Principles of auditory information-processing derived from neuroethology

1989 ◽  
Vol 146 (1) ◽  
pp. 277-286 ◽  
Author(s):  
N. Suga
Keyword(s):  
Auditory System ◽  
Doppler Shift ◽  
Basilar Membrane ◽  
Neural Mechanisms ◽  
Target Distance ◽  
Target Velocity ◽  
Auditory Information ◽  
Constant Frequency ◽  
Mustached Bat ◽  
Velocity Information

For auditory imaging, a bat emits orientation sounds (pulses) and listens to echoes. The parameters characterizing a pulse-echo pair each convey particular types of biosonar information. For example, a Doppler shift (a difference in frequency between an emitted pulse and its echo) carries velocity information. For a 61-kHz sound, a 1.0-kHz Doppler shift corresponds to 2.8 ms-1 velocity. The delay of the echo from the pulse conveys distance (range) information. A 1.0-ms echo delay corresponds to a target distance of 17 cm. The auditory system of the mustached bat, Pteronotus parnelli, from Central America solves the computational problems in analyzing these parameters by creating maps in the cerebral cortex. The pulse of the mustached bat is complex. It consists of four harmonics, each of which contains a long constant-frequency (CF) component and a short frequency-modulated (FM) component. Therefore, there are eight components in the emitted pulse (CF1-4 and FM1-4). The CF signal is particularly suited for target velocity measurement, whereas the FM signal is suited for target distance measurement. Since the eight components differ from each other in frequency, they are analyzed in parallel at different regions of the basilar membrane in the inner ear. Then, they are separately coded by primary auditory neurons and are sent up to the auditory cortex through several auditory nuclei. During the ascent of the signals through these auditory nuclei, neurons responding to the FM components process range information, while other neurons responding to the CF components process velocity information. A comparison of the data obtained from the mustached bat with those obtained from other species illustrates both the specialized neural mechanisms specific to the bat's auditory system, and the general neural mechanisms which are probably shared with many different types of animals.

Inactivation of the DSCF area of the auditory cortex with muscimol disrupts frequency discrimination in the mustached bat

1992 ◽  
Vol 68 (5) ◽  
pp. 1613-1623 ◽  
Author(s):  
H. Riquimaroux ◽  
S. J. Gaioni ◽  
N. Suga
Keyword(s):  
Auditory Cortex ◽  
Doppler Shift ◽  
Broad Band ◽  
Primary Auditory Cortex ◽  
Tone Burst ◽  
Signal Pulse ◽  
Gamma Aminobutyric Acid ◽  
Constant Frequency ◽  
Mustached Bat ◽  
Leg Flexion

1. The Jamaican mustached bat uses a biosonar signal (pulse) with eight major components: four harmonics each consisting of a long constant frequency (CF1-4) component followed by a short frequency-modulated (FM1-4) component. While flying, the bat adjusts the frequency of its pulse so as to maintain the CF2 of the Doppler-shifted echo at a frequency to which its cochlea is very sharply tuned. This Doppler shift (DS) compensation likely is mediated or influenced by the Doppler-shifted CF (DSCF) processing area of the primary auditory cortex, which only represents frequencies in the range of echo CF2s (60.6 to 62.3 kHz when the "resting" frequency of the CF2 is 61.0 kHz). 2. We trained four bats to discriminate between different trains of paired tone bursts that mimicked a bat's pulse CF2 and the accompanying echo CF2. The frequency of these CF2s ranged between 61.0 and 64.0 kHz. A discriminated shock avoidance procedure response was employed using a leg flexion. For one stimulus, the S+, the pulse and echo CF2s were the same frequency (delta f = 0, i.e., no Doppler shift). A leg flexion during the S+ turned off both the S+ and the scheduled shock. For a second stimulus, the S-, the echo CF2 was 0.05, 0.1, 0.3, 0.5, or 2.0 kHz higher than the pulse CF2. A delta f of 0.05 kHz was a frequency difference of 0.08%. No shock followed the S-, and leg flexions had no consequences. Correct responses consisted of a leg flexion during the S+ and no flexion during the S-; these responses were added together to compute the percentage of correct responses. When a bat correctly responded at better than 75% for all the delta f s, muscimol, a potent agonist of gamma-aminobutyric acid, was bilaterally applied to inactivate the DSCF area. Performance on each delta f discrimination was then measured. 3. Initial attempts to condition the bats to flex their legs to the CF tones mimicking part of the natural pulses and echoes failed. When broad-band noise bursts were substituted, however, the conditioned response was rapidly established. The noise band-width was gradually reduced and then replaced with the CF tones. Discrimination training with the tone burst trains then commenced. Throughout this procedure, the bats maintained their responding to the stimuli. The bats typically required approximately 20-30 sessions to perform consistently (> or = 75% correct responses) a discrimination involving a 2 kHz delta f.(ABSTRACT TRUNCATED AT 400 WORDS)

Doppler-shift compensation by the mustached bat: quantitative data.

1994 ◽  
Vol 188 (1) ◽  
pp. 115-129 ◽  
Author(s):  
A W Keating ◽  
O W Henson ◽  
M M Henson ◽  
W C Lancaster ◽  
D H Xie
Keyword(s):  
Doppler Shift ◽  
Quantitative Data ◽  
Second Harmonic ◽  
Frequency Component ◽  
Reference Frequency ◽  
Constant Frequency ◽  
Doppler Shifts ◽  
Mustached Bat ◽  
Pteronotus Parnellii ◽  
Significant Difference

Quantitative data for Doppler-shift compensation by Pteronotus parnellii parnellii were obtained with a device which propelled the bats at constant velocities over a distance of 12 m. The bats compensated for Doppler shifts at all velocities tested (0.1-5.0 ms-1). The main findings were (1) that compensation was usually accomplished by a progressive lowering of the approximately 61 kHz second harmonic constant-frequency component of emitted sounds in small frequency steps (93 +/- 72 Hz); (2) that the time needed to reach a steady compensation level averaged 514 +/- 230 ms and the number of pulses required to reach full compensation averaged 10.78 +/- 5.16; (3) that the animals compensated to hold the echo (reference) frequency at a value that was slightly higher than the resting frequency and slightly lower than the cochlear resonance frequency; (4) that reference frequency varied as a function of velocity, the higher the velocity of the animal, the higher was the reference frequency (slope 55 Hz m-1s-2); and (5) that the mean reference frequency was always an undercompensation. The average amount of undercompensation was 15.8%. There was a significant difference (P < or = 0.005) in Doppler-shift compensation data collected at velocities that differed by 0.1 ms-1. A velocity difference of 0.1 ms-1 corresponds to a Doppler-shift difference of about 35 Hz in the approximately 61 kHz signals reaching the ear.

Combination-sensitive neurons in the medial geniculate body of the mustached bat: encoding of relative velocity information

1991 ◽  
Vol 65 (6) ◽  
pp. 1254-1274 ◽  
Author(s):  
J. F. Olsen ◽  
N. Suga
Keyword(s):  
Doppler Shift ◽  
Cortical Neurons ◽  
Medial Geniculate Body ◽  
Distance Information ◽  
Tuning Curves ◽  
Mustached Bat ◽  
Medial Geniculate ◽  
Velocity Information ◽  
Pulse Echo ◽  
Echo Delay

1. Orientation sounds (pulses) emitted by the mustached bat (Pteronotus parnellii) consist of up to four harmonics (H1-4); each harmonic contains a constant frequency (CF) component and a terminal frequency modulated (FM) component, so that there are eight components in total (CF1-4 and FM1-4). By referring the echo from a target to the emitted pulse, the mustached bat derives velocity information from Doppler shift and distance information from echo delay. In this study, the responses of single neurons in the medial geniculate body (MGB) to synthetic biosonar signals were investigated. Stimuli consisted of CF, FM, and CF-FM sounds. Paired CF-FM sounds were used to mimic any two harmonics of pulse-echo pairs. The dorsal and medial divisions of the MGB were found to contain combination-sensitive neurons. These neurons responded poorly to individual sounds regardless of frequency and amplitude and were facilitated by paired sounds presented at particular frequencies, amplitudes and inter-component intervals (simulated echo delay). Combination-sensitive neurons were tuned to the frequencies that characterize particular components of natural biosonar signals and were classified according to the components of pulse-echo pairs that best matched the spectral selectivity of the neuron. Two classes of combination-sensitive neurons were found, CF/CF and FM-FM. This paper focuses on CF/CF combination-sensitive neurons, which extract velocity information from paired CF components, and on CF2 and CF3 neurons, which, although not combination-sensitive, are tuned to the frequencies of the CF2 and CF3 components of biosonar signals. 2. CF2 and CF3 neurons were sharply tuned in frequency. The best frequencies of the most sharply tuned CF2 neurons were all approximately equal to 61.17 kHz (SD = 370 Hz), which closely matches the frequency at which P. parnellii stabilizes the CF2 component of an echo when compensating for Doppler shift. Thus CF2 neurons are specialized for a fine analysis of Doppler-compensated echoes. 3. Tuning curves of CF2 and CF3 neurons remained narrow regardless of stimulus level. When compared at high stimulus levels (30 and 50 dB above minimum threshold), bandwidths of tuning curves of CF2 and CF3 neurons were much smaller than those of peripheral auditory neurons turned to CF2 or CF3 frequencies but were about the same as those of cortical neurons tuned to CF2 or CF3 frequencies. Thus the sharpening of neural tuning curves by the bat's central auditory system occurs within or before the MGB.(ABSTRACT TRUNCATED AT 400 WORDS)

Development of Echolocation Calls in the Mustached Bat, Pteronotus parnellii

2003 ◽  
Vol 90 (4) ◽  
pp. 2274-2290 ◽  
Author(s):  
M. Vater ◽  
M. Kössl ◽  
E. Foeller ◽  
F. Coro ◽  
E. Mora ◽  
...  
Keyword(s):  
Doppler Shift ◽  
Body Motion ◽  
Constant Frequency ◽  
Echolocation Calls ◽  
A Value ◽  
Mustached Bat ◽  
Pteronotus Parnellii ◽  
Age Related ◽  
Fm Signals ◽  
Frequency Modulations

Adult mustached bats employ Doppler-sensitive sonar to hunt fluttering prey insects in acoustically cluttered habitats. The echolocation call consists of 4–5 harmonics, each composed of a long constant frequency (CF) component flanked by brief frequency modulations (FM). The 2nd harmonic CF component (CF2) at 61 kHz is the most intense, and analyzed by an exceptionally sharply tuned auditory system. The maturation of echolocation calls and the development of Doppler-shift compensation was studied in Cuba where large maternity colonies are found in hot caves. In the 1st postnatal week, infant bats did not echolocate spontaneously but could be induced to vocalize CF-FM signals by passive body motion. The CF2 frequency emitted by the smallest specimens was at 48 kHz (i.e., 0.4 octaves lower than the adult signal). CF-FM signals were spontaneously produced in the 2nd postnatal week at a CF2 frequency of 52 kHz. The CF2 frequencies of induced and spontaneous calls shifted upward to reach a value of 60.5 kHz in the 5th postnatal week. Standard deviations of CF2 frequency were large (up to ±1.5 kHz) in the youngest bats and dropped to values of ±250 Hz at the end of the 3rd postnatal week. Some individuals in the 4th and 5th postnatal weeks emitted with adultlike frequency precision of about ±100 Hz. In the youngest bats, the 1st harmonic CF component (CF1) was up to 22 dB stronger than CF2. Adultlike relative levels of CF1 (–28 dB relative to CF2) were reached in the 5th postnatal week. In spontaneously emitted CF-FM calls, the duration of the CF2 component gradually increased with age from 5 ms to maximum values of 18 ms. Durations of the CF2 component in induced calls averaged 7 ± 2.6 ms in the 1st postnatal week and 8.2 ± 1.5 ms in the 5th postnatal week. There were no age-related changes in duration of the terminal FM sweep (3 ± 0.4 ms) in both induced and spontaneous calls. The magnitude of the terminal FM sweep in spontaneous calls was not correlated with age (mean 13.5 ± 2 kHz). Values for induced calls slightly increased with age from 11 ± 2 to 13 ± 2 kHz. The emission rate of induced CF-FM signals increased with age from values of 2.5 ± 2 to 17 ± 5 pulses/s. Values for spontaneously emitted calls were 4.4 ± 3 and 9 ± 4.5 pulses/s, respectively. Doppler-shift compensation, as tested in the pendulum task, emerged during the 4th postnatal week in young bats that were capable of very brief active flights, but before the time of active foraging outside the cave.

Neuromodulatory Feedback to the Inferior Colliculus

2018 ◽  
pp. 576-610 ◽  
Author(s):  
Laura Hurley
Keyword(s):  
Inferior Colliculus ◽  
Auditory System ◽  
Auditory Processing ◽  
Internal State ◽  
Brain Regions ◽  
Auditory Information ◽  
Situational Variables ◽  
Acoustic Stimuli ◽  
Functionally Diverse ◽  
Behavioral Information

The inferior colliculus (IC) receives prominent projections from centralized neuromodulatory systems. These systems include extra-auditory clusters of cholinergic, dopaminergic, noradrenergic, and serotonergic neurons. Although these modulatory sites are not explicitly part of the auditory system, they receive projections from primary auditory regions and are responsive to acoustic stimuli. This bidirectional influence suggests the existence of auditory-modulatory feedback loops. A characteristic of neuromodulatory centers is that they integrate inputs from anatomically widespread and functionally diverse sets of brain regions. This connectivity gives neuromodulatory systems the potential to import information into the auditory system on situational variables that accompany acoustic stimuli, such as context, internal state, or experience. Once released, neuromodulators functionally reconfigure auditory circuitry through a variety of receptors expressed by auditory neurons. In addition to shaping ascending auditory information, neuromodulation within the IC influences behaviors that arise subcortically, such as prepulse inhibition of the startle response. Neuromodulatory systems therefore provide a route for integrative behavioral information to access auditory processing from its earliest levels.

A neural model of medial geniculate body and auditory cortex detecting target distance independently of target velocity in echolocation

2000 ◽  
Vol 32-33 ◽  
pp. 833-841 ◽  
Author(s):  
Satoru Inoue ◽  
Manabu Kimyou ◽  
Yoshiki Kashimori ◽  
Osamu Hoshino ◽  
Takeshi Kambara
Keyword(s):  
Auditory Cortex ◽  
Medial Geniculate Body ◽  
Neural Model ◽  
Target Distance ◽  
Target Velocity ◽  
Medial Geniculate

Damage of the Auditory System Associated with Acute Blast Trauma

1989 ◽  
Vol 98 (5_suppl) ◽  
pp. 23-34 ◽  
Author(s):  
Michele Roberto ◽  
Roger P. Hamernik ◽  
George A. Turrentine
Keyword(s):  
Auditory System ◽  
Sound Pressure ◽  
Basilar Membrane ◽  
Organ Of Corti ◽  
Blast Waves ◽  
Mechanical Fracture ◽  
Sound Pressure Levels ◽  
Temporal Bones ◽  
Blast Trauma ◽  
The Impact

This paper reviews the results of several studies on the effects of blast wave exposure on the auditory system of the chinchilla, the pig, and the sheep. The chinchillas were exposed at peak sound pressure levels of approximately 160 dB under well-controlled laboratory conditions. A modified shock tube was used to generate the blast waves. The pigs and sheep were exposed under field conditions in an instrumented hard-walled enclosure. Blast trauma was induced by the impact of a single explosive projectile. The peak sound pressure levels varied between 178 and 209 dB. All animals were killed immediately following exposure, and their temporal bones were removed for fixation and histologic analysis using light microscopy and scanning electron microscopy. Middle ears were examined visually for damage to the conductive system. There were well-defined differences in susceptibility to acoustic trauma among species. However, common findings in each species were the acute mechanical fracture and separation of the organ of Corti from the basilar membrane, and tympanic membrane and ossicular failure.

Are the initial frequency-modulated components of the mustached bat's biosonar pulses important for ranging?

1991 ◽  
Vol 66 (6) ◽  
pp. 1951-1964 ◽  
Author(s):  
D. C. Fitzpatrick ◽  
N. Suga ◽  
H. Misawa
Keyword(s):  
Maximum Amplitude ◽  
Signal Amplitude ◽  
Frequency Component ◽  
Target Range ◽  
Constant Frequency ◽  
Range Resolution ◽  
Mustached Bat ◽  
Pteronotus Parnellii ◽  
Target Ranging ◽  
The Mean

1. FM-FM neurons in the auditory cortex of the mustached bat, Pteronotus parnellii, are specialized to process target range. They respond when the terminal frequency-modulated component (TFM) of a biosonar pulse is paired with the TFM of the echo at a particular echo delay. Recently, it has been suggested that the initial FM components (IFMs) of biosonar signals may also be important for target ranging. To examine the possible role of IFMs in target ranging, we characterized the properties of IFMs and TFMs in biosonar pulses emitted by bats swung on a pendulum. We then studied responses of FM-FM neurons to synthesized biosonar signals containing IFMs and TFMs. 2. The mustached bat's biosonar signal consists of four harmonics, of which the second (H2) is the most intense. Each harmonic has an IFM in addition to a constant-frequency component (CF) and a TFM. Therefore each pulse potentially consists of 12 components, IFM1-4, CF1-4, and TFM1-4. The IFM sweeps up while the TFM sweeps down. 3. The IFM2 and TFM2 depths (i.e., bandwidths) were measured in 217 pulses from four animals. The mean IFM2 depth was much smaller than the mean TFM2 depth, 2.87 +/- 1.52 (SD) kHz compared with 16.27 +/- 1.08 kHz, respectively. The amplitude of the IFM2 continuously increased throughout its duration and was always less than the CF2 amplitude, whereas the TFM2 was relatively constant in amplitude over approximately three-quarters of its duration and was often the most intense part of the pulse. The maximum amplitude of the IFM2 was, on average, 11 dB smaller than that of the TFM2. Because range resolution increases with depth and the maximum detectable range increases with signal amplitude, the IFMs are poorly suited for ranging compared with the TFMs. 4. FM-FM neurons (n = 77) did not respond or responded very poorly to IFMs with depths and intensities similar to those emitted on the pendulum. The mean IFM2 depth at which a just-noticeable response appeared was 4.48 +/- 1.98 kHz. Only 14% of the pulses emitted on the pendulum had IFM2 depths that exceeded the mean IFM2 depth threshold of FM-FM neurons. 5. Most FM-FM neurons responded to IFMs that had depths comparable with those of TFMs. However, when all parameters were adjusted to optimize the response to TFMs and then readjusted to maximize the response to IFMs, 52% of 27 neurons tested responded significantly better to the optimal TFMs than to the optimal IFMs (P less than 0.05, t test).(ABSTRACT TRUNCATED AT 400 WORDS)

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