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R A Normann

Publications and source records attributed to R A Normann.

At least 19 recordsLinked to original sources

Trial-by-trial discrimination of three enantiomer pairs by neural ensembles in mammalian olfactory bulb.

Populations of output neurons in the mammalian olfactory bulb (OB) exhibit distinct, widespread spatial and temporal activation patterns when stimulated with odorants. However, questions remain as to how ensembles of mitral/tufted (M/T) neurons in the mammalian OB represent odorant information. In this report, the single-trial encoding limits of random ensembles of putative single- and multiunit M/T cells in the anesthetized rat OB during presentations of enantiomers of limonene, carvone, and 2-butanol are investigated using simultaneous multielectrode recording techniques. The results of these experiments are: the individual constituents of our recorded ensembles broadly represent information about the presented odorants, the ensemble single-trial response of small spatially distributed populations of M/T neurons can readily discriminate between six different odorants, and the most consistent odorant discrimination is attained when the ensemble consists of all available units and their responses are integrated over an entire breathing cycle. These results suggest that small differences in spike counts among the ensemble members become significant when taken within the context of the entire ensemble. This may explain how ensembles of broadly tuned OB neurons contribute to olfactory perception and may explain how small numbers of individual units receiving input from distinct olfactory receptor neurons can be combined to form a robust representation of odorants.

Action Potentials↗

Coding of position by simultaneously recorded sensory neurones in the cat dorsal root ganglion.

Muscle, cutaneous and joint afferents continuously signal information about the position and movement of individual joints. How does the nervous system extract more global information, for example about the position of the foot in space? To study this question we used microelectrode arrays to record impulses simultaneously from up to 100 discriminable nerve cells in the L6 and L7 dorsal root ganglia (DRG) of the anaesthetized cat. When the hindlimb was displaced passively with a random trajectory, the firing rate of the neurones could be predicted from a linear sum of positions and velocities in Cartesian (x, y), polar or joint angular coordinates. The process could also be reversed to predict the kinematics of the limb from the firing rates of the neurones with an accuracy of 1-2 cm. Predictions of position and velocity could be combined to give an improved fit to limb position. Decoders trained using random movements successfully predicted cyclic movements and movements in which the limb was displaced from a central point to various positions in the periphery. A small number of highly informative neurones (6-8) could account for over 80% of the variance in position and a similar result was obtained in a realistic limb model. In conclusion, this work illustrates how populations of sensory receptors may encode a sense of limb position and how the firing of even a small number of neurones can be used to decode the position of the limb in space.

Action Potentials↗

High-resolution analysis of the spatio-temporal activity patterns in rat olfactory bulb evoked by enantiomer odors.

Understanding how mammals process olfactory stimuli has motivated the development of tools and techniques which permit the simultaneous study of finely structured spatial and temporal patterns of neural activity. A technique is described that uses an array of 32 penetrating microelectrodes implanted bilaterally into the dorsal aspect of rat olfactory bulb to investigate the responses of mitral and tufted neurons to stimulation with simple enantiomer odor pairs at a number of concentrations. It is shown that stable, simultaneous recordings from up to 49 single- and multi-units can be performed for periods of up to 14 h. We show that such odors evoke unique spatial and fast-temporal activity patterns which may subserve odor discrimination. This technique is extensible to other systems neuroscience investigations of olfactory sensory processing.

Animals↗

High-resolution spatio-temporal mapping of visual pathways using multi-electrode arrays.

The parallel processing of visual information was studied with penetrating microelectrode arrays. We studied the high-resolution visuotopic organization of cat primary visual cortex, and the encoding of simple visual stimuli by ensembles of ganglion cells in the isolated turtle retina. The high-resolution visuotopic organization of visual cortex is non-conformal. Regions of visual cortex separated by 400 mu may have receptive field centers that are separated by as much as 3 degrees, or they may superimpose. Ganglion cells are 'generalists', and are poor specifiers of the color of full field visual stimuli. Groups of 'luminosity' type ganglion cells can assist in the specification of stimulus color, but even individual 'chromatic' ganglion cells are not capable of quality color specification. These basic studies have relevance to the development of visual neuroprostheses based upon electrical stimulation of the retina and cortex.

Animals↗

High-resolution two-dimensional spatial mapping of cat striate cortex using a 100-microelectrode array.

Much of our understanding of the visuotopic organization of striate cortex results from single-electrode penetrations and serial recording of receptive field properties. However, the quality of these maps is limited by imprecision in quantifying electrode position, combining data from multiple laminae, and eye drift during the measurement of the receptive field properties. We have addressed these concerns by using an array of 100 closely spaced microelectrodes to investigate the two-dimensional visuotopic organization of layer IV in cat striate cortex. This array allowed simultaneous measurement of the receptive field properties of multiple single units on a regularly spaced grid. We found the relationship between cortical and visual space to be locally non-conformal: the receptive field locations associated with a closely spaced line of electrodes appeared randomly scattered in visual space. To quantify the scatter, we fitted a linear transformation of electrode sites onto the associated receptive field locations. We found that the distribution of the difference between the predicted receptive field location and the measured location had standard deviations of 0.59 degrees and 0.45 degrees in the horizontal and the vertical axes, respectively. Although individual receptive field positions differed from the predicted locations in a non-conformal sense, the trend across multiple receptive fields followed the maps described elsewhere. We found, on average, that the 13 mm2 of cortex sampled by the array mapped onto a 5.8-degrees) region of visual space. From the scaling of this map and a combination of the statistics of the receptive field size (2.7+/-1.5 degrees) and scatter, we have explored the impact of electrode spacing on the completeness and redundancy in coverage of visual space sampled by an array. The simulation indicated an array with 1.2-mm spacing would completely sample the region of visual space addressed by the array. These results have implications for neuroprosthetic applications. Assuming phosphene organization resembles the visuotopic organization, remapping of visual space may be necessary to accommodate the scatter in phosphene locations.

Animals↗

Selective stimulation of cat sciatic nerve using an array of varying-length microelectrodes.

Restoration of motor function to individuals who have had spinal cord injuries or stroke has been hampered by the lack of an interface to the peripheral nervous system. A suitable interface should provide selective stimulation of a large number of individual muscle groups with graded recruitment of force. We have developed a new neural interface, the Utah Slanted Electrode Array (USEA), that was designed to be implanted into peripheral nerves. Its goal is to provide such an interface that could be useful in rehabilitation as well as neuroscience applications. In this study, the stimulation capabilities of the USEA were evaluated in acute experiments in cat sciatic nerve. The recruitment properties and the selectivity of stimulation were examined by determining the target muscles excited by stimulation via each of the 100 electrodes in the array and using force transducers to record the force produced in these muscles. It is shown in the results that groups of up to 15 electrodes were inserted into individual fascicles. Stimulation slightly above threshold was selective to one muscle group for most individual electrodes. At higher currents, co-activation of agonist but not antagonist muscles was observed in some instances. Recruitment curves for the electrode array were broader with twitch thresholds starting at much lower currents than for cuff electrodes. In these experiments, it is also shown that certain combinations of electrode pairs, inserted into an individual fascicle, excite fiber populations with substantial overlap, whereas other pairs appear to address independent populations. We conclude that the USEA permits more selective stimulation at much lower current intensities with more graded recruitment of individual muscles than is achieved by conventional cuff electrodes.

Animals↗

Population coding in spike trains of simultaneously recorded retinal ganglion cells.

To achieve a better understanding of the parallel information processing that takes place in the nervous system, many researchers have recently begun to use multielectrode techniques to obtain high spatial- and temporal-resolution recordings of the firing patterns of neural ensembles. Apart from the complexities of acquiring and storing single unit responses from large numbers of neurons, the multielectrode technique has provided new challenges in the analysis of the responses from many simultaneously recorded neurons. This paper provides insights into the problem of coding/decoding of retinal images by ensembles of retinal ganglion cells. We have simultaneously recorded the responses of 15 ganglion cells to visual stimuli of various intensities and wavelengths and analyzed the data using discriminant analysis. Models of stimulus encoding were generated and discriminant analysis used to estimate the wavelength and intensity of the stimuli. We find that the ganglion cells we have recorded from are non-redundant encoders of these stimulus features. While single ganglion cells are poor classifiers of the stimulus parameters, examination of the responses of only a few ganglion cells greatly enhances our ability to specify the stimulus wavelength and intensity. Of the parameters studied, we find that the rate of firing of the ganglion cells provides the most information about these stimulus parameters, while the timing of the first action potential provides almost as much information. While we are not suggesting that the brain is using these variables, our results show how a population of sensory neurons can encode stimulus features and suggest that the brain could potentially deduce reliable information about stimulus features from response patterns of retinal ganglion cell populations.

Action Potentials↗

A technique to prevent dural adhesions to chronically implanted microelectrode arrays.

Minimizing relative movements between neural tissues and arrays of microelectrodes chronically implanted into them is expected to greatly enhance the capacity of the microelectrodes to record from single cortical neurons on a long-term basis. We describe a new surgical technique to minimize the formation of adhesions between the dura and an implanted electrode array using a 12 microm (0.5 mil) thick sheet of Teflon film positioned between the array and the dura. A total of 15 cats were implanted using this technique. Gross examination of 12 implant sites at the time of sacrifice failed to find evidence of adhesions between the arrays and the dura when the Teflon(R) film remained in its initial position. In six implants from which recordings were made, an average of nine of the 11 (81%) connected electrodes in each array recorded evoked neural activity after 180 days post implantation. Further, on average, two separable units were identified on each of the implanted electrodes in these arrays. No significant change was found in the density of cell bodies around implanted electrodes of four of the implanted electrode arrays. However, histological evaluation of the implant sites revealed evidence of meningeal proliferation beneath the arrays. The technique described is shown to be effective at preventing adhesions between implanted electrode arrays and improve the characteristics of chronic recordings obtained with these structures.

Animals↗

A multielectrode array for intrafascicular recording and stimulation in sciatic nerve of cats.

The feasibility of implanting an array of penetrating electrodes into peripheral nerves is studied in acute experiments in the cat sciatic nerve. A novel, silicon-based array of microelectrodes, the Utah Electrode Array, was used, which contains 25 or 100 1-mm long electrodes that project out from a silicon substrate. Electrode arrays of this complexity, when inserted in the peripheral nerve, could cause significant compression of the nerve and block the conduction of action potentials. Using a high velocity insertion technique, the electrode array was implanted into the sciatic nerve. Compound action potentials were evoked by and recorded with cuff electrodes. Compound action potentials recorded 1 h after insertion were only slightly altered from those recorded before insertion. Single units were readily extracted from evoked multiunit neural recordings in response to cutaneous stimulation and limb rotation around joints. Current injections into the nerve through the electrodes evoked muscle twitches with currents in the 10 microA range. Recording and stimulation stability were demonstrated for periods of up to 60 h. We have shown that high density arrays of electrodes can be inserted into the peripheral nerve and can provide a stable recording and stimulating interface to individual peripheral nerve axons. Such an array may be useful in future neuroscience research and potential neuroprosthetic applications.

Action Potentials↗

A 100-channel system for real time detection and storage of extracellular spike waveforms.

As extracellular electrode arrays with 100 or more active recording sites become more widely used for simultaneous recording of neural ensembles, practical data acquisition systems that can efficiently accommodate high electrode counts are needed. To reduce the high data rates associated with extracellular recordings from these arrays, various algorithms and systems have been designed to provide complete online detection and classification of extracellular spike waveforms. However, many of these algorithms require significant user supervision to ensure accurate performance. In this paper, we discuss the design and validation of a 100-channel PC-based system that can be used with arrays of extracellular electrodes such as the Utah Electrode Array. Instead of comprehensive online spike analysis, the system performs online detection and storage of the spike waveforms for offline classification. This strategy preserves the data of interest, reduces system complexity, and requires less user supervision during experiments.

Action Potentials↗

Neuronal interactions improve cortical population coding of movement direction.

Interactions among groups of neurons in primary motor cortex (MI) may convey information about motor behavior. We investigated the information carried by interactions in MI of macaque monkeys using a novel multielectrode array to record simultaneously from 12-16 neurons during an arm-reaching task. Pairs of simultaneously recorded cells revealed significant correlations in their trial-to-trial firing rate variation when estimated over broad (600 msec) time intervals. This covariation was only weakly related to the preferred directions of the individual MI neurons estimated from the firing rate and did not vary significantly with interelectrode distance. Most significantly, in a portion of cell pairs, correlation strength varied with the direction of the arm movement. We evaluated to what extent correlated activity provided additional information about movement direction beyond that available in single neuron firing rate. A multivariate statistical model successfully classified direction from single trials of neural data. However, classification was consistently better when correlations were incorporated into the model as compared to one in which neurons were treated as independent encoders. Information-theoretic analysis demonstrated that interactions caused by correlated activity carry additional information about movement direction beyond that based on the firing rates of independently acting neurons. These results also show that cortical representations incorporating higher order features of population activity would be richer than codes based solely on firing rate, if such information can exploited by the nervous system.

Action Potentials↗

A neural interface for a cortical vision prosthesis.

The development of a cortically based vision prosthesis has been hampered by a lack of basic experiments on phosphene psychophysics. This basic research has been hampered by the lack of a means to safely stimulate large numbers of cortical neurons. Recently, a number of laboratories have developed arrays of silicon microelectrodes that could enable such basic studies on phosphene psychophysics. This paper describes one such array, the Utah electrode array, and summarizes neurosurgical, physiological and histological experiments that suggest that such an array could be implanted safely in visual cortex. We also summarize a series of chronic behavioral experiments that show that modest levels of electrical currents passed into cortex via this array can evoke sensory percepts. Pending the successful outcome of biocompatibility studies using such arrays, high count arrays of penetrating microelectrodes similar to this design could provide a useful tool for studies of the psychophysics of phosphene perception in human volunteers. Such studies could provide a proof-of-concept for cortically based artificial vision.

Animals↗

Chronic intracortical microstimulation (ICMS) of cat sensory cortex using the Utah Intracortical Electrode Array.

In an effort to assess the safety and efficacy of focal intracortical microstimulation (ICMS) of cerebral cortex with an array of penetrating electrodes as might be applied to a neuroprosthetic device to aid the deaf or blind, we have chronically implanted three trained cats in primary auditory cortex with the 100-electrode Utah Intracortical Electrode Array (UIEA). Eleven of the 100 electrodes were hard-wired to a percutaneous connector for chronic access. Prior to implant, cats were trained to "lever-press" in response to pure tone auditory stimulation. After implant, this behavior was transferred to "lever-presses" in response to current injections via single electrodes of the implanted arrays. Psychometric function curves relating injected charge level to the probability of response were obtained for stimulation of 22 separate electrodes in the three implanted cats. The average threshold charge/phase required for electrical stimulus detection in each cat was, 8.5, 8.6, and 11.6 nC/phase respectively, with a maximum charge/phase of 26 nC/phase and a minimum of 1.5 nC/phase thresholds were tracked for varying time intervals, and seven electrodes from two cats were tracked for up to 100 days. Electrodes were stimulated for no more than a few minutes each day. Neural recordings taken from the same electrodes before and after multiple electrical stimulation sessions were very similar in signal/noise ratio and in the number of recordable units, suggesting that the range of electrical stimulation levels used did not damage neurons in the vicinity of the electrodes. Although a few early implants failed, we conclude that ICMS of cerebral cortex to evoke a behavioral response can be achieved with the penetrating UIEA. Further experiments in support of a sensory cortical prosthesis based on ICMS are warranted.

Animals↗

Chronic recording capability of the Utah Intracortical Electrode Array in cat sensory cortex.

The Utah Intracortical Electrode Array (UIEA) is an array of 100 penetrating silicon microelectrodes designed to focally electrically stimulate or record neurons residing in a single layer up to 1.5 mm beneath the surface of the cerebral cortex. Apart from its use as a unique tool to study parallel processing in the central nervous system, this array could form the platform for a cortical neuroprosthetic system. Although the UIEA has been used extensively in acute neural recording and stimulation experiments, its long-term performance in a chronic application has yet to be demonstrated. As an initial investigation into the feasibility of long-term cortical recording with an array of microelectrodes, we have hard-wired a subset of 11 electrodes of the UIEA to a percutaneous connector. This chronic UIEA assembly was then implanted into the cerebral cortices of ten cats for durations ranging from 2 to 13 months; over which time, both random and stimulus-evoked single and multiple unit action potentials were periodically recorded. On average, after a 6-month implant period, 60% of implanted arrays could still record some type of activity. Post-sacrifice dissections revealed a fibrous encapsulation of the UIEA. Although most implanted cortex was histologically normal, evidence of a chronic astroglial response was seen in a few cases. The results of the reported experiments indicate that the UIEA can be successfully used for limited times in a chronic recording application.

Action Potentials↗

Light adaptation and sensitivity controlling mechanisms in vertebrate photoreceptors.

The human visual system can discriminate increment and decrement light stimuli over a wide range of ambient illumination; from moonlight to bright sunlight. Several mechanisms contribute to this property but the major ones reside in the retina and more specifically within the photoreceptors themselves. Numerous studies in retinae from cold- and warm-blooded vertebrates have demonstrated the ability of the photoreceptors to respond in a graded manner to light increments and decrements even if these are applied during a background illumination that is expected to saturate the cells. In all photoreceptors regardless of type and species, three cellular mechanisms have been identified that contribute to background desensitization and light adaptation. These gain controlling mechanisms include; response-compression due to the non-linearity of the intensity-response function, biochemical modulation of the phototransduction process and pigment bleaching. The overall ability of a photoreceptor to adapt to background lights reflects the relative contribution of each of these mechanisms and the light intensity range over which they operate. In rods of most species, response-compression tends to dominate these mechanisms at light levels too weak to cause significant pigment bleaching and therefore, rods exhibit saturation. In contrast, cones are characterized by powerful background-induced modulation of the phototransduction process at moderate to bright background intensities where pigment bleaching becomes significant.Therefore, cones do not exhibit saturation even when the level of ambient illumination is raised by 6-7 log units.

Adaptation, Ocular↗

A study of printed spiral coils for neuroprosthetic transcranial telemetry applications.

We have explored the use of printed spiral coils (PSC's) for neuroprosthetic transcranial telemetry applications. We fabricated two-dimensional PSC's on a thin (25 microns) polyimide substrate using copper (35 microns) as a conducting material. All the coils had a fixed inner diameter of 1.0 cm. We fabricated two sets of coils. One set of coils consisted of 2- to 5-turn circular and square spiral coils and had different trace widths (W), different spacings (S) between adjacent traces, and different outer diameters. The other set of coils consisted of 5-turn circular spiral coils and had fixed inner and outer diameters but different W to S ratios. We measured loss resistances (Rs and Rp) and quality factors (Q) of these coils at different resonating frequencies in the range of 5-40 MHz. Over this frequency range, we observed that for fixed inner and outer diameters, the coil with the largest W achieved the lowest Rs and the highest Rp and Q. These electrical properties and the fact that these coils can conform to the complex convoluted cortical surface suggest that a PSC [15] can provide a viable alternative to a conventional wire-wound coil for neuroprosthetic transcranial telemetry applications.

Biocompatible Materials↗

The Utah intracortical Electrode Array: a recording structure for potential brain-computer interfaces.

We investigated the potential of the Utah Intracortical Electrode Array (UIEA) to provide signals for a brain-computer interface (BCI). The UIEA records from small populations of neurons which have an average signal-to-noise ratio (SNR) of 6:1. We provide specific examples that show the activities of these populations of neurons contain sufficient information to perform control tasks. Results from a simple stimulus detection task using these signals as inputs confirm that the number of neurons present in a recording is significant in determining task performance. Increasing the number of units in a recording decreases the sensitivity of the response to the stimulus; decreasing the number of units in the recording, however, increases the variability of the response to the stimulus. We conclude that recordings from small populations of neurons, not single units, provide a reliable source of sufficiently stimulus selective signals which should be suitable for a BCI. In addition, the potential for simultaneous and proportional control of a large number of external devices may be realized through the ability of an array of microelectrodes such as the UIEA to record both spatial and temporal patterns of neuronal activation.

Action Potentials↗