The tempotron: applications to visual and time-warp invariant auditory processing
Lecture
Tuesday, January 13, 2009
Hour: 12:30
Location:
Jacob Ziskind Building
The tempotron: applications to visual and time-warp invariant auditory processing
Dr. Robert Guetig
Racah Institute of Physics &
Interdisciplinary Center for Neural Computation
Hebrew University, Jerusalem
The timing of action potentials of sensory neurons contains substantial information about the eliciting stimuli. Although computational advantages of spike-timing-based neuronal codes have long been recognized, it is unclear whether and how neurons can learn to read out such representations. We propose a novel biologically plausible supervised synaptic learning rule, the tempotron, enabling neurons to efficiently learn a broad range of decision rules, even when information is embedded in the spatio-temporal structure of spike patterns and not in mean firing rates. We demonstrate the enhanced performance of the tempotron over the rate-based perceptron in reading out spike patterns from retinal ganglion cell populations. Extending the tempotron to conductance-based voltage kinetics, we show that this model can subserve time-warp invariant processing of afferent spike patterns. Furthermore, we show that the conductance-based tempotron can learn to balance excitation and inhibition to match its integration time constant to the temporal scale of a given processing task. We show that already a small population of neurons can solve the TI46 isolated digit speech recognition task with near perfect performance
How to migrate when immobilized: Novel role for Reelin in the migration of cortical neurons
Lecture
Wednesday, January 7, 2009
Hour: 12:30
Location:
Nella and Leon Benoziyo Building for Brain Research
How to migrate when immobilized: Novel role for Reelin in the migration of cortical neurons
Prof. Michael Frotscher
Institute of Anatomy & Cell Biology
University of Freiburg, Germany
Reelin, a glycoprotein of the extracellular matrix, is secreted by Cajal-Retzius cells in the marginal zone of the cortex and controls the radial migration of cortical neurons. Reelin signaling involves the lipoprotein receptors apolipoprotein E receptor 2 (ApoER2) and very low density lipoprotein receptor (VLDLR), the adapter protein Disabled1 (Dab1), and phosphatidylinositol-3-kinase (PI3K). In regulating neuronal migration, Reelin signaling eventually acts on the cytoskeleton; however, its effects on the dynamic reorganization of the cytoskeleton have remained obscure. In reeler mutants deficient in Reelin, the majority of cortical neurons are unable to migrate to their destinations, suggesting Reelin signaling to be essential for the dynamic cytoskeletal reorganization that is required for neurons to migrate.
In contrast, we show that Reelin signaling stabilizes the cytoskeleton by serine3 phosphorylation of n-cofilin, an actin-depolymerizing protein. Phosphorylation at serine3 renders n-cofilin unable to depolymerize F-actin. However, depolymerization of F-actin is required for cytoskeletal reorganization. The Reelin receptor ApoER2, Dab1, src family kinases (SFKs), and PI3K were found to be involved in n-cofilin serine3 phosphorylation. Phosphorylation of n-cofilin was observed in the leading processes of migrating neurons when they reached the Reelin-containing marginal zone. Using a stripe choice assay, we found neuronal processes to be stable on Reelin-coated stripes. In contrast, on control stripes they formed lamellipodia as a sign of ongoing growth. These new results indicate that Reelin-induced stabilization of neuronal processes anchors them to the marginal zone which is crucial for directional migration by nuclear translocation.
(Supported by the German Research Foundation, DFG: SFB 592)
Rule-Rationality versus Act-Rationality
Lecture
Tuesday, December 30, 2008
Hour: 12:30
Location:
Jacob Ziskind Building
Rule-Rationality versus Act-Rationality
Prof. Yisrael Aumann
Nobel Prize Laureate in Economics, 2005
The Center for the Study of Rationality
Hebrew University, Jerusalem
People's actions often deviate from rationality, i.e., self-interested behavior. We propose a paradigm called rule-rationality, according to which people do not maximize utility in each of their acts, but rather follow rules or modes of behavior that usually---but not always---maximize utility. Specifically, rather than choosing an act that maximizes utility among all possible acts in a given situation, people adopt rules that maximize average utility among all applicable rules, when the same rule is applied to many apparently similar situations. The distinction is analogous to that between Bentham's "act-utilitarianism'' and the "rule-utilitarianism'' of Mill, Harsanyi, and others. The genesis of such behavior is examined, and examples are given. The paradigm may provide a synthesis between rationalistic neo-classical economic theory and behavioral economics.
Nonlinearity, memory, and phase transitions in learning
Lecture
Thursday, December 25, 2008
Hour: 12:00
Location:
Nella and Leon Benoziyo Building for Brain Research
Nonlinearity, memory, and phase transitions in learning
Dr. Ilya Nemenman
Computer, Computation and Statistical Sciences Division & Center for Nonlinear Studies
Los Alamos National Laboratory
Abstracting from physiological details, I will present a theory that suggests an explanation behind critical periods in learning as a natural consequence of learning dynamics under a small and realistic set of assumptions. Surprisingly, the same theory offers an explanation for other animal learning phenomena, such as the tendency to reverse to the status quo following a transient learning experience. Additionally, the theory suggests simple experiments that can be used to prove or refute it. If the time permits, as a commercial for future results, I will finish the talk with a brief overview of recent attempts at LANL for petascale simulations of the mammalian visual cortex.
Salience-based selection: How does the brain ignore saliency?
Lecture
Tuesday, December 23, 2008
Hour: 12:30
Location:
Jacob Ziskind Building
Salience-based selection: How does the brain ignore saliency?
Dr. Carmel Mevorach
Behavioral Brain Sciences Centre
University of Birmingham UK
At any particular time the brain is bombarded with an almost infinite amount of visual information. Efficient behaviour, then, relies on a process of attentional selection which is required to filter out irrelevant stimuli and to prioritize the processing of relevant events. Importantly, this attentional prioritisation process needs to be flexible in order to be responsive to changes in behavioural relevance. Thus, bottom-up cues for attention must be modulated by top-down information, reflecting the goals of behaviour. In recent years, considerable neurobiological evidence has accumulated indicating that flexible visual selection is controlled by a fronto-parietal network within the brain. In particular, the posterior parietal cortex (PPC) has been implicated both when spatial selection is required and when selection is non-spatial. In a series of recent studies we have used converging operations to demonstrate a link between the PPC and a form of non-spatial selection – selecting on the basis of the relative salience of the stimuli. Using variants of the classic Global/Local task we orthogonally manipulated the level of shape that participants responded to and the salience of that information. Using experimental techniques such as neuropsychological studies, Trans-cranial Stimulation (TMS) and functional imaging (fMRI) we show that the PPC is sensitive to the relative saliency of the information so that selection can be based on whether the target or the distractor are more salient. Most importantly, we provide evidence for distinct roles played by the right and left PPC in selection and suppression of saliency, respectively. The data may also suggest how such complementary forms of selection are implemented in the brain.
Representation of the visual field in object-selective cortex
Lecture
Wednesday, December 17, 2008
Hour: 15:00
Location:
Arthur and Rochelle Belfer Building for Biomedical Research
Representation of the visual field in object-selective cortex
Dr. Rory Sayres
Dept of Psychology, Stanford University
Functional MRI (fMRI) studies have defined a series of visual processing regions in the human cortex, which are believed to enable visual recognition behaviors through a hierarchy of processing stages. At the higher stages in this hierarchy lie regions which preferentially respond to images of intact objects compared to other visual stimuli, a set of regions collectively termed object-selective cortex. Within object-selective cortex exist category-selective regions, which prefer particular categories of images over others (e.g., faces, body parts, houses or scenes). Initially these regions were considered non-retinotopic, but increasing evidence indicates substantial retinal position selectivity, and in some cases retinotopy, in these regions.
What is the representation of the visual field in object-selective regions? Are separate object- and category-selective regions part of a single map or embedded within a set of distinct visual field maps? We scanned seven subjects on separate experiments to localize object/category-selective regions, and measure visual field maps (GE 3T scanner). For retinotopic experiments, subjects viewed moving bar stimuli containing different stimuli, including slowly drifting checkerboards and frontal face images. The bars extended out to around 14° eccentricity from the fovea, and had a width of ~2.6°. We employed a recently-developed method for estimating population receptive fields
(pRFs) using fMRI (Dumoulin and Wandell, Neuroimage, 2008), which estimates pRF center and size for each cortical location.
Face-containing bars produced substantially larger responses than checkerboards along the fusiform gyrus, improving our ability to measure visual field maps in these regions. Eccentricity maps revealed two foveal representations, which may correspond to visual field map clusters previously identified as VO and VT (Wandell et al., Neuro-opth. Jpn., 2006). These foveas are within or adjacent to fusiform face-selective regions, and separated by smoothly-varying extra-foveal maps which are less face-selective. For several subjects, pRF sizes systematically increased with eccentricity in face-selective regions. The distribution of pRF sizes were substantially larger than in earlier visual cortex, but comparable to recent measurements made in lateral occipital cortex.
We find two spatially separate face-selective regions along the fusiform gyrus, with comparable visual field coverage, separated by a representation of intermediate eccentricities. This indicates these two regions are likely to fall within different visual field maps. Current work addresses possible effects of low-level visual features (e.g. spatial frequency) and stimulus visibility in driving the observed face-selective retinotopic responses. I will also present some preliminary data from retinotopic mapping with house-containing bars, and an examination of retinotopic organization in house- or scene-selective cortical regions.
Active sensing: from natural stimulus statistics to auditory object classification in echolocating bats
Lecture
Tuesday, December 16, 2008
Hour: 12:30
Location:
Jacob Ziskind Building
Active sensing: from natural stimulus statistics to auditory object classification in echolocating bats
Yossi Yovel
(Post-doc Ulanovsky Group)
Department of Neurobiology, WIS
Echolocating bats perceive their surroundings acoustically. They continuously emit sonar signals and analyze the returning echoes, which enables them to orient in space and acquire food in complete darkness. Natural echoes along with other natural sounds compose a major part of the bat's sensory world, and have likely played a key evolutionary role in shaping the design of the bat's echolocation system and the auditory computations in the bat brain. However, the statistics of natural complex echoes, as well as how bats utilize them, are poorly understood – especially in the context of sonar-based object classification. The goal of this work was to elucidate the natural acoustical stimuli in the bat's world. I will present data on the statistical properties of complex echoes from various classes of plants and will compare them to what is known about natural images. In addition I will use a machine learning approach to discuss ways that bats may use to classify these stimuli. Finally, I will also describe behavioral experiments that aimed to understand the strategy used by bats to classify natural stimuli.
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Interaction between the amygdala and the prefrontal cortex in emotional memory
Lecture
Tuesday, November 25, 2008
Hour: 12:30
Location:
Jacob Ziskind Building
Interaction between the amygdala and the prefrontal cortex in emotional memory
Dr. Mouna Maroun
Department of Neurobiology and Ethology
University of Haifa
The amygdala and the medial prefrontal cortex interact to guide emotional behavior. Alterations in the balance between these two structures can lead to persistent fear associations and to the development of anxiety disorders.
In this talk I will present work from my laboratory studying the interaction between these two structures in normal conditions and when exposed to a fearful or stressful experience.
We have recently found that fear and extinction learning induce differential changes in these two structures that could hint on the mechanisms by which these structures encode memories of fear and safety.
ON THE RELATIONSHIP BETWEEN MOTOR AND PERCEPTUAL BEHAVIOR –
Lecture
Wednesday, November 12, 2008
Hour: 12:00
Location:
Nella and Leon Benoziyo Building for Brain Research
ON THE RELATIONSHIP BETWEEN MOTOR AND PERCEPTUAL BEHAVIOR –
Dr. Andrei Gorea
Laboratoire Psychologie de la Perception
CNRS & Paris Descartes University
Starting with Goodale & Milner's (1992) neuropsychological observations, a large number of neuropsychological and psychophysical studies has documented a putative dissociation between perception and action. However, a closer inspection of this literature reveals a number of methodological and conceptual shortcomings. I shall present a series of experiments making use of a variety of psychophysical techniques designed to gauge the relationship between Response Times as well Saccade Perturbations and observers' Perceptual States as assessed for not-masked and masked (metacontrast) stimuli via Yes/No, Temporal Order Judgments and Anticipation Response Times paradigms. All these studies reveal a strong action-perceptual state correlation indicating that motor and perceptual responses are based on a unique internal response. A one-path-two-decisions stochastic race model drawing on standard Signal Detection Theory provides a fair account of some of these data, hence overruling the necessity of a two-paths model of visual processing.
New insights into the hallmarks of obsessive-compulsive disorder (OCD): The prevalence of incompleteness and pessimal behavior
Lecture
Tuesday, November 11, 2008
Hour: 12:15
Location:
Jacob Ziskind Building
New insights into the hallmarks of obsessive-compulsive disorder (OCD): The prevalence of incompleteness and pessimal behavior
Prof. David Eilam
Dept of Zoology, Tel Aviv University
Performance of OCD patients was compared with that of matched normal individuals who were asked to perform the same task that the patients ascribed to their performance. Sequences of consecutive functional acts were long in controls and short in OCD, whereas sequences of non-functional acts were short in controls and long in OCD. Non-functional acts accumulated as a "tail" after the natural termination of the task, supporting the notion of incompleteness as an underling mechanism in OCD. It is suggested that the identified properties are consistent with a recent hypothesis that the individual's attention in OCD shifts from a normal focus on structured actions to a pathological attraction onto the processing of basic acts, a shift that invariably overtaxes memory. Such characteristics and mechanisms of compulsive rituals may prove useful in objective assessment of psychiatric disorders, behavioral therapy, and OCD nosology.
An embedded subnetwork of highly active neurons in the cortex
Lecture
Wednesday, November 5, 2008
Hour: 14:30
Location:
Arthur and Rochelle Belfer Building for Biomedical Research
An embedded subnetwork of highly active neurons in the cortex
Dr. Lina Yassin
Dept of Biological Sciences &
Center for the Neural Basis of Cognition
Carnegie Mellon University, Pittsburgh, PA
In vivo and in vitro, spontaneous and evoked neuronal activity are sparsely distributed across neocortical networks, where only a small subset of cells show firing rates greater than 1 Hz. Understanding the stability, network connectivity, and functional properties of this active subpopulation has been hampered by an inability to identify and characterize these neurons in vitro. Here we use expression of a fosGFP transgene to identify and characterize the properties of cells with a recent history of elevated activity. Neurons that had induced fosGFP expression in vivo maintained elevated firing rates in vitro over the course of many hours. Paired-cell recordings indicated that fosGFP+ neurons have a greater likelihood of being connected to each other, both directly and indirectly. These findings indicate that highly active neuronal ensembles are maintained over long time periods and suggest that specific, identifiable groups of neurons may dominate the way information is represented in the neocortex.
Voltage-Gated Sodium Channels in Neocortical Pyramidal Neurons:
Lecture
Tuesday, November 4, 2008
Hour: 12:15
Location:
Jacob Ziskind Building
Voltage-Gated Sodium Channels in Neocortical Pyramidal Neurons:
Prof. Mike Gutnick
Koret School of Veterinary Medicine
The Hebrew University of Jerusalem, Rehovot
CARBOXYPEPTIDASE E: ROLE IN PEPTIDERGIC VESICLE TRANSPORT, NEUROPROTECTION AND CANCER
Lecture
Tuesday, October 28, 2008
Hour: 12:15
Location:
Jacob Ziskind Building
CARBOXYPEPTIDASE E: ROLE IN PEPTIDERGIC VESICLE TRANSPORT, NEUROPROTECTION AND CANCER
Dr. Y. Peng Loh
Section on Cellular Neurobiology,
Program on Developmental Neuroscience,
NICHD, NIH, Bethesda
Carboxypeptidase E (CPE) is a prohormone processing enzyme that cleaves C-terminal basic residues from peptide hormone intermediates to yield active hormones, within secretory granules of neuroendocrine cells. A transmembrane form of the enzyme has been shown to be a sorting receptor that sorts prohormones and BDNF at the trans Golgi network and targets them to the regulated secretory pathway. Recently, live cell imaging studies have demonstrated that transport of peptidergic/BDNF secretory vesicles to the release site is dependent upon CPE. The cytoplasmic tail of CPE on the vesicles binds to microtubule motors, KIF1A/KIF3A and dynein via dynactin to effect transport of prohormone/BDNF vesicles in a bidirectional manner from the soma to the process terminals and return. In addition, CPE has been found to play a neuroprotective role in adult brain. In CPE-knockout (KO) mice, degeneration of pyramidal neurons was observed in the hippocampal CA3 region of animals equal or greater than 4 weeks of age, whereas the hippocampus was intact at 3 weeks and younger. Calbindin staining indicated early termination of the mossy fibers before reaching the CA1 region, and a lack of staining of the pyramidal neurons and apical dendritic arborizations in the CA1 region of CPE-KO mice. Ex vivo studies showed that cultured hippocampal neurons transfected with an enzymatically inactive form of CPE were protected against H2O2 oxidative-stress-induced cell death but not in non-transfected or LacZ transfected neurons. Thus CPE has an anti-apoptotic role in the maintenance of survival of adult hippocampal CA3 neurons, although the mechanism of action is unknown. In hepatocellular carcinoma (HCC) cells, overexpression of CPE resulted in enhanced proliferation and migration. SiRNA knockdown of CPE expression in highly metastatic HCC cells inhibited their growth and metastasis in nude mice. These results indicate that CPE is a new mediator of tumor growth and metastasis. Thus CPE is a multi-functional protein which actions include both enzymatic and non-enzymatic to mediate various physiological functions.
Population imaging in vivo: from the awake to the anesthetized
Lecture
Tuesday, October 7, 2008
Hour: 12:15
Location:
Jacob Ziskind Building
Population imaging in vivo: from the awake to the anesthetized
Prof. Jason Kerr
Max Planck Institute, Tubingen, Germany
It is unclear how the complex spatiotemporal organization of ongoing cortical neuronal activity recorded in anesthetized animals relates to the awake animal. We therefore used two-photon population calcium imaging in awake and subsequently anesthetized rats to follow action potential firing in populations of neurons across brain states, and examined how single neurons contributed to population activity. Firing rates and spike bursting in awake rats were higher, and pair-wise correlations were lower, compared with anesthetized rats. Anesthesia modulated population-wide synchronization and the relationship between firing rate and correlation. Overall, brain activity during wakefulness cannot be inferred using anesthesia.
Decoding conscious and unconscious mental states from brain activity in humans
Lecture
Tuesday, September 23, 2008
Hour: 12:15
Location:
Jacob Ziskind Building
Decoding conscious and unconscious mental states from brain activity in humans
Prof. Dr. John-Dylan Haynes
Bernstein Center for Computational Neuroscience, Charité Berlin &
Max Planck Institute for Human Cognitive and Brain Sciences, Leipzig, Germany
Recent advances in human neuroimaging have shown that it is possible to accurately read out a person's conscious experience based only on non-invasive fMRI measurements of their brain activity. This "brain reading" is possible because each thought is associated with a unique pattern of brain activity that can serve as a "fingerprint" of this thought in the brain. By training a computer to recognize these fMRI "thought patterns" it is possible to read out what someone is currently thinking with high accuracy. Here several studies will be presented that also directly address the relationship between neural encoding of information (as measured with fMRI) and its availability for awareness. These studies include comparisons of neural and perceptual information, unconscious information processing, decoding of time courses of perception, as well as decoding of high-level mental states. This will show that it is possible to read out a person's concealed intentions and even to predict how someone is going to decide a few seconds later. Finally, the talk will discuss fundamental challenges and limitations of the field, along with the ethical question if such methods might one day be a danger to our mental privacy.
Comparing spontaneous and stimulus-evoked activities in human sensory cortex
Lecture
Tuesday, September 16, 2008
Hour: 12:15
Location:
Nella and Leon Benoziyo Building for Brain Research
Comparing spontaneous and stimulus-evoked activities in human sensory cortex
Yuval Nir (Rafi Malach Group)
Department of Neurobiology, WIS
Traditionally, the brain and sensory cortex in particular have been viewed as being primarily driven by external events, but recent studies in anesthetized animals revealed robust spontaneous activity in sensory cortex, highlighting the intrinsic nature of brain processing. Using fMRI we found widespread slow fluctuations occurring spontaneously in the human visual cortex in the absence of external stimuli. These waves exhibited a consistent and specific neuro-anatomical distribution, suggesting that they largely reflect neuronal activity rather than hemodynamic noise sources. In further studies we obtained neurophysiological recordings in neurosurgical patients, and found direct electrophysiological evidence for such slow spontaneous neuronal fluctuation in human sensory cortex. These fluctuations were evident mainly in neuronal firing rates and in LFP gamma power changes, showed unique temporal dynamics following 1/f power laws, and were found to be correlated between corresponding ‘mirror’ sites across hemispheres within specific functional networks. Overall, these results extend previous animal studies of spontaneous activity by revealing and characterizing such activity in human sensory cortex.
Strong Loops in the Neocortex
Lecture
Wednesday, August 13, 2008
Hour: 12:15
Location:
Wolfson Building for Biological Research
Strong Loops in the Neocortex
Prof. Henry Kennedy
Dept of Integrative Neuroscience
INSERM, France
Hierarchy provides a major conceptual framework for understanding structure-function relationships of the cortex (Felleman and Van Essen, Cerb Cortex 1991). Feedforward (rostral directed) projections link areas in an ascending series and have a driving influence; feedback (caudal directed) projections link areas in a descending series and have a modulatory influence. This has led to the suggestion that feedforward projections are uniquely reciprocated by feedback projections i.e no strong loops (Crick and Koch, Nature 1998). We have re-examined this issue by making retrograde tracer injections in 22 areas spanning the occipital, parietal, temporal and frontal lobes. Injections were placed in areas V1, V2, V4 TEO, STPa, STPm, STPp, AudPba, AudPbp, 5, 7a, 7b, F1, 2, 8a, 45b, 9/46d, 9/46v, 46d, F5, ProM, 24c. High frequency sampling allows determination of indices of laminar distribution (SLN) and the relative strength (FLN) of connections (Vezoli et al., The Neuroscientist 2004). Analysis shows an inverse relationship between strength of connection and distance and revealed many (30%) hitherto unknown long-distance connections. Elsewhere we have shown that cortico-cortical projections form a smooth gradient: long-distance ascending connections are strongly feedforward (high SLN XX 100%) and on approaching the injection site have progressively lower SLN values (reaching 51%); likewise long-distance descending connections are strongly feedback (low SLN XX 0) and approaching the injection site reduce SLN 49% (Barone and Kennedy, J. Neurosci. 2000). The Felleman and Van Essen data is strictly hierarchical (no strong loops). A topological model of our data shows small world features (high cluster index and short average path distances) and five strong loops. Strong loops link frontal areas with occipital (areas 45-V4, 8A-V4), temporal (areas 45-TEO, 46-TEO) and parietal (areas 8A-7A, 46-7A) areas. The areas participating in strong loops exhibit high degrees of connectivity and constitute the hubs promoting small world attributes in the cortical architecture. The strong loops make it possible to go from V4 to all higher areas and back to V4 by uniqely feedforward pathways in an average of 3 and a maximum of 8 steps. One consequence of these anti-hierarchical connections is that the computations carried out in the supragranular layers of the cortex (Douglas and Martin, Annual Rev Neurosci. 2004) can be widely distributed in large-scale cortical networks mediating top-down control.
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