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Biomedical subjects

J A Simmons

Publications and source records attributed to J A Simmons.

At least 19 recordsLinked to original sources

Steering by echolocation: a paradigm of ecological acoustics.

1. Flights of three big brown bats (Eptesicus fuscus) landing on a hand and catching a suspended mealworm were video analysed. 2. Results were consistent with the bats using the same basic control procedure in the quite different approach tasks--namely keeping tau (r) = kr and tau (a)/tau (r) = k alpha r. Here r is the current distance to the destination; alpha is the angle between the current direction of the destination and the goal direction of final approach (beta min); tau (r) = r/r, tau (alpha) = alpha/alpha; and kr, k alpha r are constants. 3. The bats were each quite consistent on a particular task (hand or mealworm) in the values they used for the control parameters kr, k alpha r and beta min. However, different values were used in the two tasks, which reflected the different behaviour required at the destination. Flights to hand required twisting and landing upside down and approach angle beta min was closer to vertical and kr was smaller and corresponded to decelerating nearly to a stop. In contrast, the mealworms were caught in mid flight and approach angle beta min was shallower and speed of approach was about constant. 4. tau (r) might be registered acoustically by tau (echo-delay) or by tau (echo-intensity). tau (alpha) might be registered by the bat's directional hearing and gravity sense. 5. The bat's learned the tasks easily, suggesting that the control procedure they used in the experiments was part and parcel of the natural skills they had developed in the wild.

Acoustics

Spatially dependent acoustic cues generated by the external ear of the big brown bat, Eptesicus fuscus.

To measure the directionality of the external ear of the echolocating bat, Eptesicus fuscus, the left or right eardrum of a dead bat was replaced by a microphone which recorded signals received from a sound source that was moved around the stationary head. The test signal was a 0.5-ms FM sweep from 100 kHz to 10 kHz (covering all frequencies in the bat's biosonar sounds). Notches and peaks in transfer functions for 7 tested ears varied systematically with changes in elevation. For the most prominent notch, center frequency decreased from about 50 kHz for elevations at or near the horizontal to 30-40 kHz for elevations 30 degrees-40 degrees below the horizontal. A second notch shifted from about 85 kHz to 70 kHz over these same elevations. Above the horizontal, a peak that flanks these notches changed in amplitude by 15 dB with changes in elevation. Removal of the tragus from the external ear disrupted the systematic movement of notch frequencies with elevation but did not disrupt changes in the peak's amplitude. Smaller changes in notch frequency also occurred with changes in azimuth, so monaural notch information alone cannot determine the position of sound sources away from the median plane. However, because bats routinely keep the head pointed at the target's azimuth, median-plane localization occurs with monaural cues delivered to the two ears. Corresponding changes with elevation occurred in the impulse-response, which consists of a series of 3-6 peaks spaced 10-20 microseconds apart. The time separation of two prominent impulse peaks systematically increased from 22-26 microseconds above the horizontal to about 36-40 microseconds below the horizontal, and removal of the tragus disrupted this time shift below the horizontal.

Animals

Jamming bat echolocation: the dogbane tiger moth Cycnia tenera times its clicks to the terminal attack calls of the big brown bat Eptesicus fuscus.

Certain tiger moths emit high-frequency clicks to an attacking bat, causing it to break off its pursuit. The sounds may either orient the bat by providing it with information that it uses to make an attack decision (aposematism) or they may disorient the bat by interrupting the normal flow of echo information required to complete a successful capture (startle, jamming). At what point during a bat's attack does an arctiid emit its clicks? If the sounds are aposematic, the moth should emit them early in the attack echolocation sequence in order to allow the bat time to understand their meaning. If, however, the sounds disrupt the bat's echo-processing behaviour, one would expect them to be emitted later in the attack to maximize their confusion effects. To test this, we exposed dogbane tiger moths (Cycnia tenera) to a recording of the echolocation sequence emitted by a big brown bat (Eptesicus fuscus) as it attacked a stationary target. Our results demonstrate that, at normal echolocation intensities, C. tenera does not respond to approach calls but waits until the terminal phase of the attack before emitting its clicks. This timing is evident whether the moth is stationary or flying and is largely independent of the intensity of the echolocation calls. These results support the hypothesis of a jamming effect (e.g. 'phantom echoes') and suggest that, to determine experimentally the effects of arctiid clicks on bats, it is important that the bats be tested under conditions that simulate the natural context in which this defence operates.

Acoustics

Experimental calculations of droplet diffusion in a low-pressure cloud chamber.

A low-pressure cloud chamber was used for several years to display the tracks created by the passage of ionizing particles through vapors of interest. The spatial distributions of the ions that were formed were of special interest, but the accuracy with which these distributions could be determined was reduced by the presence of diffusion. This meant that the droplets, when photographed, had moved significantly away from the point of creation of the parent ion. In the present investigation photographs obtained by previous workers have been analyzed in an attempt to quantify the extent to which the droplets had diffused. The results suggest that the diffusion, when converted to standard density (1000 kg/m3), was independent of the pressure inside the cloud chamber and the mixture used. It could be represented by a one-dimensional root-mean-square diffusion distance whose value was calculated to be 2.42 +/- 0.04 nm. Values for the diffusion of thermalized electrons (< approximately 4 eV) before capture to form negative ions were also calculated. They appeared to lie in the range 3.5-5.0 nm, and were again independent of the pressure and nature of the mixture. The magnitude of the diffusion was large enough to mask any measurable prediffusion structure for a distance in the region of 10 nm radially around the track path of the alpha-particle and proton tracks analyzed.

Alpha Particles

Sonar gain control and echo detection thresholds in the echolocating bat, Eptesicus fuscus.

The echolocating bat, Eptesicus fuscus, detects sonar echoes with a sensitivity that changes according to the time elapsed between broadcasting of each sonar signal and reception of echoes. When tested in an electronic target simulator on a two-choice echo-detection task, the bat's threshold improved by 11.5 dB as echo delay changed from 2.3 to 4.6 ms (target ranges of 40 and 80 cm). Earlier experiments measured the change in detection threshold for delays from 1 to 6.4 ms (target ranges from about 17 to 110 cm) and obtained about 11 dB of improvement per doubling of delay. The new experiments used electronic delay lines to simulate echo delay, thus avoiding movement of loudspeakers to different distances and the possible creation of delay-dependent backward masking between stimulus echoes and cluttering echoes from the loudspeaker surfaces. The slope of the threshold shift defines an echo gain control that keeps echoes from point targets at a fixed sensation level--reducing sensitivity by 11 to 12 dB as echo amplitude increases by 12 dB per halving of range during the bat's approach to the target. A recent experiment using loudness discrimination of echoes at 70 to 80 dB SPL (roughly 50 dB above threshold) found a slope of about 6 dB per halving of range, so the gain-control effect may be level dependent. The observed effect is operationally equivalent to forward masking of echoes by the transmission, but any events correlated with vocalization which impair hearing sensitivity for a short interval following transmissions could cause a decline in sensitivity to echoes. Contractions of the bat's middle-ear muscles synchronized to transmissions may account for the observed threshold shift, at least for a span of echo delays associated with the most critical portion of the approach stage of pursuit. Forward masking by the sonar transmissions may contribute to the threshold shift, too, but middle-ear muscle contractions do occur and must be a significant part of the cause.

Animals

Cell migration in the rat embryonic neocortex.

Three-dimensional reconstructions of the normal rat embryonic (E) neocortex on days E15, E17, E19, and E21, using Skandha (software designed by J. Prothero, University of Washington, Seattle), show that the neocortical ventricular zone shrinks rapidly in the medial direction during cortical morphogenesis. [3H]thymidine autoradiography indicates that the shrinkage of the ventricular zone occurs before neurons in lateral and ventrolateral parts of layers IV-II are generated. Consequently, most of these neurons originate 400-1000 microns medial to their settling sites in the cortical plate. Embryos killed at daily intervals up to E21 after a single injection of [3H]thymidine on either E17 or E18 revealed the presence of a prominent migratory path, the lateral cortical stream, used by neurons migrating to the lateral and ventrolateral cortical plate; neurons migrating to the dorsal cortical plate follow a direct radial path. Arrival times of neurons in the cortical plate depend on the migratory path and are proportional to the overall distance travelled. Neurons that migrate only radially arrive in the dorsal cortical plate in two days (shortest route). Neurons that migrate laterally arrive in the lateral cortical plate in 3 days (longer route) and in the ventrolateral cortical plate in 4 days (longest route). [3H]thymidine autoradiography also shows that cells generated in the neocortical ventricular zone migrate in the lateral cortical stream for 5 or more days and accumulate in a reservoir. Cells leave the reservoir to enter the piriform cortex and destinations (as yet undetermined) in the basal telencephalon. The lateral cortical stream is found wherever the neocortical primordium surrounds the basal ganglia and is absent behind the basal ganglia. A computer analysis of nuclear orientation in anterior and posterior parts of the intermediate zone in the dorsal neocortex between days E17 and E22 shows that horizontally oriented nuclei are more common anteriorly where many cells are migrating laterally than posteriorly where most cells are migrating radially.

Animals

Convergence of temporal and spectral information into acoustic images of complex sonar targets perceived by the echolocating bat, Eptesicus fuscus.

1. FM echolocating bats (Eptesicus fuscus) were trained to discriminate between a two-component complex target and a one-component simple target simulated by electronically-returned echoes in a series of experiments that explore the composition of the image of the two-component target. In Experiment I, echoes for each target were presented sequentially, and the bats had to compare a stored image of one target with that of the other. The bats made errors when the range of the simple target corresponded to the range of either glint in the complex target, indicating that some trace of the parts of one image interfered with perception of the other image. In Experiment II, echoes were presented simultaneously as well as sequentially, permitting direct masking of echoes from one target to the other. Changes in echo amplitude produced shifts in apparent range whose pattern depended upon the mode of echo presentation. 2. Eptesicus perceives images of complex sonar targets that explicitly represent the location and spacing of discrete glints located at different ranges. The bat perceives the target's structure in terms of its range profile along a psychological range axis using a combination of echo delay and echo spectral representations that together resemble a spectrogram of the FM echoes. The image itself is expressed entirely along a range scale that is defined with reference to echo delay. Spectral information contributes to the image by providing estimates of the range separation of glints, but it is transformed into these estimates. 3. Perceived absolute range is encoded by the timing of neural discharges and is vulnerable to shifts caused by neural amplitude-latency trading, which was estimated at 13 to 18 microseconds per dB from N1 and N4 auditory evoked potentials in Eptesicus. Spectral cues representing the separation of glints within the target are transformed into estimates of delay separations before being incorporated into the image. However, because they are encoded by neural frequency tuning rather than the time-of-occurrence of neural discharges, the perceived range separation of glints in images is not vulnerable to amplitude-latency shifts. 4. The bat perceives an image that is displayed in the domain of time or range. The image receives no evident spectral contribution beyond what is transformed into delay estimates. Although the initial auditory representation of FM echoes is spectrogram-like, the time, frequency, and amplitude dimensions of the spectrogram appear to be compressed into an image that has only time and amplitude dimensions.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation

Discrimination of jittered sonar echoes by the echolocating bat, Eptesicus fuscus: the shape of target images in echolocation.

1. Behavioral experiments with jittering echoes examined acoustic images of sonar targets in the echolocating bat, Eptesicus fuscus, along the echo delay or target range axis. Echo phase, amplitude, bandwidth, and signal-to-noise ratio were manipulated to assess the underlying auditory processes for image formation. 2. Fine delay acuity is about 10 ns. Calibration and control procedures indicate that this represents temporal acuity rather than spectral discrimination. Jitter discrimination curves change in phase when the phase of one jittering echo is shifted by 180 degrees relative to the other, showing that echo phase is involved in delay estimation. At an echo detectability index of about 36 dB, fine acuity is 40 ns, which is approximately as predicted for the delay accuracy of an ideal receiver. 3. Compound performance curves for 0 degrees and 180 degrees phase conditions match the crosscorrelation function of the echoes. The locations of both 0 degrees and 180 degrees phase peaks in the performance curves shift along the time axis by an amount that matches neural amplitude-latency trading in Eptesicus, confirming a temporal basis for jitter discrimination.

Acoustic Stimulation

A view of the world through the bat's ear: the formation of acoustic images in echolocation.

Echolocating bats perceive objects as acoustic images derived from echoes of the ultrasonic sounds they emit. They can detect, track, identify, and intercept flying insects using sonar. Many species, such as the big brown bat, Eptesicus fuscus, emit frequency-modulated sonar sounds and perceive the distance to targets, or target range, from the delay of echoes. For Eptesicus, a point-target's image has a sharpness along the range axis that is determined by the acuity of echo-delay perception, which is about 10 ns under favorable conditions. The image as a whole has a fine range structure that corresponds to the cross-correlation function between emissions and echoes. A complex target- which has reflecting points, called "glints", located at slightly different distances and reflects echoes containing overlapping components with slightly different delays--is perceived in terms of its range profile. The separation of the glints along the range dimension is encoded by the shape of the echo spectrum created by interference between overlapping echo components. However, Eptesicus transforms the echo spectrum back into an estimate of the original delay separation of echo components. The bat thus converts spectral cues into elements of an image expressed in terms of range. The absolute range of the nearest glint is encoded by the arrival time of the earliest echo component, and the spectrally encoded range separation of additional glints is referred to this time-encoded reference range for the image as a whole. Each individual glint is represented by a cross-correlation function for its own echo component, the nearest of which is computed directly from arrival-time measurements while further ones are computed by transformation of the echo spectrum. The bat then sums the cross-correlation functions for multiple glints to form the entire image of the complex target. Range and shape are two distinct features of targets that are separately encoded by the bat's auditory system, but the bat perceives unitary images that require fusion of these features to create a synthetic psychological dimension of range. The bat's use of cross-correlation-like images reveals neural computations that achieve fusion of stimulus features and offers an example of high-level operations involved in the formation of perceptual "wholes".

Animals

Clutter interference and the integration time of echoes in the echolocating bat, Eptesicus fuscus.

The ability of the echolocating bat, Eptesicus fuscus, to detect a sonar target is affected by the presence of other targets along the same axis at slightly different ranges. If echoes from one target arrive at about the same delay as echoes from another target, clutter interference occurs and one set of echoes masks the other. Although the bat's sonar emissions and the echoes themselves are 2 to 5 ms long, echoes (of approximately equal sensation levels--around 15 dB SL) only interfere with each other if they arrive within 200 to 400 microseconds of the same arrival time. This figure is an estimate of the integration time of the bat's sonar receiver for echoes. The fine structure of the clutter-interference data reflects the reinforcement and cancellation of echoes according to their time separation. When clutter interference first occurs, the waveforms of test and cluttering echoes already overlap for much of their duration. The masking effect underlying clutter interference appears specifically due to overlap, not between raw echo waveforms, but between the patterns of mechanical excitation created when echoes pass through bandpass filters equivalent to auditory-nerve tuning curves. While the time scale of clutter interference is substantially shorter than the duration of echo waveforms, it still is much longer than the eventual width of a target's range-axis image expressed in terms of echo delay.

Animals

The acoustic basis for target discrimination by FM echolocating bats.

Past experiments show that echolocating bats of the species Myotis lucifugus and Eptesicus fuscus can discriminate among airborne sonar targets presented in the context of pursuit maneuvers for the interception of prey. These bats distinguish between edible mealworms and inedible spheres of various sizes. Myotis can distinguish between disks and mealworms similar enough in size that the bat's performance requires the ability to perceive the acoustic equivalent of target shape. Previously observed small differences in the spectrum of echoes from mealworms and disks appear insufficient to distinguish these targets at the performance levels achieved by bats. We measured the acoustic properties of the targets by broadcasting ultrasonic impulses at mealworms, spheres, and disks and recording their echoes, displaying the results in terms of impulse echo waveforms and the frequency response of targets derived from the target transfer function. The echoes from disks and mealworms at various orientations convey the range-axis profile of the target (number and spacing of reflecting points or glints distributed at different ranges) in terms of the impulse structure of their waveforms and in terms of the locations and spacing of notches or nulls in their spectra. For targets that bats can discriminate and that reflect echoes which do not clearly differ in overall amplitude, the targets appear distinguishable from the acoustic representation of their range profile, which is a feature of targets that bats can perceive with great acuity.

Animals

The volumes of rat and human lung cells as measured by an image analyser.

Previous studies of the morphometry of lung cells have required long and tedious measurements involving serial sectioning and the reconstruction of models. However the advent of computer-based image analysers offers a more straightforward, although less direct, method of making such measurements. In the present study a Magiscan has been used to investigate the shapes and sizes of the nuclei from the lung cells of three mammalian species. Associated cellular volumes have also been derived. Based on the measurements of about two hundred cells the mean volumes of rat, beagle and human lung cell nuclei were estimated to be 67 microns 3, 41 microns 3 and 78 microns 3 respectively. Cell volumes were estimated to be about 500 microns 3. It was also confirmed that a sphere is a very poor approximation for the shape of a nucleus, spheroids representing the observations much more closely.

Animals

Clutter interference along the target range axis in the echolocating bat, Eptesicus fuscus.

The sensitivity of the echolocating bat, Eptesicus fuscus, for detection of a sonar target is impaired by the presence of additional targets located at similar distances. At a range of 54 cm, sensitivity to one target declines if the range separation to other targets is smaller than 8-9 cm. This loss of sensitivity is an example of clutter interference along the range axis, created by simultaneous masking of one set of echoes by another. Echoes that fall within an experimentally defined critical range band may sum together to contribute collectively to detection in that band. Echoes falling into separate bands may be detected independently. Acoustic glints within a band appear to be grouped together to be perceived as a single range-extended target of complex structure. Range bands may thus define what a "target" is by specifying within-target and between-target differences in range. The width of critical range bands appears to depend upon target range, with wider bands at greater ranges.

Animals

Analysis of the dose-response relationships following the irradiation of amino acids.

A comparison is made between results recently reported on the induction of free radicals in irradiated alanine and studies previously published on irradiated amino acids. It is shown that there is good qualitative agreement between the old and the new measurements although several quantitative inconsistencies are noted. A query is raised as to whether the new data can truly be interpreted in terms of the theory of track structure when some of the effects observed are not fully taken into account.

Amino Acids

Sonar tracking of horizontally moving targets by the big brown bat Eptesicus fuscus.

When following a moving target, echolocating bats (Eptesicus fuscus) keep their heads aimed at the target's position. This tracking behavior seems not to involve predicting the target's trajectory, but is achieved by the bat's pointing its head at the target's last known position. The bat obtains frequent position updates by emitting sonar signals at a high rate. After the lag between head and target positions and the nonunity tracking gain were corrected for, bats' tracking accuracy in the horizontal plane was +/- 1.6 degree.

Animals

Microdosimetry of alpha-irradiated lung.

Measurements have been made on the fractions of tissue affected and distributions of energy deposited when rat and human lung are irradiated by alpha particles. Thin sections of alveolar tissue were examined in an image analyser and an imaginary 239PuO2 particle was placed in an alveolar sac. The hypothetical alpha particles were followed to the ends of their tracks so the effects of the material's real structure could be examined in detail. It is shown that a fair comparison exists with results previously obtained with hamster lung. In addition, the measurements actually made with human lung correlate well with predictions based on the rat lung measurements. Finally, it is confirmed that the approximation of a uniform, structureless lung is a poor one.

Alpha Particles