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Biomedical subjects

W J Freeman

Publications and source records attributed to W J Freeman.

At least 37 records · Page 2Linked to original sources
PubMed1994

Role of chaotic dynamics in neural plasticity.

Mathematical models are essential for the study of complex neural systems at all levels of the hierarchy from macromolecules through neurons to brain systems. ANN are readily available, but most of them are inappropriate for modeling brain function in normal behavior, because they stem from studies of neural systems in anesthetized or paralyzed animals which are capable only of reflux output. That class of models lacks the goal-directed, self-organizing properties of neural systems in behaving animals. In contrast to the stability of ANN and their reliance on asymptotic convergence to steady states (point attractors) and periodic oscillations (limit cycle attractors), BNN are intrinsically unstable. They continuously generate 'spontaneous' aperiodic activity that manifests the operation of chaotic dynamics undergoing repeated state transitions. Observations on the activity patterns of sensory cortex reveal that the perceptual outputs of BNN are by construction of spatial patterns and dynamic trajectories and not by computation using symbolic representations. Chaotic dynamics plays essential roles both in the construction of perceptions and in the continuing update of cortical populations, which requires selective synaptic modification during associative learning and habituation. Simultaneous multichannel recording from the olfactory bulb and cortex has given the following experimental results. (1) The cortical activity that relates to the perception of a sensory stimulus is carried macroscopically by a smaller number of single neurons e.g. 'units', 'feature detectors'. (2) The macroscopic activity reflects the meaning and significance of the stimulus for the experimental subject and not the stimulus as it is known to the observer. (3) The activity carries the meaning in spatial patterns, not in time series (the difference between a phonograph or radio and movie or TV). (4) The spatial patterns of activity that accompany previously learned stimuli or responses are changed by the introduction of new stimuli and also by modifications in reinforcement contingencies. There is no invariance in the memory store within the populations. (5) The patterns of activity are created by dynamic neural interactions in sensory cortex, not by registration or filtering of stimuli. There is no evidence for storage, retrieval, cross-correlation or logical tree search. (6) The dynamics is chaotic, not merely noisy, so that each act of perception involves a new construction by the cortex not by mere information processing. From these findings we infer that chaotic dynamics plays a crucial role in the construction of the associational contexts comprising the memory systems of experimental subjects.

Animals↗
PubMed1991

Asymmetric sigmoid non-linearity in the rat olfactory system.

The statistical relationship between multi-unit spike activity and simultaneously recorded local dendritic field potentials in the olfactory system of the waking rat was studied with chronically placed electrodes. The relationship had the form of a sigmoid increase in axonal firing probability conditional on the amplitude of dendritic potentials. These data were fitted with an asymmetric sigmoid curve previously derived from the Hodgkin-Huxley equations. The curve was fitted using non-linear regression to optimize its parameter: the maximal firing rate. The maximal rate also gave the steepness of the slope of the sigmoid. Pulse trains were recorded from excitatory and inhibitory neurons in the olfactory cortex (including the anterior olfactory nucleus, the prepyriform cortex and the lateral entorhinal area) as identified by the phase relations of the pulse probability and the dendritic potentials, and from the excitatory neurons in the bulb (the inhibitory granule cells do not give extracellularly detectable action potentials). All these neurons are known to interact in disynaptic negative feedback loops giving rise to oscillations. The same sigmoid function fit the data from both types of neurons in all locations. The curves for neurons in all parts of the olfactory cortex had a 3-fold higher slope and maximal value than the curves from bulbar neurons. The significances of this difference and of the asymmetric sigmoid are discussed in terms of models for olfactory oscillatory dynamics and pattern recognition.

Anesthesia↗
PubMed1990

Correlations between unit firing and EEG in the rat olfactory system.

The olfactory EEG of awake animals displays oscillatory bursts of activity in the gamma- (30-100 Hz) range. The bursts are correlated with inflow of air over the receptor layer in the nose. None of the inputs to the cortices that display these oscillations carries periodic signals in the gamma-range. Thus these bursts are generated locally, either by neuronal feedback interactions or by coupling of oscillatory neurons. In the first case if the oscillations are generated by negative feedback, then two classes of cells must exist: excitatory neurons and inhibitory neurons with the same frequency of oscillation but with a quarter cycle phase lag by the inhibitory cells from the excitatory cells. On the other hand, if the EEG's result from coupling of cells that are intrinsically oscillatory, there should be a broad but monomodal distribution of phase values. In order to determine the origin of these bursts, we performed simultaneous recordings of EEG and multi-unit spikes in the 4 parts of the olfactory system (olfactory bulb, anterior olfactory nucleus, prepyriform cortex and lateral entorhinal area) of awake and motivated rats. For each sample, the EEG and the multi-unit spikes were recorded from the same local neighborhood. The multi-unit electrode recorded pulses from the principal output neurons of the respective cortical areas. In all locations tested, the oscillations in pulse probabilities of firing were found to have the same frequency as the dominant EEG frequency. In all 4 structures two sets of cells were found. One set displayed pulses in phase with the EEG and the other set displayed pulses that led or lagged the EEG by approximately 1/4 cycle. These data confirm the negative feedback interaction model rather than the coupled oscillator model for the generation of the bursts in the olfactory system. The relevance of these findings to other cortical systems, in casu the visual cortex is discussed.

Animals↗
PubMed1989

Spatial EEG correlates of nonassociative and associative olfactory learning in rabbits.

Recent studies have shown that spatially distributed olfactory bulbar activity correlates with odor-specific behavioral responding (Coopersmith & Leon, 1984; Freeman & Grajski, 1987; Freeman & Schneider, 1982; Freeman & Viana di Prisco, 1986; Grajski, Breiman, Viana di Prisco, & Freeman, 1986; Gray, Freeman, & Skinner, 1986; Sullivan & Leon, 1986; Viana di Prisco & Freeman, 1985). The present studies established olfactory bulbar spatial electroencephalogram (EEG) correlates of nonassociative and associative learning in odorant stimulation in rabbits. Behavior was quantified by measuring magnitude and probability of the sniff response. It was shown that (a) olfactory bulbar spatial EEG amplitude patterns do not simply reflect odor (peripheral) stimulation, (b) repeated presentations of a nonreinforced odor initially reveal a transient EEG pattern change but the pattern change does not recur after the subject has habituated to the odor, and (c) repeated presentations of a reinforced odor (mild cutaneous shock), with a second nonreinforced odor serving as a control, reveal that coexisting, odor-specific spatial EEG amplitude patterns emerge with the acquisition of differential behavioral responding.

Animals↗
PubMed1987

Spatial patterns of visual cortical fast EEG during conditioned reflex in a rhesus monkey.

A preliminary assay was made of the existence of time-space coherence patterns of fast EEG activity in the visual cortex of a Rhesus monkey. The primary intent of the present study was to evaluate the similarities and differences in relation to the olfactory bulb, where such coherences have been described and have been demonstrated to be associated with behaviour. Segments 1.5 s in duration were recorded simultaneously without averaging from 16 to 35 subdural electrodes fixed over the left occipital lobe in an array 3.6 cm X 2.8 cm. Each segment was taken during the delivery of a visual conditioned stimulus (CS) and the performance of a conditioned response (CR) by a well-trained Rhesus monkey. The EEGs appeared chaotic with irregular bursts lasting 75-200 ms, resembling those in the olfactory EEG but with lower peak frequencies. Fourier spectra showed broad distributions of power resembling '1/f noise' with multiple peaks in the range of 20-40 Hz. Time intervals were selected where coherent activity seemed to be present at a number of electrodes. A dominant component waveform that was common to all channels was extracted by principal components analysis (PCA) of each segment. The distribution of the power of this component across the electrodes (the factor loadings) was used to describe the spatial pattern of the coherent cortical activity. Statistical analyses suggested that different patterns could be associated to the CS and the CR, as has been found in the olfactory system. These patterns remained stable over a 6 week recording interval. The patterns can be better discriminated, when the factor loadings of each channel are normalized to zero mean and unit variance, to discard a basic pattern of power distribution, which may reflect anatomical and electrode positioning factors that are related to behavioral information processing by the cortex. The wide spatial distribution of the common patterns found suggests that EEG patterns that manifest differing states of the visual cortex may also be accessible with scalp electrodes.

Animals↗
PubMed1987

Induction and maintenance of epileptiform activity in the rabbit olfactory bulb depends on centrifugal input.

A technique of cryogenic blockade was used in waking rabbits to produce complete and reversible isolation of the olfactory bulb from the rest of the brain. During cooling of the olfactory peduncle epileptiform activity occurred spontaneously in the pyriform cortex in 3 out of 20 sessions, but never in the bulb. Following removal of the cryoblockade, during the seizure state, epileptiform discharges appeared simultaneously in the bulb and pyriform cortex. In the control state, without cooling of the peduncle, epileptiform activity could be evoked in the bulb and cortex by intense electrical stimulation of either the bulb or the lateral olfactory tract. During the cryoblockade, however, intense stimulation of the bulb failed to evoke seizure-like discharges. The results demonstrate a dependency on more central olfactory structures for the induction and maintenance of epileptiform activity in the olfactory bulb.

Animals↗
PubMed1987

Simulation of chaotic EEG patterns with a dynamic model of the olfactory system.

The main parts of the central olfactory system are the bulb (OB), anterior nucleus (AON), and prepyriform cortex (PC). Each part consists of a mass of excitatory or inhibitory neurons that is modelled in its noninteractive state by a 2nd order ordinary differential equation (ODE) having a static nonlinearity. The model is called a KOe or a KOi set respectively; it is evaluated in the "open loop" state under deep anesthesia. Interactions in waking states are represented by coupled KO sets, respectively KIe (mutual excitation) and KIi (mutual inhibition). The coupled KIe and KIi sets form a KII set, which suffices to represent the dynamics of the OB, AON, and PC separately. The coupling of these three structures by both excitatory and inhibitory feedback loops forms a KIII set. The solutions to this high-dimensional system of ODEs suffice to simulate the chaotic patterns of the EEG, including the normal low-level background activity, the high-level relatively coherent "bursts" of oscillation that accompany reception of input to the bulb, and a degenerate state of an epileptic seizure determined by a toroidal chaotic attractor. An example is given of the Ruelle-Takens-Newhouse route to chaos in the olfactory system. Due to the simplicity and generality of the elements of the model and their interconnections, the model can serve as the starting point for other neural systems that generate deterministic chaotic activity.

Animals↗
PubMed1987

Relation of olfactory EEG to behavior: spatial analysis.

The hypothesis that odor-specific patterns of mitral cell activity during odor discrimination might be found in the corresponding spatial patterns of electroencephalogram (EEG) amplitude over a surgically accessible segment of the bulbar surface was tested in rabbits with chronically implanted electrode arrays. The spatial spectrum of the bulbar EEG was derived and compared with the spectrum predicted for the granule cell generator. Spatial filters were devised to identify, enhance, or remove the granule cell contribution to the EEG. Spatial deconvolution was applied to the filtered granule cell activity patterns to correct for distortion caused by volume conduction. The results indicated that the bulb generated odor-specific spatial patterns in rabbits trained to discriminate between two odors. The odor-specific information was not localizable to subsets of channels. This suggested that the discriminative output of the bulb involved the entire structure, even though the receptor input was delivered to limited subsets of mitral cells.

Animals↗
PubMed1987

Relation of olfactory EEG to behavior: factor analysis.

Rabbits were conditioned to lick (CR+) in response to one odor (CS+); another odor (CS-) served as a discriminative control (CR-). Electroencephalograms (EEGs) were recorded from arrays of 64 electrodes on the olfactory bulb in three stages, each with six sessions: in Stage I, odors A+ and B-; in Stage II, odors C+ and B-; and in Stage III, odors C+ and A-. Spatial EEG amplitude patterns were measured for multiple control (C), CS+, and CS- EEG bursts in each trial. Data were transformed via factor analysis and expressed by factor scores as spatial patterns specified by factor loadings. In discriminant analysis of the factor scores, we correctly classified the C and CS bursts on the average by 65-80% from all trials for each subject and session and by 75-90% for trials with correct CRs. The latter was confirmed with a stepwise linear discriminant analysis of the original 64-variable data. Factor patterns were relatively invariant within but changed between stages. The results implied that stable spatial patterns of bulbar activity emerged in respect to CSs under reinforcement and persisted until the stimulus-response contingencies were changed.

Animals↗
PubMed1986

Petit mal seizure spikes in olfactory bulb and cortex caused by runaway inhibition after exhaustion of excitation.

The olfactory bulb (OB), anterior olfactory nucleus (AON) and prepyriform cortex (PC) maintain 3 kinds of feedback among their populations of excitatory and inhibitory neurons: negative feedback, mutual excitation, and mutual inhibition. At normal levels of synaptic input these are balanced and give rise to chaotic and near-sinusoidal oscillatory EEG activity. Under intense repetitive electrical stimulation of the lateral olfactory tract (LOT), there is failure of the afferent excitatory terminals, perhaps due to transmitter depletion. In this circumstance there is deficient excitatory input under the condition of a high level of sustained activity among mutually inhibitory neurons. An instability develops in which some inhibitory neurons become more disinhibited (excited) and others more inhibited (less active) to the point of a paroxysmal discharge that is manifested in a massive compound IPSP of the excitatory neurons. The paroxysm terminates abruptly, but by mechanisms still unclear repeats at a rate of about 3/s for 10-70 s. It is accompanied by simultaneous ipsilateral twitching of the eyelids and muzzle, salivation, tearing, arrest, and lack of responding to sensory stimuli but without loss of posture, resembling absence in humans. It does not result from runaway mutual excitation, and it rarely culminates in full-blown convulsions. Similar spikes usually also occur in the OB and AON; the sequences of spikes appear to entrain. These normal and seizure EEGs are simulated with a network of non-linear differential equations, that is designed in conformance with the anatomy and physiology of the olfactory system. The seizure appears as an emergent property of the OB, AON and PC interactive system, that is due to an induced asymmetry in the feedback network that controls normal background activity.

Animals↗
PubMed1986

Chemical dependencies of learning in the rabbit olfactory bulb: acquisition of the transient spatial pattern change depends on norepinephrine.

Intracerebral cannulas were implanted in both olfactory bulbs of 6 rabbits. A surface electrode-array (8 X 8) was implanted epidurally on the lateral surface of the left bulb. Each rabbit was conditioned to respond to sniffing to an odor paired with cutaneous shock while receiving continuous intrabulbar infusion of either vehicle or propranolol (100 microM at 1 microliter/hr) in vehicle. After two training sessions to the original odor, a response to a new odor was conditioned under the influence of the alternate infusate. Electroencephalographic (EEG) activity was sampled on inspirations before and during odor presentations. During vehicle infusion a transient alteration in the pattern of activity was acquired that occurred during the second and third inspirations following presentation of the reinforced odor. The acquisition did not occur when propranolol was infused. No significant pattern changes occurred with unreinforced odors in either condition. There was no local anesthetic effect of the racemic mixture of propranolol found for any type of electric activity, including antidromic spike activity observed in an independent control group. Intrabulbar norepinephrine injection (100 microM, 10 microL) resulted in an amplitude increase of the bulbar 40-80-Hz EEG and a potentiation of the transient spatial pattern change to a novel odor, when compared with those observed during vehicle infusion. It is concluded that norepinephrine released under centrifugal control may act to prevent or delay habituation that otherwise occurs rapidly to unreinforced odors.

Animals↗
PubMed1986

Relation of olfactory EEG to behavior: time series analysis.

Oscillatory electroencephalographic bursts were measured from 64 electrodes implanted on the olfactory bulbs of rabbits. Oscillatory bursts that occurred before and during presentation of odorant conditioned stimuli (CSs) were selected in brief segments. Comparisons between the 64 traces and their spectra showed that, despite amplitude differences between channels, every burst had a common waveform over the entire array. The spectra showed 2 to 5 distinct peaks in each burst. Each trace was fitted with the sum of 5 cosines to express the burst in ten 8 X 8 matrices of amplitude and phase values at its peak frequencies. Two types of burst were identified. Those with dominant frequencies greater than 55 Hz had one narrow dominant spectral peak and reproducible spatial patterns of its amplitude within subgroups of bursts relating to control and odorant CS conditions. Those with dominant frequencies less than 55 Hz were disorderly; their spectra were broad, and their spatial patterns of amplitude did not reproduce within subgroups. A behavioral assay showed that the high- and not the low-frequency bursts contained odor-specific information.

Animals↗
PubMed1985

Spatial EEG patterns, non-linear dynamics and perception: the neo-Sherringtonian view.

Spatial analysis with preamplifier arrays and computers offers fresh perspectives on brain function. Realization of its potential depends on development of appropriate procedures for data processing and display, experimental paradigms to serve as benchmarks, and theories of brain function to predict what to look for and how to distinguish valid results from artifacts. Measurement of EEGs from arrays of 64 electrodes chronically implanted on the olfactory bulbs of rabbits that are trained to discriminate odorant conditioned stimuli show that the odorants induce spatially distinctive amplitude patterns of neural activity. The odor-specific information density is inferred to be uniform over the whole main bulb. The neural dynamics that produce these activity patterns emerge from the synaptically interactive sheet of excitatory mitral and inhibitory granule cells with distributed input and output tracts and with static nonlinearities deriving from the nerve impulse mechanism. Excitatory synapses between mitral cells are subject to modification when odorants are paired with unconditioned stimuli, thus forming nerve cell assemblies. Odorant-specific information established by a stimulus locally in the bulbar unit activity is integrated with past experience by an assembly, disseminated over the entire bulb on the order of 100 mm2 in area in a time period of 2.5 ms, and sustained for a time period on the order of 0.1 s. An arbitrary spatial sample on the order of 20% of bulbar EEG activity captures the entire integrated information albeit at lesser resolution than the whole. This synaptic mechanism of local input and global output may be common to all of the cerebral cortex. The implications are discussed for neocortical sensory systems, motor pattern generators, and goal-directed behavior in the context of self-organizing non-linear dynamic systems.

Aging↗
PubMed1985

Odor-related bulbar EEG spatial pattern analysis during appetitive conditioning in rabbits.

Mildly thirsty rabbits were classically conditioned by reinforcement with water to give a discriminative licking response to the presentation of odors. The jaw movement component of the licking conditioned response (JM CR) was elicited only by the reinforced odor; an increase in the relative frequency of sniffing (RR CR) occurred to both reinforced (CS+) and nonreinforced (CS-) odors. Oscillatory electroencephalographic bursts of high-frequency (40-80 Hz) potentials were recorded epidurally from the lateral olfactory bulb with 64-electrode arrays (8 X 8, 3.5 X 3.5 mm) chronically implanted. Emphasis was on comparing bursts during odor presentation with bursts preceding odor arrival on each trial. A "detection" burst was characterized as occurring immediately after odor arrival and before the sniff response. "Discrimination" bursts occurred during the RR CR and before the JM CR onset. Significant air-odor burst differences (together with sniffing) occurred through up to six sessions for both CS+ and CS- odors for "discrimination" bursts but not for "detection" bursts.

Animals↗