Stimulus Integration and Parsing in the Primate Auditory Midbrain

Integrating information from multiple sources is a crucial function of the brain. Examples of such integration include multiple stimuli of different modalties, such as visual and auditory, multiple stimuli of the same modality, such as auditory and auditory, and integrating stimuli from the sensory organs (i.e. ears) with stimuli delivered from brain-machine interfaces.

The overall aim of this body of work is to empirically examine stimulus integration in these three domains to inform our broader understanding of how and when the brain combines information from multiple sources.

First, I examine visually-guided auditory, a problem with implications for the general problem in learning of how the brain determines what lesson to learn (and what lessons not to learn). For example, sound localization is a behavior that is partially learned with the aid of vision. This process requires correctly matching a visual location to that of a sound. This is an intrinsically circular problem when sound location is itself uncertain and the visual scene is rife with possible visual matches. Here, we develop a simple paradigm using visual guidance of sound localization to gain insight into how the brain confronts this type of circularity. We tested two competing hypotheses. 1: The brain guides sound location learning based on the synchrony or simultaneity of auditory-visual stimuli, potentially involving a Hebbian associative mechanism. 2: The brain uses a ‘guess and check’ heuristic in which visual feedback that is obtained after an eye movement to a sound alters future performance, perhaps by recruiting the brain’s reward-related circuitry. We assessed the effects of exposure to visual stimuli spatially mismatched from sounds on performance of an interleaved auditory-only saccade task. We found that when humans and monkeys were provided the visual stimulus asynchronously with the sound but as feedback to an auditory-guided saccade, they shifted their subsequent auditory-only performance toward the direction of the visual cue by 1.3-1.7 degrees, or 22-28% of the original 6 degree visual-auditory mismatch. In contrast when the visual stimulus was presented synchronously with the sound but extinguished too quickly to provide this feedback, there was little change in subsequent auditory-only performance. Our results suggest that the outcome of our own actions is vital to localizing sounds correctly. Contrary to previous expectations, visual calibration of auditory space does not appear to require visual-auditory associations based on synchrony/simultaneity.

My next line of research examines how electrical stimulation of the inferior colliculus influences perception of sounds in a nonhuman primate. The central nucleus of the inferior colliculus is the major ascending relay of auditory information before it reaches the forebrain, and thus an ideal target for understanding low-level information processing prior to the forebrain, as almost all auditory signals pass through the central nucleus of the inferior colliculus before reaching the forebrain. Thus, the inferior colliculus is the ideal structure to examine to understand the format of the inputs into the forebrain and, by extension, the processing of auditory scenes that occurs in the brainstem. Therefore, the inferior colliculus was an attractive target for understanding stimulus integration in the ascending auditory pathway.

Moreover, understanding the relationship between the auditory selectivity of neurons and their contribution to perception is critical to the design of effective auditory brain prosthetics. These prosthetics seek to mimic natural activity patterns to achieve desired perceptual outcomes. We measured the contribution of inferior colliculus (IC) sites to perception using combined recording and electrical stimulation. Monkeys performed a frequency-based discrimination task, reporting whether a probe sound was higher or lower in frequency than a reference sound. Stimulation pulses were paired with the probe sound on 50% of trials (0.5-80 µA, 100-300 Hz, n=172 IC locations in 3 rhesus monkeys). Electrical stimulation tended to bias the animals’ judgments in a fashion that was coarsely but significantly correlated with the best frequency of the stimulation site in comparison to the reference frequency employed in the task. Although there was considerable variability in the effects of stimulation (including impairments in performance and shifts in performance away from the direction predicted based on the site’s response properties), the results indicate that stimulation of the IC can evoke percepts correlated with the frequency tuning properties of the IC. Consistent with the implications of recent human studies, the main avenue for improvement for the auditory midbrain implant suggested by our findings is to increase the number and spatial extent of electrodes, to increase the size of the region that can be electrically activated and provide a greater range of evoked percepts.

My next line of research employs a frequency-tagging approach to examine the extent to which multiple sound sources are combined (or segregated) in the nonhuman primate inferior colliculus. In the single-sound case, most inferior colliculus neurons respond and entrain to sounds in a very broad region of space, and many are entirely spatially insensitive, so it is unknown how the neurons will respond to a situation with more than one sound. I use multiple AM stimuli of different frequencies, which the inferior colliculus represents using a spike timing code. This allows me to measure spike timing in the inferior colliculus to determine which sound source is responsible for neural activity in an auditory scene containing multiple sounds. Using this approach, I find that the same neurons that are tuned to broad regions of space in the single sound condition become dramatically more selective in the dual sound condition, preferentially entraining spikes to stimuli from a smaller region of space. I will examine the possibility that there may be a conceptual linkage between this finding and the finding of receptive field shifts in the visual system.

In chapter 5, I will comment on these findings more generally, compare them to existing theoretical models, and discuss what these results tell us about processing in the central nervous system in a multi-stimulus situation. My results suggest that the brain is flexible in its processing and can adapt its integration schema to fit the available cues and the demands of the task.

Other Than a Citizen: Vernacular Poetics in Postwar America

Few symbols of 1950s-1960s America remain as central to our contemporary conception of Cold War culture as the iconic ranch-style suburban home. While the house took center stage in the Nixon/Khrushchev kitchen debates as a symbol of modern efficiency and capitalist values, its popularity depended largely upon its obvious appropriation of vernacular architecture from the 19th century, those California haciendas and Texas dogtrots that dotted the American west. Contractors like William Levitt modernized the historical common houses, hermetically sealing their porous construction, all while using the ranch-style roots of the dwelling to galvanize a myth of an indigenous American culture. At a moment of intense occupational bureaucracy, political uncertainty and atomized social life, the rancher gave a self-identifying white consumer base reason to believe they could master their own plot in the expansive frontier. Only one example of America’s mid-century love affair with commodified vernacular forms, the ranch-style home represents a broad effort on the part of corporate and governmental interest groups to transform the vernacular into a style that expresses a distinctly homogenous vision of American culture. “Other than a Citizen” begins with an anatomy of that transformation, and then turns to the work of four poets who sought to reclaim the vernacular from that process of standardization and use it to countermand the containment-era strategies of Cold War America.

In four chapters, I trace references to common speech and verbal expressivity in the poetry and poetic theory of Charles Olson, Robert Duncan, LeRoi Jones/Amiri Baraka and Gwendolyn Brooks, against the historical backdrop of the Free-Speech Movement and the rise of mass-culture. When poets frame nonliterary speech within the literary page, they encounter the inability of writing to capture the vital ephemerality of verbal expression. Rather than treat this limitation as an impediment, the writers in my study use the poem to dramatize the fugitivity of speech, emphasizing it as a disruptive counterpoint to the technologies of capture. Where critics such as Houston Baker interpret the vernacular strictly in terms of resistance, I take a cue from the poets and argue that the vernacular, rooted etymologically at the intersection of domestic security and enslaved margin, represents a gestalt form, capable at once of establishing centralized power and sparking minor protest. My argument also expands upon Michael North’s exploration of the influence of minstrelsy and regionalism on the development of modernist literary technique in The Dialect of Modernism. As he focuses on writers from the early 20th century, I account for the next generation, whose America was not a culturally inferior collection of immigrants but an imperial power, replete with economic, political and artistic dominance. Instead of settling for an essentially American idiom, the poets in my study saw in the vernacular not phonetic misspellings, slang terminology and fragmented syntax, but the potential to provoke and thereby frame a more ethical mode of social life, straining against the regimentation of citizenship.

My attention to the vernacular argues for an alignment among writers who have been segregated by the assumption that race and aesthetics are mutually exclusive categories. In reading these writers alongside one another, “Other than a Citizen” shows how the avant-garde concepts of projective poetics and composition by field develop out of an interest in black expressivity. Conversely, I trace black radicalism and its emphasis on sociality back to the communalism practiced at the experimental arts college in Black Mountain, North Carolina, where Olson and Duncan taught. In pressing for this connection, my work reveals the racial politics embedded within the speech-based aesthetics of the postwar era, while foregrounding the aesthetic dimension of militant protest.

Not unlike today, the popular rhetoric of the Cold War insists that to be a citizen involves defending one’s status as a rightful member of an exclusionary nation. To be other than a citizen, as the poets in my study make clear, begins with eschewing the false certainty that accompanies categorical nominalization. In promoting a model of mutually dependent participation, these poets lay the groundwork for an alternative model of civic belonging, where volition and reciprocity replace compliance and self-sufficiency. In reading their lines, we become all the more aware of the cracks that run the length of our load-bearing walls.

Passive Acoustics: A Multifaceted Tool for Marine Mammal Conservation

Marine mammals exploit the efficiency of sound propagation in the marine environment for essential activities like communication and navigation. For this reason, passive acoustics has particularly high potential for marine mammal studies, especially those aimed at population management and conservation. Despite the rapid realization of this potential through a growing number of studies, much crucial information remains unknown or poorly understood. This research attempts to address two key knowledge gaps, using the well-studied bottlenose dolphin (Tursiops truncatus) as a model species, and underwater acoustic recordings collected on four fixed autonomous sensors deployed at multiple locations in Sarasota Bay, Florida, between September 2012 and August 2013. Underwater noise can hinder dolphin communication. The ability of these animals to overcome this obstacle was examined using recorded noise and dolphin whistles. I found that bottlenose dolphins are able to compensate for increased noise in their environment using a wide range of strategies employed in a singular fashion or in various combinations, depending on the frequency content of the noise, noise source, and time of day. These strategies include modifying whistle frequency characteristics, increasing whistle duration, and increasing whistle redundancy. Recordings were also used to evaluate the performance of six recently developed passive acoustic abundance estimation methods, by comparing their results to the true abundance of animals, obtained via a census conducted within the same area and time period. The methods employed were broadly divided into two categories – those involving direct counts of animals, and those involving counts of cues (signature whistles). The animal-based methods were traditional capture-recapture, spatially explicit capture-recapture (SECR), and an approach that blends the “snapshot” method and mark-recapture distance sampling, referred to here as (SMRDS). The cue-based methods were conventional distance sampling (CDS), an acoustic modeling approach involving the use of the passive sonar equation, and SECR. In the latter approach, detection probability was modelled as a function of sound transmission loss, rather than the Euclidean distance typically used. Of these methods, while SMRDS produced the most accurate estimate, SECR demonstrated the greatest potential for broad applicability to other species and locations, with minimal to no auxiliary data, such as distance from sound source to detector(s), which is often difficult to obtain. This was especially true when this method was compared to traditional capture-recapture results, which greatly underestimated abundance, despite attempts to account for major unmodelled heterogeneity. Furthermore, the incorporation of non-Euclidean distance significantly improved model accuracy. The acoustic modelling approach performed similarly to CDS, but both methods also strongly underestimated abundance. In particular, CDS proved to be inefficient. This approach requires at least 3 sensors for localization at a single point. It was also difficult to obtain accurate distances, and the sample size was greatly reduced by the failure to detect some whistles on all three recorders. As a result, this approach is not recommended for marine mammal abundance estimation when few recorders are available, or in high sound attenuation environments with relatively low sample sizes. It is hoped that these results lead to more informed management decisions, and therefore, more effective species conservation.