Adaptations to changes in the acoustic scene of the echolocating bat

Our natural environment is noisy and in order to navigate it successfully, we must filter out the important components so that we may guide our next steps. In analyzing our acoustic scene, one of the most common challenges is to segregate speech communication sounds from background noise; this process is not unique to humans. Echolocating bats emit high frequency biosonar signals and listen to echoes returning off objects in their environment. The sound wave they receive is a merging of echoes reflecting off target prey and other scattered objects, conspecific calls and echoes, and any naturally-occurring environmental noises. The bat is faced with the challenge of segregating this complex sound wave into the components of interest to adapt its flight and echolocation behavior in response to fast and dynamic environmental changes. In this thesis, we employ two approaches to investigate the mechanisms that may aid the bat in analyzing its acoustic scene. First, we test the bat’s adaptations to changes of controlled echo-acoustic flow patterns, similar to those it may encounter when flying along forest edges and among clutter. Our findings show that big brown bats adapt their flight paths in response to the intervals between echoes, and suggest that there is a limit to how close objects can be spaced, before the bat does not represent them as distinct any longer. Further, we consider how bats that use different echolocation signals may navigate similar environments, and provide evidence of species-specific flight and echolocation adaptations. Second, we research how temporal patterning of echolocation calls is affected during competitive foraging of paired bats in open and cluttered environments. Our findings show that “silent behavior”, the ceasing of emitting echolocation calls, which had previously been proposed as a mechanism to avoid acoustic interference, or to “eavesdrop” on another bat, may not be as common as has been reported

ORIENTING IN 3D SPACE: BEHAVIORAL AND NEUROPHYSIOLOGICAL STUDIES IN BIG BROWN BATS

In their natural environment, animals engage in a wide range of behavioral tasks that require them to orient to stimuli in three-dimensional space, such as navigating around obstacles, reaching for objects and escaping from predators. Echolocating bats, for example, have evolved a high-resolution 3D acoustic orienting system that allows them to localize and track small moving targets in azimuth, elevation and range. The bat’s active control over the features of its echolocation signals contributes directly to the information represented in its sonar receiver, and its adaptive adjustments in sonar signal design provide a window into the acoustic features that are important for different behavioral tasks. When bats inspect sonar objects and require accurate 3D localization of targets, they produce sonar sound groups (SSGs), which are clusters of sonar calls produced at short intervals and flanked by long interval calls. SSGs are hypothesized to enhance the bat’s range resolution, but this hypothesis has not been directly tested. We first, in Chapter 2, provide a comprehensive comparison of SSG production of bats flying in the field and in the lab under different environmental conditions. Further, in Chapter 3, we devise an experiment to specifically compare SSG production under conditions when target motion is predictable and unpredictable, with the latter mimicking natural conditions where bats chase erratically moving prey. Data from both of these studies are consistent with the hypothesis that SSGs improve the bat’s spatio-temporal resolution of target range, and provide a behavioral foundation for the analysis and interpretation of neural recording data in chapters 4 and 6. The complex orienting behaviors exhibited by animals can be understood as a feedback loop between sensing and action. A primary brain structure involved in sensorimotor integration is the midbrain superior colliculus (SC). The SC is a widely studied brain region and has been implicated in species-specific orienting behaviors. However, most studies of the SC have investigated its functional organization using synthetic 2D (azimuth and elevation) stimuli in restrained animals, leaving gaps in our knowledge of how 3D space (azimuth, elevation and distance) is represented in the CNS. In contrast, the representation of stimulus distance in the auditory systems of bats has been widely studied. Almost all of these studies have been conducted in passively listening bats, thus severing the loop between sensing and action and leaving gaps in our knowledge regarding how target distance is represented in the auditory system of actively echolocating bats. In chapters 4, 5 and 6, we attempt to fill gaps in our knowledge by recording from the SC of free flying echolocating bats engaged in a naturalistic navigation task where bats produce SSGs. In chapter 4, we provide a framework to compute time-of-arrival and direction of the instantaneous echo stimuli received at the bats ears. In chapters 5 and 6, we provide an algorithm to classify neural activity in the SC as sensory, sensorimotor and premotor and then compute spatial receptive fields of SC neurons. Our results show that neurons in the SC of the free-flying echolocating bat respond selectively to stimulus azimuth, elevation and range. Importantly, we find that SC neuron response profiles are modulated by the bat’s behavioral state, indicated by the production of SSG. Broadly, we use both behavior and electrophysiology to understand the action-perception loop that supports spatial orientation by echolocation. We believe that the results and methodological advances presented here will open doors to further studies of sensorimotor integration in freely behaving animals

Exploring Animal Behavior Through Sound: Volume 1

This open-access book empowers its readers to explore the acoustic world of animals. By listening to the sounds of nature, we can study animal behavior, distribution, and demographics; their habitat characteristics and needs; and the effects of noise. Sound recording is an efficient and affordable tool, independent of daylight and weather; and recorders may be left in place for many months at a time, continuously collecting data on animals and their environment. This book builds the skills and knowledge necessary to collect and interpret acoustic data from terrestrial and marine environments. Beginning with a history of sound recording, the chapters provide an overview of off-the-shelf recording equipment and analysis tools (including automated signal detectors and statistical methods); audiometric methods; acoustic terminology, quantities, and units; sound propagation in air and under water; soundscapes of terrestrial and marine habitats; animal acoustic and vibrational communication; echolocation; and the effects of noise. This book will be useful to students and researchers of animal ecology who wish to add acoustics to their toolbox, as well as to environmental managers in industry and government

Exploring Animal Behavior Through Sound: Volume 1

This open-access book empowers its readers to explore the acoustic world of animals. By listening to the sounds of nature, we can study animal behavior, distribution, and demographics; their habitat characteristics and needs; and the effects of noise. Sound recording is an efficient and affordable tool, independent of daylight and weather; and recorders may be left in place for many months at a time, continuously collecting data on animals and their environment. This book builds the skills and knowledge necessary to collect and interpret acoustic data from terrestrial and marine environments. Beginning with a history of sound recording, the chapters provide an overview of off-the-shelf recording equipment and analysis tools (including automated signal detectors and statistical methods); audiometric methods; acoustic terminology, quantities, and units; sound propagation in air and under water; soundscapes of terrestrial and marine habitats; animal acoustic and vibrational communication; echolocation; and the effects of noise. This book will be useful to students and researchers of animal ecology who wish to add acoustics to their toolbox, as well as to environmental managers in industry and government