scholarly journals Amplitude discrimination is predictably affected by echo frequency filtering in wideband echolocating bats

2021 ◽  
Author(s):  
Amaro Tuninetti ◽  
Andrea Megela Simmons ◽  
James A Simmons

Big brown bats emit wideband frequency modulated (FM) ultrasonic pulses for echolocation. They perceive target range from echo delay and target size from echo amplitude. Their sounds contain two prominent down-sweeping harmonic sweeps (FM1, ~55-22 kHz; FM2, ~100-55 kHz), which are affected differently by propagation out to the target and back to the bat. FM2 is attenuated more than FM1 during propagation. Bats anchor target ranging asymmetrically on the low frequencies in FM1, while FM2 only contributes if FM1 is present as well. These experiments tested whether the bat's ability to discriminate target size from the amplitude of echoes is affected by selectively attenuating upper or lower frequencies. Bats were trained to perform an echo amplitude discrimination task with virtual echo targets 83 cm away. While echo delay was held constant and echo amplitude was varied to estimate threshold, either lower FM1 frequencies or higher FM2 frequencies were attenuated. The results parallel effects seen in echo delay experiments; bats' performance was significantly poorer when the lower frequencies in echoes were attenuated, compared to higher frequencies. The bat's ability to distinguish between virtual targets at the same simulated range from echoes arriving at the same delay indicates a high level of focused attention for perceptual isolation of one and suppression of the other.

2020 ◽  
Vol 117 (29) ◽  
pp. 17288-17295 ◽  
Author(s):  
Chen Ming ◽  
Mary E. Bates ◽  
James A. Simmons

Big brown bats transmit wideband FM biosonar sounds that sweep from 55 to 25 kHz (first harmonic, FM1) and from 110 to 50 kHz (second harmonic, FM2). FM1 is required to perceive echo delay for target ranging; FM2 contributes only if corresponding FM1 frequencies are present. We show that echoes need only the lowest FM1 broadcast frequencies of 25 to 30 kHz for delay perception. If these frequencies are removed, no delay is perceived. Bats begin echo processing at the lowest frequencies and accumulate perceptual acuity over successively higher frequencies, but they cannot proceed without the low-frequency starting point in their broadcasts. This reveals a solution to pulse-echo ambiguity, a serious problem for radar or sonar. In dense, extended biosonar scenes, bats have to emit sounds rapidly to avoid collisions with near objects. But if a new broadcast is emitted when echoes of the previous broadcast still are arriving, echoes from both broadcasts intermingle, creating ambiguity about which echo corresponds to which broadcast. Frequency hopping by several kilohertz from one broadcast to the next can segregate overlapping narrowband echo streams, but wideband FM echoes ordinarily do not segregate because their spectra still overlap. By starting echo processing at the lowest frequencies in frequency-hopped broadcasts, echoes of the higher hopped broadcast are prevented from being accepted by lower hopped broadcasts, and ambiguity is avoided. The bat-inspired spectrogram correlation and transformation (SCAT) model also begins at the lowest frequencies; echoes that lack them are eliminated from processing of delay and no longer cause ambiguity.


1991 ◽  
Vol 66 (6) ◽  
pp. 1951-1964 ◽  
Author(s):  
D. C. Fitzpatrick ◽  
N. Suga ◽  
H. Misawa

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)


2012 ◽  
Vol 108 (10) ◽  
pp. 2641-2652 ◽  
Author(s):  
K. Heimonen ◽  
E.-V. Immonen ◽  
R. V. Frolov ◽  
I. Salmela ◽  
M. Juusola ◽  
...  

In dim light, scarcity of photons typically leads to poor vision. Nonetheless, many animals show visually guided behavior with dim environments. We investigated the signaling properties of photoreceptors of the dark active cockroach ( Periplaneta americana) using intracellular and whole-cell patch-clamp recordings to determine whether they show selective functional adaptations to dark. Expectedly, dark-adapted photoreceptors generated large and slow responses to single photons. However, when light adapted, responses of both phototransduction and the nontransductive membrane to white noise (WN)-modulated stimuli remained slow with corner frequencies ∼20 Hz. This promotes temporal integration of light inputs and maintains high sensitivity of vision. Adaptive changes in dynamics were limited to dim conditions. Characteristically, both step and frequency responses stayed effectively unchanged for intensities >1,000 photons/s/photoreceptor. A signal-to-noise ratio (SNR) of the light responses was transiently higher at frequencies <5 Hz for ∼5 s after light onset but deteriorated to a lower value upon longer stimulation. Naturalistic light stimuli, as opposed to WN, evoked markedly larger responses with higher SNRs at low frequencies. This allowed realistic estimates of information transfer rates, which saturated at ∼100 bits/s at low-light intensities. We found, therefore, selective adaptations beneficial for vision in dim environments in cockroach photoreceptors: large amplitude of single-photon responses, constant high level of temporal integration of light inputs, saturation of response properties at low intensities, and only transiently efficient encoding of light contrasts. The results also suggest that the sources of the large functional variability among different photoreceptors reside mostly in phototransduction processes and not in the properties of the nontransductive membrane.


2017 ◽  
Vol 866 ◽  
pp. 272-276
Author(s):  
Naphat Albutt ◽  
Suejit Pechprasarn ◽  
Phimonkhae Chobdee ◽  
Thanapong Sareein

The dielectric permittivity (ε) of Y2NiMnO6 ceramics prepared by sintering at 1400 °C over 6 to 24 hours was investigated. The response of the ceramics was measured from 1 kHz to 3 MHz, with the influence of a fixed dc bias from 0 to 1.5 V and temperature from 40 °C to -60 oC. Increasing dc bias was found to reduce ε' at low frequencies, while at higher frequencies the dc bias had less influence on ε'. At 40 °C a sharp transition from high to low ε' occurred starting at ~100 kHz, as the temperature of the ceramic was lowered, the transition shifted to lower frequencies. This behaviour is attributed to the charge ordering of Ni2+ and Mn4+ ions and the thermal effect on the ions energy.


Weed Science ◽  
2010 ◽  
Vol 58 (1) ◽  
pp. 30-38 ◽  
Author(s):  
Vince M. Davis ◽  
Greg R. Kruger ◽  
Steven G. Hallett ◽  
Patrick J. Tranel ◽  
William G. Johnson

Horseweed has rapidly become a major weed in soybean and cotton production fields of the United States, and Indiana farmers ranked horseweed as one of the five worst weeds in their fields during a mail survey in 2003. Glyphosate resistance in horseweed is conferred by a single, incompletely dominant gene. Horseweed populations possess a high level of variability in their response to glyphosate. Horseweed has also evolved resistance to acetolactate synthase (ALS) inhibitors, and biotypes resistant to ALS-inhibiting herbicides and glyphosate are in many of the same areas. An experiment was designed to determine whether glyphosate resistance can be transferred by pollen. We found glyphosate-resistant plants in 1.1 to 3.8% of the progeny. Segregation ratios fit the expected 3 : 1 resistant : sensitive ratios confirming that glyphosate resistance in horseweed can transfer to closely located glyphosate-susceptible biotypes under open-pollinated conditions at low frequencies. The hypothesis of a follow-up experiment was that first-generation progeny of parent plants, selected on a continuum of low to high phenotypic response to glyphosate, will inherit respective low to high phenotypic responses to glyphosate. The variability in field-collected populations (low-level to high-level glyphosate resistance) ranged from 2 to 14 times the commonly recommended field use rate of glyphosate. However, low- and high-levels of glyphosate resistance were not observed in first-generation progeny. We conclude that differential glyphosate responses observed among parental populations was due to different frequencies of the resistance allele within the populations, rather than the presence of different resistance alleles.


1973 ◽  
Vol 59 (3) ◽  
pp. 617-629
Author(s):  
C. J. BROKAW ◽  
R. JOSSLIN

1. Treatment of Ciona spermatozoa with low concentrations of Triton X-100 (less than 0·01 %) causes them to beat at lower than normal frequencies. The wavelength of the flagellar bending waves remains constant over the range from 10 to 40 Hz. There is a small increase in wavelength at lower frequencies; in the range of 1·5-6·2 Hz, the wavelength averaged 114% of the normal value for Ciona spermatozoa. The angle of bend of the bent regions of the flagellar bending waves remained constant within ± 10% over this range of frequencies. 2. Decapitated sperm flagella from Lytechinus beat at a continually declining frequency as they exhaust their content of ATP. Both wavelength and bend angle retain normal values until the frequency falls below about 8 Hz. Both parameters increase at lower frequencies, with a sharp increase below 3 Hz. 3. ATP-reactivated spermatozoa from Lytechinus show relatively small changes in wavelength and bend angle as the frequency is varied over the range from 5 to 25 Hz by varying the ATP concentration. 4. Constancy of wavelength over a wide range of frequencies is consistent with the hypothesis that wavelength is determined by the relative values of viscous bending resistance within a flagellum and external viscosity. 5. No satisfactory explanation is available at present for the constancy of bend angle over a wide range of frequencies nor for the changes in wave parameters which are observed at low frequencies.


1997 ◽  
Vol 41 (10) ◽  
pp. 2244-2250 ◽  
Author(s):  
F M Aarestrup ◽  
E M Nielsen ◽  
M Madsen ◽  
J Engberg

The MICs of 16 antimicrobial agents were determined for 202 Campylobacter jejuni isolates, 123 Campylobacter coli isolates, and 6 Campylobacter lari isolates from humans and food animals in Denmark. The C. jejuni isolates originated from humans (75), broilers (95), cattle (29), and pigs (3); the C. coli isolates originated from humans (7), broilers (17), and pigs (99); and the C. lari isolates originated from broilers (5) and cattle (1). All isolates were susceptible to apramycin, neomycin, and gentamicin. Only a few C. jejuni isolates were resistant to one or more antimicrobial agents. Resistance to tetracycline was more common among C. jejuni isolates from humans (11%) than among C. jejuni isolates from animals (0 to 2%). More resistance to streptomycin was found among C. jejuni isolates from cattle (10%) than among those from humans (4%) or broilers (1%). A greater proportion of C. coli than of C. jejuni isolates were resistant to the other antimicrobial agents tested. Isolates were in most cases either coresistant to tylosin, spiramycin, and erythromycin or susceptible to all three antibiotics. More macrolide-resistant isolates were observed among C. coli isolates from swine (79%) than among C. coli isolates from broilers (18%) and humans (14%). Twenty-four percent of C. coli isolates from pigs were resistant to enrofloxacin, whereas 29% of C. coli isolates from humans and none from broilers were resistant. More resistance to streptomycin was observed among C. coli isolates from swine (48%) than among C. coli isolates from broilers (6%) or humans (0%). The six C. lari isolates were susceptible to all antimicrobial agents except ampicillin and nalidixic acid. This study showed that antimicrobial resistance was found only at relatively low frequencies among C. jejuni and C. lari isolates. Among C. coli isolates, especially from swine, there was a high level of resistance to macrolides and streptomycin. Furthermore, this study showed differences in the resistance to antimicrobial agents among Campylobacter isolates of different origins.


1989 ◽  
Vol 61 (1) ◽  
pp. 202-207 ◽  
Author(s):  
H. Edamatsu ◽  
M. Kawasaki ◽  
N. Suga

1. The orientation sound (pulse) of the mustached bat, Pteronotus parnellii parnellii, consists of long constant-frequency components (CF1-4) and short frequency-modulated components (FM1-4). The auditory cortex of this bat contains several combination-sensitive areas: FM-FM, DF, VA, VF, and CF/CF. The FM-FM area consists of neurons tuned to a combination of the pulse FM1 and the echo FMn (n = 2, 3, or 4) and has an echo-delay (target-range) axis. Our preliminary anatomical studies with tritiated amino acids suggest that the FM-FM area projects to the dorsal fringe (DF) area, which in turn projects to the ventral fringe (VF) area. The aim of our study was to characterize the response properties of VF neurons and to explore the functional organization of the VF area. Acoustic stimuli delivered to the bats were CF tones, FM sounds, and their combinations mimicking the pulse emitted by the mustached bat and the echo. 2. Like the FM-FM and DF areas, the VF area is composed of three types of FM-FM combination-sensitive neurons: FM1-FM2, FM1-FM3, and FM1-FM4. These neurons show little or no response to a pulse alone, echo alone, single CF tone or single FM sound. They do, however, show a strong facilitative response to a pulse-echo pair with a particular echo delay. The essential components in the pulse-echo pair for facilitation are the FM1 of the pulse and the FMn of the echo.(ABSTRACT TRUNCATED AT 250 WORDS)


2018 ◽  
Vol 120 (3) ◽  
pp. 1323-1339 ◽  
Author(s):  
Silvio Macías ◽  
Jinhong Luo ◽  
Cynthia F. Moss

Echolocating bats must process temporal streams of sonar sounds to represent objects along the range axis. Neuronal echo-delay tuning, the putative mechanism of sonar ranging, has been characterized in the inferior colliculus (IC) of the mustached bat, an insectivorous species that produces echolocation calls consisting of constant frequency and frequency modulated (FM) components, but not in species that use FM signals alone. This raises questions about the mechanisms that give rise to echo-delay tuning in insectivorous bats that use different signal designs. To investigate whether stimulus context may account for species differences in echo-delay selectivity, we characterized single-unit responses in the IC of awake passively listening FM bats, Eptesicus fuscus, to broadcasts of natural sonar call-echo sequences, which contained dynamic changes in signal duration, interval, spectrotemporal structure, and echo-delay. In E. fuscus, neural selectivity to call-echo delay emerges in a population of IC neurons when stimulated with call-echo pairs presented at intervals mimicking those in a natural sonar sequence. To determine whether echo-delay selectivity also depends on the spectrotemporal features of individual sounds within natural sonar sequences, we studied responses to computer-generated echolocation signals that controlled for call interval, duration, bandwidth, sweep rate, and echo-delay. A subpopulation of IC neurons responded selectively to the combination of the spectrotemporal structure of natural call-echo pairs and their temporal patterning within a dynamic sonar sequence. These new findings suggest that the FM bat’s fine control over biosonar signal parameters may modulate IC neuronal selectivity to the dimension of echo-delay. NEW & NOTEWORTHY Echolocating bats perform precise auditory temporal computations to estimate their distance to objects. Here, we report that response selectivity of neurons in the inferior colliculus of a frequency modulated bat to call-echo delay, or target range tuning, depends on the temporal patterning and spectrotemporal features of sound elements in a natural echolocation sequence. We suggest that echo responses to objects at different distances are gated by the bat’s active control over the spectrotemporal patterning of its sonar emissions.


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