Discrimination of surface-structured targets by the echolocating batMyotis myotis during flight (original) (raw)
Related papers
Representation of Perceptual Dimensions of Insect Prey During Terminal Pursuit by Echolocating Bats
The Biological Bulletin, 1996
The echolocating big brown bat, Eptesicus fusciis, broadcasts brief frequency-modulated (FM) ultrasonic sounds and perceives objects from echoes of these sounds returning to its ears. Eptesicus is an insectivorous species that uses sonar to locate and track flying prey. Although the bat normally hunts in open areas, it nevertheless is capable of chasing insects into cluttered environments such as vegetation, where it completes interceptions in much the same manner as in the open except that it has to avoid the obstacles as well as catch the insect. During pursuit, the bat shortens its sonar signals and increases their rate of emission as it closes in to seize the target, and it keeps its head pointed at the insect throughout the maneuver. In the terminal stage of interception, the bat makes rapid adjustments in its flightpath and body posture to capture the insect, and these reactions occur whether the bat is pursuing its prey in the open or close to obstacles such as vegetation. Insects can be distinguished from other objects by the spectrum and phase of their echoes, and Eptesicus is very good at discriminating these acoustic features. To identify the insect in the open, but especially to distinguish which object is the insect in clutter, the bat must have some means for representing these features throughout the interception maneuver. Moreover, continuity for perception of these features is necessary to keep track of the prey in complex surroundings, so the nature of the auditory representations for the spectrum and phase of echoes has to be conserved across the approach, tracking, and terminal stages. The first problem is that representation of changes
Biosonar discrimination of fine surface textures by echolocating free-tailed bats
Frontiers in Ecology and Evolution
Echolocating bats are able to discriminate between different surface textures based on the spectral properties of returning echoes. This capability is likely to be important for recognizing prey and for finding suitably perching sites along smooth cave walls. Previous studies showed that bats may exploit echo spectral interference patterns in returning echoes to classify surface textures, but a systematic assessment of the limits of their discrimination performance is lacking and may provide important clues about the neural mechanisms by which bats reconstruct target features based on echo acoustic cues. We trained three Mexican free-tailed bats (Tadarida brasiliensis) on a Y-maze to discriminate between the surfaces of 10 different sheets of aluminum-oxide abrasive sandpapers differing in standardized grit sizes ranging from 40 grit (coarse, 425 μm mean particle diameter) to 240 grit (fine, 54 μm mean particle diameter). Bats were rewarded for choosing the coarsest of two choices. ...
Evidence for spatial representation of object shape by echolocating bats (Eptesicus fuscus)
The Journal of the Acoustical Society of America, 2008
Big brown bats were trained in a two-choice task to locate a two-cylinder dipole object with a constant 5 cm spacing in the presence of either a one-cylinder monopole or another two-cylinder dipole with a shorter spacing. For the dipole versus monopole task, the objects were either stationary or in motion during each trial. The dipole and monopole objects varied from trial to trial in the left-right position while also roving in range ͑10-40 cm͒, cross range separation ͑15-40 cm͒, and dipole aspect angle ͑0°-90°͒. These manipulations prevented any single feature of the acoustic stimuli from being a stable indicator of which object was the correct choice. After accounting for effects of masking between echoes from pairs of cylinders at similar distances, the bats discriminated the 5 cm dipole from both the monopole and dipole alternatives with performance independent of aspect angle, implying a distal, spatial object representation rather than a proximal, acoustic object representation.
Acoustic scanning of natural scenes by echolocation in the big brown bat, Eptesicus fuscus
Journal of Experimental Biology, 2009
Echolocation allows bats to orient and localize prey in complete darkness. The sonar beam of the big brown bat, Eptesicus fuscus, is directional but broad enough to provide audible echo information from within a 60-90 deg. cone. This suggests that the big brown bat could interrogate a natural scene without fixating each important object separately. We tested this idea by measuring the directional aim and duration of the bat's sonar beam as it performed in a dual task, obstacle avoidance and insect capture. Bats were trained to fly through one of two openings in a fine net to take a tethered insect at variable distances behind the net. The bats sequentially scanned the edges of the net opening and the prey by centering the axis of their sonar beam with an accuracy of ~5 deg. The bats also shifted the duration of their sonar calls, revealing sequential sampling along the range axis. Changes in duration and directional aim were correlated, showing that the bat first inspected the hole, and then shifted its gaze to the more distant insect, before flying through the net opening. Contrary to expectation based on the sonar beam width, big brown bats encountering a complex environment accurately pointed and shifted their sonar gaze to sequentially inspect closely spaced objects in a manner similar to visual animals using saccades and fixations to scan a scene. The findings presented here from a specialized orientation system, echolocation, offer insights into general principles of active sensing across sensory modalities for the perception of natural scenes.
Adaptive behavior for texture discrimination by the free-flying big brown bat, Eptesicus fuscus
Journal of Comparative Physiology A, 2011
This study examined behavioral strategies for texture discrimination by echolocation in free-flying bats. Big brown bats, Eptesicus fuscus, were trained to discriminate a smooth 16 mm diameter object (S?) from a size-matched textured object (S-), both of which were tethered in random locations in a flight room. The bat's three-dimensional flight path was reconstructed using stereo images from high-speed video recordings, and the bat's sonar vocalizations were recorded for each trial and analyzed off-line. A microphone array permitted reconstruction of the sonar beam pattern, allowing us to study the bat's directional gaze and inspection of the objects. Bats learned the discrimination, but performance varied with S-. In acoustic studies of the objects, the S? and S-stimuli were ensonified with frequency-modulated sonar pulses. Mean intensity differences between S? and S-were within 4 dB. Performance data, combined with analyses of echo recordings, suggest that the big brown bat listens to changes in sound spectra from echo to echo to discriminate between objects. Bats adapted their sonar calls as they inspected the stimuli, and their sonar behavior resembled that of animals foraging for insects. Analysis of sonar beam-directing behavior in certain trials clearly showed that the bat sequentially inspected S? and S-.
Trawling bats exploit an echo-acoustic ground effect
Frontiers in physiology, 2013
A water surface acts not only as an optic mirror but also as an acoustic mirror. Echolocation calls emitted by bats at low heights above water are reflected away from the bat, and hence the background clutter is reduced. Moreover, targets on the surface create an enhanced echo. Here, we formally quantified the effect of the surface and target height on both target detection and -discrimination in a combined laboratory and field approach with Myotis daubentonii. In a two-alternative, forced-choice paradigm, the bats had to detect a mealworm and discriminate it from an inedible dummy (20 mm PVC disc). Psychophysical performance was measured as a function of height above either smooth surfaces (water or PVC) or above a clutter surface (artificial grass). At low heights above the clutter surface (10, 20, or 35 cm), the bats' detection performance was worse than above a smooth surface. At a height of 50 cm, the surface structure had no influence on target detection. Above the clutter...
Complex echo classification by echo-locating bats: a review
Echo-locating bats constantly emit ultrasonic pulses and analyze the returning echoes to detect, localize, and classify objects in their surroundings. Echo classification is essential for bats' everyday life; for instance, it enables bats to use acoustical landmarks for navigation and to recognize food sources from other objects. Most of the research of echo based object classification in echo-locating bats was done in the context of simple artificial objects. These objects might represent prey, flower, or fruit and are characterized by simple echoes with a single up to several reflectors. Bats, however, must also be able to use echoes that return from complex structures such as plants or other types of background. Such echoes are characterized by superpositions of many reflections that can only be described using a stochastic statistical approach. Scientists have only lately started to address the issue of complex echo classification by echo-locating bats. Some behavioral evidence showing that bats can classify complex echoes has been accumulated and several hypotheses have been suggested as to how they do so. Here, we present a first review of this data. We raise some hypotheses regarding possible interpretations of the data and point out necessary future directions that should be pursued.
Object-Oriented Echo Perception and Cortical Representation in Echolocating Bats
PLoS Biology, 2007
Echolocating bats can identify three-dimensional objects exclusively through the analysis of acoustic echoes of their ultrasonic emissions. However, objects of the same structure can differ in size, and the auditory system must achieve a size-invariant, normalized object representation for reliable object recognition. This study describes both the behavioral classification and the cortical neural representation of echoes of complex virtual objects that vary in object size. In a phantom-target playback experiment, it is shown that the bat Phyllostomus discolor spontaneously classifies most scaled versions of objects according to trained standards. This psychophysical performance is reflected in the electrophysiological responses of a population of cortical units that showed an object-size invariant response (14/109 units, 13%). These units respond preferentially to echoes from objects in which echo duration (encoding object depth) and echo amplitude (encoding object surface area) co-varies in a meaningful manner. These results indicate that at the level of the bat's auditory cortex, an object-oriented rather than a stimulus-parameter-oriented representation of echoes is achieved.
Gleaning insectivorous bats that forage by using echolocation within dense forest vegetation face the sensorial challenge of acoustic masking effects. Active perception of silent and motionless prey in acoustically cluttered environments by echolocation alone has thus been regarded impossible. The gleaning insectivorous bat Micronycteris microtis however, forages in dense understory vegetation and preys on insects, including dragonflies, which rest silent and motionless on vegetation. From behavioural experiments, we show that M. microtis uses echolocation as the sole sensorial modality for successful prey perception within a complex acoustic environment. All individuals performed a stereotypical three-dimensional hovering flight in front of prey items, while continuously emitting short, multi-harmonic, broadband echolocation calls. We observed a high precision in target localization which suggests that M. microtis perceives a detailed acoustic image of the prey based on shape, surface structure and material. Our experiments provide, to our knowledge, the first evidence that a gleaning bat uses echolocation alone for successful detection, classification and precise localization of silent and motionless prey in acoustic clutter. Overall, we conclude that the three-dimensional hovering flight of M. microtis in combination with a frequent emission of short, high-frequency echolocation calls is the key for active prey perception in acoustically highly cluttered environments.
Scientific Reports
Using echolocation, bats receive acoustic information on their surroundings, which is assumed to help them sophisticatedly navigate complex environments. In this study, to understand spatial learning and acoustic sensing in bats, we investigated how flight and echolocation control changed in Rhinolophus ferrumequinum nippon as they learnt about their surroundings in an obstacle course that they flew through repeatedly. In these experiments, two testing environments (acoustically permeable and acoustically reflective) were prepared using chains and acrylic boards as obstacles to evaluate the interactive effects of spatial learning and flight environments. We found that bats reduced the meandering width of their flights and pulse emissions, and also seemed to reduce their shifts in pulse direction as they learnt more about their environments in both conditions. Throughout all our experiments, the bats with slower flight speeds tended to emit more pulses, which suggests that the number...
Echolocating bats inspect and discriminate landmark features to guide navigation
The Journal of Experimental Biology, 2019
Landmark-guided navigation is a common behavioral strategy for way-finding, yet prior studies have not examined how animals collect sensory information to discriminate landmark features. We investigated this question in animals that rely on active sensing to guide navigation. Four echolocating bats (Eptesicus fuscus) were trained to use an acoustic landmark to find and navigate through a net opening for a food reward. In experimental trials, an object serving as a landmark was placed adjacent to a net opening and a distractor object next to a barrier (covered opening). The location of the opening, barrier, and objects were moved between trials, but the spatial relationships between the landmark and opening, and between the distractor and barrier were maintained. In probe trials, the landmark was placed next to a barrier, while the distractor was placed next to the opening, to test whether the bats relied on the landmark to guide navigation. Vocal and flight behaviors were recorded w...
Probing the natural scene by echolocation in bats
2010
a direct impact on the information available to its acoustic imaging system. In turn, the bat's perception of the echo scene guides its adjustments of the features of subsequent sonar vocalizations. Therefore, the bat's adaptive sonar behavior can shed light on the fundamental processes that underlie auditory scene analysis by echolocation. Indeed, the bat's active sonar behavior allows us to listen in on the signals that are used to perform a variety of auditory tasks. Here we review laboratory and field studies that point to the features echolocating bats use to represent information about a complex acoustic environment and the general importance of an animal's actions to its perception of natural scenes.
The Journal of experimental biology, 2014
Formerly thought to be a strictly insectivorous trawling bat, recent studies have shown that Myotis capaccinii also preys on fish. To determine whether differences exist in bat flight behaviour, prey handling and echolocation characteristics when catching fish and insects of different size, we conducted a field experiment focused on the last stage of prey capture. We used synchronized video and ultrasound recordings to measure several flight and dip features as well as echolocation characteristics, focusing on terminal buzz phase I, characterized by a call rate exceeding 100 Hz, and buzz phase II, characterized by a drop in the fundamental well below 20 kHz and a repetition rate exceeding 150 Hz. When capturing insects, bats used both parts of the terminal phase to the same extent, and performed short and superficial drags on the water surface. In contrast, when preying on fish, buzz I was longer and buzz II shorter, and the bats made longer and deeper dips. These variations suggest...
2003
The use of ultrasonic echolocation (sonar) in air is seriously constrained by the attenuation of high frequency sounds and unwanted echoes from the background (called clutter). Therefore, in many situations, echolocating bats have to rely on other sensory cues. The aim of this thesis is to investigate the use of vision by echolocating bats. Bat eyes are generally small, especially among aerial hawking insectivores, with the exception of members of the family Emballonuridae. In gleaning, and in frugivorous species, however, the eyes tend to be larger and more prominent. The eyes of all bats are well adapted to low illumination, having mainly rod-based retinas, large corneal surfaces and lenses, and generally large receptor fields. Bats can easily detect small differences in brightness on clear nights, and the visual acuity remains relatively good in dim illuminations. The visual resolving power (as obtained from counts of retinal ganglion cells or by optomotor response tests) varies ...
Hunting bats adjust their echolocation to receive weak prey echoes for clutter reduction
Science Advances, 2021
How animals extract information from their surroundings to guide motor patterns is central to their survival. Here, we use echo-recording tags to show how wild hunting bats adjust their sensory strategies to their prey and natural environment. When searching, bats maximize the chances of detecting small prey by using large sensory volumes. During prey pursuit, they trade spatial for temporal information by reducing sensory volumes while increasing update rate and redundancy of their sensory scenes. These adjustments lead to very weak prey echoes that bats protect from interference by segregating prey sensory streams from the background using a combination of fast-acting sensory and motor strategies. Counterintuitively, these weak sensory scenes allow bats to be efficient hunters close to background clutter broadening the niches available to hunt for insects.
Related topics
Cited by
Adaptive behavior for texture discrimination by the free-flying big brown bat, Eptesicus fuscus
Journal of Comparative Physiology A, 2011
This study examined behavioral strategies for texture discrimination by echolocation in free-flying bats. Big brown bats, Eptesicus fuscus, were trained to discriminate a smooth 16 mm diameter object (S?) from a size-matched textured object (S-), both of which were tethered in random locations in a flight room. The bat's three-dimensional flight path was reconstructed using stereo images from high-speed video recordings, and the bat's sonar vocalizations were recorded for each trial and analyzed off-line. A microphone array permitted reconstruction of the sonar beam pattern, allowing us to study the bat's directional gaze and inspection of the objects. Bats learned the discrimination, but performance varied with S-. In acoustic studies of the objects, the S? and S-stimuli were ensonified with frequency-modulated sonar pulses. Mean intensity differences between S? and S-were within 4 dB. Performance data, combined with analyses of echo recordings, suggest that the big brown bat listens to changes in sound spectra from echo to echo to discriminate between objects. Bats adapted their sonar calls as they inspected the stimuli, and their sonar behavior resembled that of animals foraging for insects. Analysis of sonar beam-directing behavior in certain trials clearly showed that the bat sequentially inspected S? and S-.
Echo-delay resolution in sonar images of the big brown bat, Eptesicus fuscus
Proceedings of the National Academy of Sciences, 1998
broadcast ultrasonic frequency-modulated (FM) biosonar sounds (20-100 kHz frequencies; 10-50 s periods) and perceive target range from echo delay. Knowing the acuity for delay resolution is essential to understand how bats process echoes because they perceive target shape and texture from the delay separation of multiple reflections. Bats can separately perceive the delays of two concurrent electronically generated echoes arriving as little as 2 s apart, thus resolving reflecting points as close together as 0.3 mm in range (two-point threshold). This two-point resolution is roughly five times smaller than the shortest periods in the bat's sounds. Because the bat's broadcasts are 2,000-4,500 s long, the echoes themselves overlap and interfere with each other, to merge together into a single sound whose spectrum is shaped by their mutual interference depending on the size of the time separation. To separately perceive the delays of overlapping echoes, the bat has to recover information about their very small delay separation that was transferred into the spectrum when the two echoes interfered with each other, thus explicitly reconstructing the range profile of targets from the echo spectrum. However, the bat's 2-s resolution limit is so short that the available spectral cues are extremely limited. Resolution of delay seems overly sharp just for interception of flying insects, which suggests that the bat's biosonar images are of higher quality to suit a wider variety of orientation tasks, and that biosonar echo processing is correspondingly more sophisticated than has been suspected.
Age-dependent gene expression in the inner ear of big brown bats (Eptesicus fuscus)
PloS one, 2017
For echolocating bats, hearing is essential for survival. Specializations for detecting and processing high frequency sounds are apparent throughout their auditory systems. Recent studies on echolocating mammals have reported evidence of parallel evolution in some hearing-related genes in which distantly related groups of echolocating animals (bats and toothed whales), cluster together in gene trees due to apparent amino acid convergence. However, molecular adaptations can occur not only in coding sequences, but also in the regulation of gene expression. The aim of this study was to examine the expression of hearing-related genes in the inner ear of developing big brown bats, Eptesicus fuscus, during the period in which echolocation vocalizations increase dramatically in frequency. We found that seven genes were significantly upregulated in juveniles relative to adults, and that the expression of four genes through development correlated with estimated age. Compared to available dat...
Echolocation: a personal historical perspective
Canadian Journal of Zoology, 2018
The remarkable ability of echolocating bats to use echoes of emitted sounds as a substitute for vision is one of the most fascinating stories in neuroethology. Based in part on personal experience, the author discusses key discoveries in the early decades after the discovery of the phenomenon by Griffin and Galambos in 1941. Advances in our understanding of this remarkable phenomenon illustrate well the importance of comparative approaches, technological advances, and the synergistic interaction between behavior and electrophysiology.
Proceedings of the Royal Society B: Biological Sciences, 2013
Gleaning insectivorous bats that forage by using echolocation within dense forest vegetation face the sensorial challenge of acoustic masking effects. Active perception of silent and motionless prey in acoustically cluttered environments by echolocation alone has thus been regarded impossible. The gleaning insectivorous bat Micronycteris microtis however, forages in dense understory vegetation and preys on insects, including dragonflies, which rest silent and motionless on vegetation. From behavioural experiments, we show that M. microtis uses echolocation as the sole sensorial modality for successful prey perception within a complex acoustic environment. All individuals performed a stereotypical three-dimensional hovering flight in front of prey items, while continuously emitting short, multi-harmonic, broadband echolocation calls. We observed a high precision in target localization which suggests that M. microtis perceives a detailed acoustic image of the prey based on shape, surf...
Journal of neurophysiology, 2002
The acoustic environment for an echolocating bat can contain multiple objects that reflect echoes so closely separated in time that they are almost completely overlapping. This results in a single echo with a spectrum characterized by deep notches due to interference. The object of this study was to document the possible selectivity, or lack thereof, of auditory neurons to the temporal separation of biosonar signals on a coarse (ms) and fine (micros) temporal scale. We recorded single-unit activity from the auditory cortex of big brown bats while presenting four protocol designs using wideband FM signals. The protocols simulated a pair of partially overlapping echoes where the separation between the first and second echo varied between 0 and 72 micros, a pulse followed by a single echo at varying delay from 0 to 30 ms, a pulse followed at a fixed delay by a pair of partially overlapping echoes that had a varying temporal separation of 0-72 micros, and a pulse followed, with a varyin...
Temporal coding of echo spectral shape in the bat auditory cortex
PLOS Biology, 2020
Echolocating bats rely upon spectral interference patterns in echoes to reconstruct fine details of a reflecting object’s shape. However, the acoustic modulations required to do this are extremely brief, raising questions about how their auditory cortex encodes and processes such rapid and fine spectrotemporal details. Here, we tested the hypothesis that biosonar target shape representation in the primary auditory cortex (A1) is more reliably encoded by changes in spike timing (latency) than spike rates and that latency is sufficiently precise to support a synchronization-based ensemble representation of this critical auditory object feature space. To test this, we measured how the spatiotemporal activation patterns of A1 changed when naturalistic spectral notches were inserted into echo mimic stimuli. Neurons tuned to notch frequencies were predicted to exhibit longer latencies and lower mean firing rates due to lower signal amplitudes at their preferred frequencies, and both were ...
Biosonar discrimination of fine surface textures by echolocating free-tailed bats
Frontiers in Ecology and Evolution
Echolocating bats are able to discriminate between different surface textures based on the spectral properties of returning echoes. This capability is likely to be important for recognizing prey and for finding suitably perching sites along smooth cave walls. Previous studies showed that bats may exploit echo spectral interference patterns in returning echoes to classify surface textures, but a systematic assessment of the limits of their discrimination performance is lacking and may provide important clues about the neural mechanisms by which bats reconstruct target features based on echo acoustic cues. We trained three Mexican free-tailed bats (Tadarida brasiliensis) on a Y-maze to discriminate between the surfaces of 10 different sheets of aluminum-oxide abrasive sandpapers differing in standardized grit sizes ranging from 40 grit (coarse, 425 μm mean particle diameter) to 240 grit (fine, 54 μm mean particle diameter). Bats were rewarded for choosing the coarsest of two choices. ...
Ecosystem services provided by bats
Annals of the New York Academy of Sciences, 2011
Ecosystem services are the benefits obtained from the environment that increase human well-being. Economic valuation is conducted by measuring the human welfare gains or losses that result from changes in the provision of ecosystem services. Bats have long been postulated to play important roles in arthropod suppression, seed dispersal, and pollination; however, only recently have these ecosystem services begun to be thoroughly evaluated. Here, we review the available literature on the ecological and economic impact of ecosystem services provided by bats. We describe dietary preferences, foraging behaviors, adaptations, and phylogenetic histories of insectivorous, frugivorous, and nectarivorous bats worldwide in the context of their respective ecosystem services. For each trophic ensemble, we discuss the consequences of these ecological interactions on both natural and agricultural systems. Throughout this review, we highlight the research needed to fully determine the ecosystem services in question. Finally, we provide a comprehensive overview of economic valuation of ecosystem services. Unfortunately, few studies estimating the economic value of ecosystem services provided by bats have been conducted to date; however, we outline a framework that could be used in future studies to more fully address this question. Consumptive goods provided by bats, such as food and guano, are often exchanged in markets where the market price indicates an economic value. Nonmarket valuation methods can be used to estimate the economic value of nonconsumptive services, including inputs to agricultural production and recreational activities. Information on the ecological and economic value of ecosystem services provided by bats can be used to inform decisions regarding where and when to protect or restore bat populations and associated habitats, as well as to improve public perception of bats.
Biotropica, 2022
Predation can effectively limit insect herbivores with cascading effects on plant community composition and diversity of tropical rainforests. Assessing variation in predation is therefore important to understand the mechanisms structuring complex rainforest ecosystems. Variation in predation with time of day may provide herbivores with temporal enemy-free space. Trichomes (plant hairs) may provide spatial enemy-free space by increasing climbing resistance for walking arthropod predators and by scattering bat echolocation calls. Artificial model prey is commonly used to measure predation pressure on insect herbivores. Whether model prey shape is sufficient to deceive predators and whether attacks represent actual predation however remain unresolved. We used artificial, plasticine prey to assess temporal and spatial variation in predation in two Panamanian rainforests and tested whether model prey shape is as important for prey recognition by predators as often assumed. We assessed the effect of prey shape and size, time of day, and trichomes on predation by comparing attacks on caterpillar-and humanoid-shaped figurines. We find higher nocturnal than diurnal predation in one but not the other forest, suggesting that herbivores may benefit from enemy-free space during the day in some forests. We find no evidence for an effect of trichomes on predation in the two plant species tested. Equal attack numbers on caterpillar-and humanoid-shaped objects challenge the idea that the visual resemblance of model prey alone is sufficient to deceive predators. We conclude that attacks on model prey represent a variety of responses to novel objects (e.g. exploration, aggression, possibly predation) and urge caution when interpreting their results.
A comprehensive computational model of animal biosonar signal processing
PLOS Computational Biology, 2021
Computational models of animal biosonar seek to identify critical aspects of echo processing responsible for the superior, real-time performance of echolocating bats and dolphins in target tracking and clutter rejection. The Spectrogram Correlation and Transformation (SCAT) model replicates aspects of biosonar imaging in both species by processing wideband biosonar sounds and echoes with auditory mechanisms identified from experiments with bats. The model acquires broadband biosonar broadcasts and echoes, represents them as time-frequency spectrograms using parallel bandpass filters, translates the filtered signals into ten parallel amplitude threshold levels, and then operates on the resulting timeof-occurrence values at each frequency to estimate overall echo range delay. It uses the structure of the echo spectrum by depicting it as a series of local frequency nulls arranged regularly along the frequency axis of the spectrograms after dechirping them relative to the broadcast. Computations take place entirely on the timing of threshold-crossing events for each echo relative to threshold-events for the broadcast. Threshold-crossing times take into account amplitude-latency trading, a physiological feature absent from conventional digital signal processing. Amplitude-latency trading transposes the profile of amplitudes across frequencies into a profile of time-registrations across frequencies. Target shape is extracted from the spacing of the object's individual acoustic reflecting points, or glints, using the mutual interference pattern of peaks and nulls in the echo spectrum. These are merged with the overall range-delay estimate to produce a delay-based reconstruction of the object's distance as well as its glints. Clutter echoes indiscriminately activate multiple parts in the nulldetecting system, which then produces the equivalent glint-delay spacings in images, thus blurring the overall echo-delay estimates by adding spurious glint delays to the image. Blurring acts as an anticorrelation process that rejects clutter intrusion into perceptions.