Analysis of temporal structure in spike trains of visual cortical area MT

The temporal structure of neuronal spike trains in the visual cortex can provide detailed information about the stimulus and about the neuronal implementation of visual processing. Spike trains recorded from the macaque motion area MT in previous studies (Newsome et al., 1989a; Britten et al., 1992; Zohary et al., 1994) are analyzed here in the context of the dynamic random dot stimulus which was used to evoke them. If the stimulus is incoherent, the spike trains can be highly modulated and precisely locked in time to the stimulus. In contrast, the coherent motion stimulus creates little or no temporal modulation and allows us to study patterns in the spike train that may be intrinsic to the cortical circuitry in area MT. Long gaps in the spike train evoked by the preferred direction motion stimulus are found, and they appear to be symmetrical to bursts in the response to the anti-preferred direction of motion. A novel cross-correlation technique is used to establish that the gaps are correlated between pairs of neurons. Temporal modulation is also found in psychophysical experiments using a modified stimulus. A model is made that can account for the temporal modulation in terms of the computational theory of biological image motion processing. A frequency domain analysis of the stimulus reveals that it contains a repeated power spectrum that may account for psychophysical and electrophysiological observations.

Some neurons tend to fire bursts of action potentials while others avoid burst firing. Using numerical and analytical models of spike trains as Poisson processes with the addition of refractory periods and bursting, we are able to account for peaks in the power spectrum near 40 Hz without assuming the existence of an underlying oscillatory signal. A preliminary examination of the local field potential reveals that stimulus-locked oscillation appears briefly at the beginning of the trial.

Latent variable models for neural data analysis

The brain is perhaps the most complex system to have ever been subjected to rigorous scientific investigation. The scale is staggering: over 10^11 neurons, each making an average of 10^3 synapses, with computation occurring on scales ranging from a single dendritic spine, to an entire cortical area. Slowly, we are beginning to acquire experimental tools that can gather the massive amounts of data needed to characterize this system. However, to understand and interpret these data will also require substantial strides in inferential and statistical techniques. This dissertation attempts to meet this need, extending and applying the modern tools of latent variable modeling to problems in neural data analysis.

It is divided into two parts. The first begins with an exposition of the general techniques of latent variable modeling. A new, extremely general, optimization algorithm is proposed - called Relaxation Expectation Maximization (REM) - that may be used to learn the optimal parameter values of arbitrary latent variable models. This algorithm appears to alleviate the common problem of convergence to local, sub-optimal, likelihood maxima. REM leads to a natural framework for model size selection; in combination with standard model selection techniques the quality of fits may be further improved, while the appropriate model size is automatically and efficiently determined. Next, a new latent variable model, the mixture of sparse hidden Markov models, is introduced, and approximate inference and learning algorithms are derived for it. This model is applied in the second part of the thesis.

The second part brings the technology of part I to bear on two important problems in experimental neuroscience. The first is known as spike sorting; this is the problem of separating the spikes from different neurons embedded within an extracellular recording. The dissertation offers the first thorough statistical analysis of this problem, which then yields the first powerful probabilistic solution. The second problem addressed is that of characterizing the distribution of spike trains recorded from the same neuron under identical experimental conditions. A latent variable model is proposed. Inference and learning in this model leads to new principled algorithms for smoothing and clustering of spike data.

Response of multiple simultaneously recorded macaque area LIP neurons in a memory saccade task

Cells in the lateral intraparietal cortex (LIP) of rhesus macaques respond vigorously and in spatially-tuned fashion to briefly memorized visual stimuli. Responses to stimulus presentation, memory maintenance, and task completion are seen, in varying combination from neuron to neuron. To help elucidate this functional segmentation a new system for simultaneous recording from multiple neighboring neurons was developed. The two parts of this dissertation discuss the technical achievements and scientific discoveries, respectively.

Technology. Simultanous recordings from multiple neighboring neurons were made with four-wire bundle electrodes, or tetrodes, which were adapted to the awake behaving primate preparation. Signals from these electrodes were partitionable into a background process with a 1/f-like spectrum and foreground spiking activity spanning 300-6000 Hz. Continuous voltage recordings were sorted into spike trains using a state-of-the-art clustering algorithm, producing a mean of 3 cells per site. The algorithm classified 96% of spikes correctly when tetrode recordings were confirmed with simultaneous intracellular signals. Recording locations were verified with a new technique that creates electrolytic lesions visible in magnetic resonance imaging, eliminating the need for histological processing. In anticipation of future multi-tetrode work, the chronic chamber microdrive, a device for long-term tetrode delivery, was developed.

Science. Simultaneously recorded neighboring LIP neurons were found to have similar preferred targets in the memory saccade paradigm, but dissimilar peristimulus time histograms, PSTH). A majority of neighboring cell pairs had a difference in preferred directions of under 45° while the trial time of maximal response showed a broader distribution, suggesting homogeneity of tuning with het erogeneity of function. A continuum of response characteristics was present, rather than a set of specific response types; however, a mapping experiment suggests this may be because a given cell's PSTH changes shape as well as amplitude through the response field. Spike train autocovariance was tuned over target and changed through trial epoch, suggesting different mechanisms during memory versus background periods. Mean frequency-domain spike-to-spike coherence was concentrated below 50 Hz with a significant maximum of 0.08; mean time-domain coherence had a narrow peak in the range ±10 ms with a significant maximum of 0.03. Time-domain coherence was found to be untuned for short lags (10 ms), but significantly tuned at larger lags (50 ms).

A study of the proposed Diamond Creek Reservoir

Politically the Colorado river is an interstate as well as an international stream. Physically the basin divides itself distinctly into three sections. The upper section from head waters to the mouth of San Juan comprises about 40 percent of the total of the basin and affords about 87 percent of the total runoff, or an average of about 15 000 000 acre feet per annum. High mountains and cold weather are found in this section. The middle section from the mouth of San Juan to the mouth of the Williams comprises about 35 percent of the total area of the basin and supplies about 7 percent of the annual runoff. Narrow canyons and mild weather prevail in this section. The lower third of the basin is composed of mainly hot arid plains of low altitude. It comprises some 25 percent of the total area of the basin and furnishes about 6 percent of the average annual runoff.

The proposed Diamond Creek reservoir is located in the middle section and is wholly within the boundary of Arizona. The site is at the mouth of Diamond Creek and is only 16 m. from Beach Spring, a station on the Santa Fe railroad. It is solely a power project with a limited storage capacity. The dam which creats the reservoir is of the gravity type to be constructed across the river. The walls and foundation are of granite. For a dam of 290 feet in height, the back water will be about 25 m. up the river.

The power house will be placed right below the dam perpendicular to the axis of the river. It is entirely a concrete structure. The power installation would consist of eighteen 37 500 H.P. vertical, variable head turbines, directly connected to 28 000 kwa. 110 000 v. 3 phase, 60 cycle generators with necessary switching and auxiliary apparatus. Each unit is to be fed by a separate penstock wholly embedded into the masonry.

Concerning the power market, the main electric transmission lines would extend to Prescott, Phoenix, Mesa, Florence etc. The mining regions of the mountains of Arizona would be the most adequate market. The demand of power in the above named places might not be large at present. It will, from the observation of the writer, rapidly increase with the wonderful advancement of all kinds of industrial development.

All these things being comparatively feasible, there is one difficult problem: that is the silt. At the Diamond Creek dam site the average annual silt discharge is about 82 650 acre feet. The geographical conditions, however, will not permit silt deposites right in the reservoir. So this design will be made under the assumption given in Section 4.

The silt condition and the change of lower course of the Colorado are much like those of the Yellow River in China. But one thing is different. On the Colorado most of the canyon walls are of granite, while those on the Yellow are of alluvial loess: so it is very hard, if not impossible, to get a favorable dam site on the lower part. As a visitor to this country, I should like to see the full development of the Colorado: but how about THE YELLOW!

Report of an investigation of a plan of elimination of grade crossings on the A.T. & S.F. Ry. in the city of Pasadena, California

The problem of the Atchison, Topeka, and Santa Fe railroad in Pasadena is a very dynamic one, as is readily recognized by engineers, city officials, and laymen. The route of the railroad was first laid out in the eighties and because of certain liberal concessions granted by the City of Pasadena, the right-of-way was located through Pasadena, despite the fact that the grade coming into the city either from Los Angeles or San Bernardino was enormous. Some years later, other transcontinental routes of the Santa Fe out of Los Angles were sought, and a right-of-way was obtained by way of Fullerton and Riverside to San Bernardino, where this route joins the one from Los Angeles through Pasadena. This route, however, is ten miles longer than the one through Pasadena, which means a considerable loss of time in a short diversion of approximately only sixty miles in length.