Biosonar behavior of mustached bats swung on a pendulum prior to cortical ablation

1990 ◽  
Vol 64 (6) ◽  
pp. 1801-1817 ◽  
Author(s):  
S. J. Gaioni ◽  
H. Riquimaroux ◽  
N. Suga
Keyword(s):  
Pulse Duration ◽  
Auditory Processing ◽  
Target Identification ◽  
Frequency Component ◽  
Signal Pulse ◽  
Echo Intensity ◽  
Constant Frequency ◽  
Large Target ◽  
Pulse Intensity ◽  
Pendulum Swing

1. The biosonar signal (pulse) of the mustached bat, Pteronotus parnellii parnellii, has four harmonics (H1-4), each consisting of a long constant-frequency component (CF1-4) followed by a short frequency-modulated component (FM1-4). As the bat approaches a target, it systematically modifies its pulses to optimize the extraction of information from the echoes. These behavioral responses include 1) Doppler-shift (DS) compensation in which the bat adjusts the frequency of its pulses to correct for the DS in the echoes. This maintains the echo CF2 at a frequency to which the bat's cochlea is very sharply tuned, slightly above the CF2 frequency of the bat's pulses when it is at rest (Frest, approximately 61 kHz); 2) echo intensity compensation, in which the bat lowers its pulse intensity as it approaches a large target, thus maintaining the echo intensity within a suitable range for auditory processing; and 3) and 4) duration and rate adjustments, in which the bat first increases its pulse duration to facilitate target identification, then shortens its pulse duration while increasing its pulse rate to facilitate target analysis. 2. We examined these responses, especially DS compensation, by swinging bats on a pendulum toward a large target over a distance of 3.6 m. Eight bats were given 15-30 swings per day for 6-25 days. 1) On 97% of all swings the bats showed strong DS compensation as the pendulum approached the target. They did not show DS compensation on the backswing. 2) On 40-50% of all swings, the bats clearly displayed the other responses. The bats typically increased their pulse intensity a small amount early in the pendulum swing, then decreased pulse intensity by as much as 18 dB as the target was more closely approached. They increased their pulse intensity during the backswing. 3) Pulse duration increased from approximately 20 to 23 ms early in the forward swing, decreased to approximately 18 ms as the target was more closely approached, and then increased to 20 ms by the end of the backswing. 4) The instantaneous repetition rate increased from approximately 17 pulses/s at the start of the forward swing to approximately 28 pulses/s near the target, then decreased to approximately 10 pulses/s by the end of the backswing. Pulses usually occurred in trains of 1-2 pulses, with longer trains occasionally occurring near the target. 3. The maximum DS on the pendulum was 1.34 kHz, and the maximum DS compensation was 146 +/- 98 (SD) Hz less than this value.(ABSTRACT TRUNCATED AT 400 WORDS)

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)

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)

Generation of intense spectral continuum and isolated attosecond pulse by selecting single harmonic emission peak

2019 ◽  
Vol 33 (35) ◽  
pp. 1950444
Author(s):  
Liqiang Feng ◽  
Yi Li
Keyword(s):  
Pulse Duration ◽  
Laser Intensity ◽  
Attosecond Pulse ◽  
Resonance Ionization ◽  
Lower Intensity ◽  
Harmonic Intensity ◽  
Chirped Pulse ◽  
Harmonic Emission ◽  
Pulse Intensity ◽  
Isolated Attosecond Pulse

Generally, due to the interference of different harmonic emission peaks (HEPs), the intensity of high-order harmonic spectrum cannot keep enhancing as the pulse intensity increases. Thus, in this paper, a potential method to obtain an intense spectral continuum and isolated attosecond pulse (IAP) by selecting single HEP has been theoretically investigated. First, we choose the harmonic cutoff and the harmonic intensity from the optimal single-color laser intensity as the referenced values. Next, by properly choosing a lower-intensity negative chirped pulse, we see that the harmonic cutoff from this field is similar as that from the referenced field. Moreover, the spectral continuum is contributed by single HEP. However, the intensity of the spectral continuum from the negative chirped pulse is lower than that from the referenced field. As the pulse duration of the chirped pulse increases, the similar harmonic cutoff can also be found by using a much lower-intensity negative chirped pulse. However, the intensity of the spectral continuum is decreased compared with that from the shorter chirped pulse duration. Further, with the introduction of an IR or UV controlling pulse, the intensity of the spectral continuum can be enhanced up to the referenced value. Moreover, the longer the pulse duration of the controlling pulse is used, the lower the controlling pulse intensity is needed. Also, due to the UV resonance ionization, much lower UV intensity is needed to enhance the harmonic yield compared with that for adding IR controlling pulse case. All in all, the total laser intensity of the combined field (the fundamental pulse + the controlling pulse) is lower than that of the referenced field. Most importantly, the signal of the spectral continuum is only coming from single HEP, which can support the generations of intense IAPs with the durations of 30 as.

Frequency and amplitude representations in anterior primary auditory cortex of the mustached bat

1983 ◽  
Vol 50 (5) ◽  
pp. 1182-1196 ◽  
Author(s):  
A. Asanuma ◽  
D. Wong ◽  
N. Suga
Keyword(s):  
Auditory Cortex ◽  
Second Harmonic ◽  
Free Field ◽  
Primary Auditory Cortex ◽  
Frequency Component ◽  
Constant Frequency ◽  
Mustached Bat ◽  
Pteronotus Parnellii ◽  
Frequency Representation ◽  
Pure Tones

The orientation sound emitted by the Panamanian mustached bat, Pteronotus parnellii rubiginosus, consists of four harmonics. The third harmonic is 6-12 dB weaker than the predominant second harmonic and consists of a long constant-frequency component (CF3) at about 92 kHz and a short frequency-modulated component (FM3) sweeping from about 92 to 74 kHz. Our primary aim is to examine how CF3 and FM3 are represented in a region of the primary auditory cortex anterior to the Doppler-shifted constant-frequency (DSCF) area. Extracellular recordings of neuronal responses from the unanesthetized animal were obtained during free-field stimulation of the ears with pure tones. FM sounds, and signals simulating their orientation sounds and echoes. Response properties of neurons and tonotopic and amplitopic representations were examined in the primary and the anteroventral nonprimary auditory cortex. In the anterior primary auditory cortex, neurons responded strongly to single pure tones but showed no facilitative responses to paired stimuli. Neurons with best frequencies from 110 to 90 kHz were tonotopically organized rostrocaudally, with higher frequencies located more rostrally. Neurons tuned to 92-94 kHz were overpresented, whereas neurons tuned to sound between 64 and 91 kHz were rarely found. Consequently a striking discontinuity in frequency representation from 91 to 64 kHz was found across the anterior DSCF border. Most neurons exhibited monotonic impulse-count functions and responded maximally to sound pressure level (SPL). There were also neurons that responded best to weak sounds but unlike the DSCF area, amplitopic representation was not found. Thus, the DSCF area is quite unique not only in its extensive representation of frequencies in the second harmonic CF component but also in its amplitopic representation. The anteroventral nonprimary auditory cortex consisted of neurons broadly tuned to pure tones between 88 and 99 kHz. Neither tonotopic nor amplitopic representation was observed. Caudal to this area and near the anteroventral border of the DSCF area, a small cluster of FM-FM neurons sensitive to particular echo delays was identified. The responses of these neurons fluctuated significantly during repetitive stimulation.

Delay lines and amplitude selectivity are created in subthalamic auditory nuclei: the brachium of the inferior colliculus of the mustached bat

1993 ◽  
Vol 69 (5) ◽  
pp. 1713-1724 ◽  
Author(s):  
N. Kuwabara ◽  
N. Suga
Keyword(s):  
Inferior Colliculus ◽  
Subthalamic Nucleus ◽  
Frequency Component ◽  
Nerve Fibers ◽  
Coincidence Detection ◽  
Delay Lines ◽  
Constant Frequency ◽  
Response Latencies ◽  
Mustached Bat ◽  
Species Specific

1. The biosonar pulse of the mustached bat, Pteronotus parnellii parnellii, consists of four harmonics of a constant-frequency component (CF1-4) followed by a frequency-modulated component (FM1-4). FM-FM combination-sensitive neurons in the auditory cortex and the medical geniculate body (MGB) show facilitative responses to certain combinations of FM components in a pulse-echo pair. They are tuned to particular delays of echo FMn (EFMn) (n = 2, 3, or 4) from pulse FM1 (PFM1). The neural mechanisms for creating their response properties involve delay lines, coincidence detection, and multiplication. Coincidence detection and multiplication take place in the MGB. It is not yet known where and how delay lines are created. The first aim of the present studies is to examine whether delay lines are created by subthalamic nuclei. FM-FM neurons are tuned to not only echo delays but also echo amplitudes. Therefore, the second aim of the present studies is to examine the extent to which amplitude selectivity is created by subthalamic nuclei. Responses of single nerve fibers to acoustic stimuli were recorded from the brachium of the inferior colliculus (BIC) using tungsten wire microelectrodes, and their response latencies and best amplitudes were measured. 2. All BIC fibers responded strongly to single tone bursts. No FM-FM combination-sensitive neurons were found in the BIC. The best frequencies of BIC fibers were predominantly within the frequency ranges of four harmonics of the species-specific biosonar pulse. 3. The response latencies of BIC fibers tuned to FM1 were more diverse (3.5-15.0 ms) than those of BIC fibers tuned to FMn (3.8-6.5 ms). This difference in latency distribution was independent of stimulus amplitude. These data are consistent with the theory that delay lines utilized by FM-FM neurons are created by neurons tuned to the "FM1 frequency," and indicate that the delay lines are mostly, if not all, created in a subthalamic nucleus or nuclei. 4. The best amplitudes of BIC fibers tuned to FM1 or CF1 were 63.2 +/- 4.5 (SE) dB SPL, and those of BIC fibers tuned to FMn or CFn were 48.2 +/- 10.7 dB SPL. The distribution of the best amplitudes of BIC fibers were very similar to those of FM-FM and CF/CF neurons in the MGB. These data indicate that the amplitude selectivity of thalamic FM-FM and CF/CF neurons is mainly a product of a subthalamic nucleus or nuclei.(ABSTRACT TRUNCATED AT 400 WORDS)

scholarly journals The resting frequency of echolocation signals changes with body temperature in the hipposiderid bat, Hipposideros armiger

2022 ◽  
Author(s):  
Diana Schoeppler ◽  
Annette Denzinger ◽  
Hans-Ulrich Schnitzler
Keyword(s):  
Body Temperature ◽  
Second Harmonic ◽  
Circadian Variation ◽  
Frequency Component ◽  
Reference Frequency ◽  
Constant Frequency ◽  
Emission Frequency ◽  
Warm Up ◽  
Vocal Control ◽  
And Function

Doppler shift (DS) compensating bats adjust in flight the second harmonic of the constant-frequency component (CF2) of their echolocation signals so that the frequency of the Doppler shifted echoes returning from ahead is kept constant with high precision (0.1-0.2%) at the so-called reference frequency (fref). This feedback adjustment is mediated by an audio-vocal control system which correlates with a maximal activation of the foveal resonance area in the cochlea. Stationary bats adjust the average CF2 with similar precision at the resting frequency (frest), which is slightly below the fref. Over a variety of time periods (from minutes up to years) variations of the coupled fref and frest have been observed, and were attributed to age, social influences and behavioural situations in rhinolophids and hipposiderids, and to body temperature effects and flight activity in Pteronotus parnellii. We assume that, for all DS compensating bats, a change in body temperature has a strong effect on the activation state of the foveal resonance area in the cochlea which leads to a concomitant change in emission frequency. We tested our hypothesis in a hipposiderid bat, Hipposideros armiger, and measured how the circadian variation of body temperature at activation phases affected frest. With a miniature temperature logger, we recorded the skin temperature on the back of the bats simultaneously with echolocation signals produced. During warm-up from torpor strong temperature increases were accompanied by an increase in frest, of up to 1.44 kHz. We discuss the implications of our results for the organization and function of the audio-vocal control systems of all DS compensating bats.

Electron-Nuclear Dynamics on Amplitude and Frequency Modulation of Molecular High-Order Harmonic Generation from H2+ and its Isotopes

2017 ◽  
Vol 72 (10) ◽  
pp. 941-953
Author(s):  
Hang Liu ◽  
Liqiang Feng
Keyword(s):  
Pulse Duration ◽  
Harmonic Generation ◽  
Vibrational State ◽  
High Order ◽  
Intensity Increase ◽  
Nuclear Mass ◽  
High Order Harmonic Generation ◽  
Nuclear Dynamics ◽  
Pulse Intensity ◽  
High Order Harmonic

AbstractElectron-nuclear dynamics of molecular high-order harmonic generation from H2+ and its isotopes has been theoretically investigated beyond the Born-Oppenheimer approximations. The results show that (i) due to the different ionisation probabilities and the harmonic emission times, the intensities of the harmonics from H2+ and its isotopes are very sensitive to the initial vibrational state, the pulse duration, and the pulse intensity. (ii) Due to the nonadiabatic effects in molecular high-order harmonic generation, the red-shifts of the harmonics can be found in the lower pulse intensity. With the increase of the pulse intensity, the harmonics are from the red-shifts to the blue-shifts. Moreover, as the pulse duration increases, the blue-shifts of the harmonics can be enhanced. As the initial vibrational state increases, the red-shifts of the harmonics can be decreased, whereas the blue-shifts of the harmonics can be enhanced. However, the shifts of the harmonics are decreased as the nuclear mass increases. (iii) Due to the coupled electron-nuclear dynamics in molecules, the spatial symmetry of the system is broken. As a result, non-odd harmonics can be generated at the larger internuclear distance. With the increase of the initial vibrational state or the nuclear mass, the generation of the non-odd harmonics can be enhanced and reduced, respectively. As the pulse duration or the pulse intensity increase, the generation of the non-odd harmonics can be enhanced. However, the intensities of the non-odd harmonics are decreased when using the longer pulse duration with the much higher laser intensity.

Combination-sensitive neurons in the ventroanterior area of the auditory cortex of the mustached bat

1988 ◽  
Vol 60 (6) ◽  
pp. 1908-1923 ◽  
Author(s):  
K. Tsuzuki ◽  
N. Suga
Keyword(s):  
Auditory Cortex ◽  
Functional Organization ◽  
Primary Auditory Cortex ◽  
Great Majority ◽  
Signal Pulse ◽  
Constant Frequency ◽  
Response Properties ◽  
Tuning Curves ◽  
Mustached Bat ◽  
Anterior Division

1. Because the ventroanterior (VA) area is one of the target areas of the FM-FM area in the auditory cortex of the mustached bat, Pteronotus parnellii parnellii, response properties of combination-sensitive neurons in this area were studied with constant-frequency (CF) tones, frequency-modulated (FM) sounds, and sounds similar to the bat's biosonar signal (pulse), which consisted of long CF components (CF1-4) and short FM components (FM1-4). CF1-4 and FM1-4 are the components in the four harmonics (H1-4) of the pulse. 2. Combination-sensitive neurons are clustered in a small area immediately anteroventral to the Doppler-shifted CF processing (DSCF) area and posteroventral to the anterior division of the primary auditory cortex. Because this cluster in the VA area is small, it was difficult to record a sufficient number of combination-sensitive neurons to explore the functional organization of the cluster, but it was found that the response properties of these VA neurons were unique. 3. Combination-sensitive neurons in the VA area are tuned to particular combinations of signal elements similar to the first and second harmonics of the pulse and/or echo. Unlike neurons in the FM-FM, dorsal fringe (DF), and CF/CF areas, no neurons in the VA area are tuned to the signal elements in the first and third or fourth harmonics. 4. The great majority of combination-sensitive neurons in the VA area can not be easily classified into either FM-FM or CF/CF neurons, because they show facilitative responses to combinations of CF1/CF2, FM1-FM2, and FM1-CF2. Therefore, they are called H1-H2 neurons. In the FM-FM and CF/CF areas, all the neurons could be easily classified as FM-FM or CF/CF. This uniqueness of H1-H2 neurons is related to the fact that their best frequencies for facilitation are predominantly between 61.0 and 62.0 kHz, i.e., within the frequency range of stabilized Doppler-shifted echo CF2. 5. In addition to 27 H1-H2 neurons, 7 FM1-FM2 neurons were also recorded in the VA area. The best delays of these H1-H2 and FM1-FM2 neurons measured with FM1-FM2 pairs are between 1 and 10 ms. Unlike neurons in the FM-FM and DF areas, their delay-tuning curves are very broad, even if their best delays are short, and extend beyond zero delay to several millisecond "negative" delays of the FM2 from the FM1, i.e., several millisecond delays of the FM1 from the FM2.(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.

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