Naturalistic Stimulus Structure Determines the Integration of Audiovisual Looming Signals in Binocular Rivalry

Rapid integration of biologically relevant information is crucial for the survival of an organism. Most prominently, humans should be biased to attend and respond to looming stimuli that signal approaching danger (e.g. predator) and hence require rapid action. This psychophysics study used binocular rivalry to investigate the perceptual advantage of looming (relative to receding) visual signals (i.e. looming bias) and how this bias can be influenced by concurrent auditory looming/receding stimuli and the statistical structure of the auditory and visual signals.Subjects were dichoptically presented with looming/receding visual stimuli that were paired with looming or receding sounds. The visual signals conformed to two different statistical structures: (1) a ‘simple’ random-dot kinematogram showing a starfield and (2) a “naturalistic” visual Shepard stimulus. Likewise, the looming/receding sound was (1) a simple amplitude- and frequency-modulated (AM-FM) tone or (2) a complex Shepard tone. Our results show that the perceptual looming bias (i.e. the increase in dominance times for looming versus receding percepts) is amplified by looming sounds, yet reduced and even converted into a receding bias by receding sounds. Moreover, the influence of looming/receding sounds on the visual looming bias depends on the statistical structure of both the visual and auditory signals. It is enhanced when audiovisual signals are Shepard stimuli.In conclusion, visual perception prioritizes processing of biologically significant looming stimuli especially when paired with looming auditory signals. Critically, these audiovisual interactions are amplified for statistically complex signals that are more naturalistic and known to engage neural processing at multiple levels of the cortical hierarchy.     

The effect of looming and receding sounds on the perceived in-depth orientation of depth-ambiguous biological motion figures

Background

The focus in the research on biological motion perception traditionally has been restricted to the visual modality. Recent neurophysiological and behavioural evidence, however, supports the idea that actions are not represented merely visually but rather audiovisually. The goal of the present study was to test whether the perceived in-depth orientation of depth-ambiguous point-light walkers (plws) is affected by the presentation of looming or receding sounds synchronized with the footsteps.

Methodology/Principal Findings

In Experiment 1 orthographic frontal/back projections of plws were presented either without sound or with sounds of which the intensity level was rising (looming), falling (receding) or stationary. Despite instructions to ignore the sounds and to only report the visually perceived in-depth orientation, plws accompanied with looming sounds were more often judged to be facing the viewer whereas plws paired with receding sounds were more often judged to be facing away from the viewer. To test whether the effects observed in Experiment 1 act at a perceptual level rather than at the decisional level, in Experiment 2 observers perceptually compared orthographic plws without sound or paired with either looming or receding sounds to plws without sound but with perspective cues making them objectively either facing towards or facing away from the viewer. Judging whether either an orthographic plw or a plw with looming (receding) perspective cues is visually most looming becomes harder (easier) when the orthographic plw is paired with looming sounds.

Conclusions/Significance

The present results suggest that looming and receding sounds alter the judgements of the in-depth orientation of depth-ambiguous point-light walkers. While looming sounds are demonstrated to act at a perceptual level and make plws look more looming, it remains a challenge for future research to clarify at what level in the processing hierarchy receding sounds affect how observers judge the in-depth perception of plws.     

Neural processing of auditory looming in the human brain

Acoustic intensity change, along with interaural, spectral, and reverberation information, is an important cue for the perception of auditory motion. Approaching sound sources produce increases in intensity, and receding sound sources produce corresponding decreases. Human listeners typically overestimate increasing compared to equivalent decreasing sound intensity and underestimate the time to contact of approaching sound sources. These characteristics could provide a selective advantage by increasing the margin of safety for response to looming objects. Here, we used dynamic intensity and functional magnetic resonance imaging to examine the neural underpinnings of the perceptual priority for rising intensity. We found that, consistent with activation by horizontal and vertical auditory apparent motion paradigms, rising and falling intensity activated the right temporal plane more than constant intensity. Rising compared to falling intensity activated a distributed neural network subserving space recognition, auditory motion perception, and attention and comprising the superior temporal sulci and the middle temporal gyri, the right temporoparietal junction, the right motor and premotor cortices, the left cerebellar cortex, and a circumscribed region in the midbrain. This anisotropic processing of acoustic intensity change may reflect the salience of rising intensity produced by looming sources in natural environments.     

Auditory neuroscience: the salience of looming sounds

Sounds that move towards us have a greater biological salience than those that move away. Recent studies in human and non-human primates have demonstrated a perceptual and behavioural priority for such looming sounds that is also reflected in an asymmetric pattern of cortical activation.     

Spatial coding of the predicted impact location of a looming object

Avoiding or intercepting looming objects implies a precise estimate of both time until contact and impact location. In natural situations, extrapolating a movement trajectory relative to some egocentric landmark requires taking into account variations in retinal input associated with moment-to-moment changes in body posture. Here, human observers predicted the impact location on their face of an approaching stimulus mounted on a robotic arm, while we systematically manipulated the relation between eye, head, and trunk orientation. The projected impact point on the observer's face was estimated most accurately when the target originated from a location aligned with both the head and eye axes. Eccentric targets with respect to either axis resulted in a systematic perceptual bias ipsilateral to the trajectory's origin. We conclude that (1) predicting the impact point of a looming target requires combining retinal information with eye position information, (2) that this computation is accomplished accurately for some, but not all, possible combinations of these cues, (3) that the representation of looming trajectories is not formed in a single, canonical reference frame, and (4) that the observed perceptual biases could reflect an automatic adaptation for interceptive/defensive actions within near peripersonal space.     

Fear influences perceived reaching to targets in audition,but not vision

The superordinate mechanism view of emotions predicts that fear should influence perception to carry out the evolved function of overcoming immediate threats. Previous work demonstrates that fear does adaptively influence visual perception. However, there are recurring situations in which auditory perception is used for overcoming immediate threats (e.g., avoiding predators after dark). Some research suggests that the auditory system, independent of fear, is adaptively biased to hear approaching sounds as closer than equidistant receding sounds (a.k.a. the looming bias). The present study investigated whether fear, as a superordinate mechanism, influences auditory perception such that sounds are perceived to be closer, ultimately providing an advantage when avoiding immediate threats. Participants judged whether or not they could reach to an aurally or visually perceived target while either in a fearful or neutral state. The results demonstrated that while in a fearful state, participants judged targets to be closer to them, but only when the target was perceived aurally. This finding extends previous work on adaptive biases in auditory perception to include the influence of fear.     

Distortions of subjective time perception within and across senses

Background

The ability to estimate the passage of time is of fundamental importance for perceptual and cognitive processes. One experience of time is the perception of duration, which is not isomorphic to physical duration and can be distorted by a number of factors. Yet, the critical features generating these perceptual shifts in subjective duration are not understood.

Methodology/Findings

We used prospective duration judgments within and across sensory modalities to examine the effect of stimulus predictability and feature change on the perception of duration. First, we found robust distortions of perceived duration in auditory, visual and auditory-visual presentations despite the predictability of the feature changes in the stimuli. For example, a looming disc embedded in a series of steady discs led to time dilation, whereas a steady disc embedded in a series of looming discs led to time compression. Second, we addressed whether visual (auditory) inputs could alter the perception of duration of auditory (visual) inputs. When participants were presented with incongruent audio-visual stimuli, the perceived duration of auditory events could be shortened or lengthened by the presence of conflicting visual information; however, the perceived duration of visual events was seldom distorted by the presence of auditory information and was never perceived shorter than their actual durations.

Conclusions/Significance

These results support the existence of multisensory interactions in the perception of duration and, importantly, suggest that vision can modify auditory temporal perception in a pure timing task. Insofar as distortions in subjective duration can neither be accounted for by the unpredictability of an auditory, visual or auditory-visual event, we propose that it is the intrinsic features of the stimulus that critically affect subjective time distortions.     

Rules from Words: A Dynamic Neural Basis for a Lawful Linguistic Process

Listeners show a reliable bias towards interpreting speech sounds in a way that conforms to linguistic restrictions (phonotactic constraints) on the permissible patterning of speech sounds in a language. This perceptual bias may enforce and strengthen the systematicity that is the hallmark of phonological representation. Using Granger causality analysis of magnetic resonance imaging (MRI)- constrained magnetoencephalography (MEG) and electroencephalography (EEG) data, we tested the differential predictions of rule-based, frequency–based, and top-down lexical influence-driven explanations of processes that produce phonotactic biases in phoneme categorization. Consistent with the top-down lexical influence account, brain regions associated with the representation of words had a stronger influence on acoustic-phonetic regions in trials that led to the identification of phonotactically legal (versus illegal) word-initial consonant clusters. Regions associated with the application of linguistic rules had no such effect. Similarly, high frequency phoneme clusters failed to produce stronger feedforward influences by acoustic-phonetic regions on areas associated with higher linguistic representation. These results suggest that top-down lexical influences contribute to the systematicity of phonological representation.     

Perceptual distance and the constancy of size and stereoscopic depth

The relationship between distance and size perception is unclear because of conflicting results of tests investigating the size-distance invariance hypothesis (SDIH), according to which perceived size is proportional to perceived distance. We propose that response bias with regard to measures of perceived distance is at the root of the conflict. Rather than employ the usual method of magnitude estimation, the bias-free two-alternative forced choice (2AFC) method was used to determine the precision (1/sigma) of discriminating depth at different distances. The results led us to define perceptual distance as a bias free power function of physical distance, with an exponent of approximately 0.5. Similar measures involving size differences among stimuli of equal angular size yield the same power function of distance. In addition, size discrimination is noisier than depth discrimination, suggesting that distance information is processed prior to angular size. Size constancy implies that the perceived size is proportional to perceptual distance. Moreover, given a constant relative disparity, depth constancy implies that perceived depth is proportional to the square of perceptual distance. However, the function relating the uncertainties of depth and of size discrimination to distance is the same. Hence, depth and size constancy may be accounted for by the same underlying law.     

Anthropogenic noise affects risk assessment and attention: the distracted prey hypothesis

Many studies have focused on the effects of anthropogenic noise on animal communication, but only a few have looked at its effect on other behavioural systems. We designed a playback experiment to test the effect of noise on predation risk assessment. We found that in response to boat motor playback, Caribbean hermit crabs (Coenobita clypeatus) allowed a simulated predator to approach closer before they hid. Two hypotheses may explain how boat noise affected risk assessment: it masked an approaching predator''s sound; and/or it reallocated some of the crabs'' finite attention, effectively distracting them, and thus preventing them from responding to an approaching threat. We found no support for the first hypothesis: a silent looming object still got closer during boat motor playbacks than during silence. However, we found support for the attentional hypothesis: when we added flashing lights to the boat motor noise to further distract the hermit crabs, we were able to approach the crabs more closely than with the noise alone. Anthropogenic sounds may thus distract prey and make them more vulnerable to predation.     

Modulation discrimination interference and auditory grouping.

The detection of a change in the modulation pattern of a (target) carrier frequency, fc (for example a change in the depth of amplitude or frequency modulation, AM or FM) can be adversely affected by the presence of other modulated sounds (maskers) at frequencies remote from fc, an effect called modulation discrimination interference (MDI). MDI cannot be explained in terms of interaction of the sounds in the peripheral auditory system. It may result partly from a tendency for sounds which are modulated in a similar way to be perceptually 'grouped', i.e. heard as a single sound. To test this idea, MDI for the detection of a change in AM depth was measured as a function of stimulus variables known to affect perceptual grouping, namely overall duration and onset and offset asynchrony between the masking and target sounds. In parallel experiments, subjects were presented with a series of pairs of sounds, the target alone and the target with maskers, and were asked to rate how clearly the modulation of the target could be heard in the complex mixture. The results suggest that two factors contribute to MDI. One factor is difficulty in hearing a pitch corresponding to the target frequency. This factor appears to be strongly affected by perceptual grouping. Its effects can be reduced or abolished by asynchronous gating of the target and masker. The second factor is a specific difficulty in hearing the modulation of the target, or in distinguishing that modulation from the modulation of other sounds that are present. This factor has effects even under conditions promoting perceptual segregation of the target and masker.     

Pre-existing sensory biases in the spectral domain in frogs: empirical results and methodological considerations

In many species of anurans, advertisement calls excite only one of the two inner-ear organs. One prediction of the pre-existing bias hypothesis is that signal innovations that additionally excite the “untapped” organ will be more behaviorally effective than normal calls. However, recent studies have shown that females of three species with single-peaked calls that stimulate only the basilar papilla (BP) preferred single-peaked synthetic calls with a frequency typical of conspecific calls to two-peaked calls that also stimulated the amphibian papilla (AP). We report that in spring peepers (Pseudacris crucifer) that also produce single-peaked calls, females did not show a preference in choices between single-peaked and two-peaked synthetic calls. Thus, the addition of energy exciting the AP had a neutral effect on signal attractiveness. Together, these results are unsupportive of the pre-existing bias hypothesis. An alternative hypothesis is that positive fitness consequences of responding to sounds providing extraordinary spectral stimulation are required for a novel call to become established as a mate-attracting signal. Testing these ideas requires a taxonomically broader examination of responses to sounds with novel spectral complexity, and attention to some methodological details will improve the comparability of such studies.     

Stimulus Familiarity Affects Perceptual Restoration in the European Starling (Sturnus vulgaris)

Background

Humans can easily restore a speech signal that is temporally masked by an interfering sound (e.g., a cough masking parts of a word in a conversation), and listeners have the illusion that the speech continues through the interfering sound. This perceptual restoration for human speech is affected by prior experience. Here we provide evidence for perceptual restoration in complex vocalizations of a songbird that are acquired by vocal learning in a similar way as humans learn their language.

Methodology/Principal Findings

European starlings were trained in a same/different paradigm to report salient differences between successive sounds. The birds'' response latency for discriminating between a stimulus pair is an indicator for the salience of the difference, and these latencies can be used to evaluate perceptual distances using multi-dimensional scaling. For familiar motifs the birds showed a large perceptual distance if discriminating between song motifs that were muted for brief time periods and complete motifs. If the muted periods were filled with noise, the perceptual distance was reduced. For unfamiliar motifs no such difference was observed.

Conclusions/Significance

The results suggest that starlings are able to perceptually restore partly masked sounds and, similarly to humans, rely on prior experience. They may be a suitable model to study the mechanism underlying experience-dependent perceptual restoration.     

Phonological representations are unconsciously used when processing complex, non-speech signals

Neuroimaging studies of speech processing increasingly rely on artificial speech-like sounds whose perceptual status as speech or non-speech is assigned by simple subjective judgments; brain activation patterns are interpreted according to these status assignments. The naïve perceptual status of one such stimulus, spectrally-rotated speech (not consciously perceived as speech by naïve subjects), was evaluated in discrimination and forced identification experiments. Discrimination of variation in spectrally-rotated syllables in one group of naïve subjects was strongly related to the pattern of similarities in phonological identification of the same stimuli provided by a second, independent group of naïve subjects, suggesting either that (1) naïve rotated syllable perception involves phonetic-like processing, or (2) that perception is solely based on physical acoustic similarity, and similar sounds are provided with similar phonetic identities. Analysis of acoustic (Euclidean distances of center frequency values of formants) and phonetic similarities in the perception of the vowel portions of the rotated syllables revealed that discrimination was significantly and independently influenced by both acoustic and phonological information. We conclude that simple subjective assessments of artificial speech-like sounds can be misleading, as perception of such sounds may initially and unconsciously utilize speech-like, phonological processing.     

Reliability of Health-Related Physical Fitness Tests among Colombian Children and Adolescents: The FUPRECOL Study

Substantial evidence indicates that youth physical fitness levels are an important marker of lifestyle and cardio-metabolic health profiles and predict future risk of chronic diseases. The reliability physical fitness tests have not been explored in Latino-American youth population. This study’s aim was to examine the reliability of health-related physical fitness tests that were used in the Colombian health promotion “Fuprecol study”. Participants were 229 Colombian youth (boys n = 124 and girls n = 105) aged 9 to 17.9 years old. Five components of health-related physical fitness were measured: 1) morphological component: height, weight, body mass index (BMI), waist circumference, triceps skinfold, subscapular skinfold, and body fat (%) via impedance; 2) musculoskeletal component: handgrip and standing long jump test; 3) motor component: speed/agility test (4x10 m shuttle run); 4) flexibility component (hamstring and lumbar extensibility, sit-and-reach test); 5) cardiorespiratory component: 20-meter shuttle-run test (SRT) to estimate maximal oxygen consumption. The tests were performed two times, 1 week apart on the same day of the week, except for the SRT which was performed only once. Intra-observer technical errors of measurement (TEMs) and inter-rater (reliability) were assessed in the morphological component. Reliability for the Musculoskeletal, motor and cardiorespiratory fitness components was examined using Bland–Altman tests. For the morphological component, TEMs were small and reliability was greater than 95% of all cases. For the musculoskeletal, motor, flexibility and cardiorespiratory components, we found adequate reliability patterns in terms of systematic errors (bias) and random error (95% limits of agreement). When the fitness assessments were performed twice, the systematic error was nearly 0 for all tests, except for the sit and reach (mean difference: -1.03% [95% CI = -4.35% to -2.28%]. The results from this study indicate that the “Fuprecol study” health-related physical fitness battery, administered by physical education teachers, was reliable for measuring health-related components of fitness in children and adolescents aged 9–17.9 years old in a school setting in Colombia.     

The initial phase of auditory and visual scene analysis

Auditory streaming and visual plaids have been used extensively to study perceptual organization in each modality. Both stimuli can produce bistable alternations between grouped (one object) and split (two objects) interpretations. They also share two peculiar features: (i) at the onset of stimulus presentation, organization starts with a systematic bias towards the grouped interpretation; (ii) this first percept has 'inertia'; it lasts longer than the subsequent ones. As a result, the probability of forming different objects builds up over time, a landmark of both behavioural and neurophysiological data on auditory streaming. Here we show that first percept bias and inertia are independent. In plaid perception, inertia is due to a depth ordering ambiguity in the transparent (split) interpretation that makes plaid perception tristable rather than bistable: experimental manipulations removing the depth ambiguity suppressed inertia. However, the first percept bias persisted. We attempted a similar manipulation for auditory streaming by introducing level differences between streams, to bias which stream would appear in the perceptual foreground. Here both inertia and first percept bias persisted. We thus argue that the critical common feature of the onset of perceptual organization is the grouping bias, which may be related to the transition from temporally/spatially local to temporally/spatially global computation.     

Imprinting on time-structured acoustic stimuli in ducklings

Filial imprinting is a dedicated learning process that lacks explicit reinforcement. The phenomenon itself is narrowly heritably canalized, but its content, the representation of the parental object, reflects the circumstances of the newborn. Imprinting has recently been shown to be even more subtle and complex than previously envisaged, since ducklings and chicks are now known to select and represent for later generalization abstract conceptual properties of the objects they perceive as neonates, including movement pattern, heterogeneity and inter-component relationships of same or different. Here, we investigate day-old Mallard (Anas platyrhynchos) ducklings’ bias towards imprinting on acoustic stimuli made from mallards’ vocalizations as opposed to white noise, whether they imprint on the temporal structure of brief acoustic stimuli of either kind, and whether they generalize timing information across the two sounds. Our data are consistent with a strong innate preference for natural sounds, but do not reliably establish sensitivity to temporal relations. This fits with the view that imprinting includes the establishment of representations of both primary percepts and selective abstract properties of their early perceptual input, meshing together genetically transmitted prior pre-dispositions with active selection and processing of the perceptual input.     

Motor Cost Influences Perceptual Decisions

Perceptual decision making has been widely studied using tasks in which subjects are asked to discriminate a visual stimulus and instructed to report their decision with a movement. In these studies, performance is measured by assessing the accuracy of the participants’ choices as a function of the ambiguity of the visual stimulus. Typically, the reporting movement is considered as a mere means of reporting the decision with no influence on the decision-making process. However, recent studies have shown that even subtle differences of biomechanical costs between movements may influence how we select between them. Here we investigated whether this purely motor cost could also influence decisions in a perceptual discrimination task in detriment of accuracy. In other words, are perceptual decisions only dependent on the visual stimulus and entirely orthogonal to motor costs? Here we show the results of a psychophysical experiment in which human subjects were presented with a random dot motion discrimination task and asked to report the perceived motion direction using movements of different biomechanical cost. We found that the pattern of decisions exhibited a significant bias towards the movement of lower cost, even when this bias reduced performance accuracy. This strongly suggests that motor costs influence decision making in visual discrimination tasks for which its contribution is neither instructed nor beneficial.     

Early activation of adult organ differentiation during delay of metamorphosis in solitary ascidians, and consequences for juvenile growth

Abstract. Many animals have the ability to delay metamorphosis when conditions are unfavorable. This strategy carries obvious benefits, but may also result in severe consequences for lecithotrophic larvae that run low on time and energy. Precocious activation of postlarval developmental programs—so-called anticipatory development—may be adaptive and increase the survival of older, energy-depleted larvae by allowing more rapid metamorphosis. Three of six solitary ascidian species displayed extensive anticipatory development of postlarval structures, similar to heterochronies normally observed for colonial species. The capacity for anticipatory development may be linked to the length of competent period, taxonomic group, or both: members of suborder Phlebobranchia exhibited extensive anticipatory development and long competent periods, but members of suborder Stolidobranchia exhibited little or no anticipatory development and had shorter competent periods. Delay of metamorphosis of up to 3 d did not negatively impact postlarval and juvenile growth rates for any of three species tested, regardless of taxonomic group or length of competent period, although a longer, 7-d, delay resulted in slower postlarval growth in Ciona intestinalis . Anticipatory development of postlarval structures may ameliorate the negative consequences of delay of metamorphosis in C. intestinalis and Ascidiella aspersa , but Molgula socialis showed neither anticipatory development nor a negative impact of metamorphic delay on postlarval fitness. This is the first demonstration that anticipatory development of postlarval structures, normally associated only with colonial ascidians, can occur as a normal part of the development of solitary ascidians.     

Reducing environmental bias when measuring natural selection

Abstract.— Crucial to understanding the process of natural selection is characterizing phenotypic selection. Measures of phenotypic selection can be biased by environmental variation among individuals that causes a spurious correlation between a trait and fitness. One solution is analyzing genotypic data, rather than phenotypic data. Genotypic data, however, are difficult to gather, can be gathered from few species, and typically have low statistical power. Environmental correlations may act through traits other than through fitness itself. A path analytic framework, which includes measures of such traits, may reduce environmental bias in estimates of selection coefficients. We tested the efficacy of path analysis to reduce bias by re-analyzing three experiments where both phenotypic and genotypic data were available. All three consisted of plant species (Impatiens capensis, Arabidopsis thaliana , and Raphanus sativus) grown in experimental plots or the greenhouse. We found that selection coefficients estimated by path analysis using phenotypic data were highly correlated with those based on genotypic data with little systematic bias in estimating the strength of selection. Although not a panacea, using path analysis can substantially reduce environmental biases in estimates of selection coefficients. Such confidence in phenotypic selection estimates is critical for progress in the study of natural selection.