Perceptual organization of acoustic stimuli by budgerigars (Melopsittacus undulatus): II. Vocal signals

Operant conditioning and multidimensional scaling procedures were used to study auditory perception of complex sounds in the budgerigar. In a same-different discrimination task, budgerigars learned to discriminate among natural vocal signals. Multidimensional scaling procedures were used to arrange these complex acoustic stimuli in a two-dimensional space reflecting perceptual organization. Results show that budgerigars group vocal stimuli according to functional and acoustical categories. Studies with only contact calls show that birds also make within-category discriminations. The acoustic cues in contact calls most salient to budgerigars appear to be quite complex. There is a suggestion that the sex of the signaler may also be encoded in these calls. The results from budgerigars were compared with the results from humans tested on some of the same sets of complex sounds.     

Perceptual organization of acoustic stimuli by budgerigars (Melopsittacus undulatus): I. Pure tones

A new combination of operant conditioning and psychophysical scaling procedures was used to study auditory perception in a small bird. In a same-different discrimination task, budgerigars learned to discriminate among pure tones that varied along one or more acoustic dimensions. Response latencies were used to generate a matrix of interstimulus similarities. Multidimensional scaling procedures were used to arrange these acoustic stimuli in a multidimensional space that supposedly reflects the bird's perceptual organization. For tones that varied in intensity, duration, and frequency simultaneously, budgerigars were much more sensitive to frequency changes. From a set of tones that varied only in intensity, it was possible to calculate the growth of loudness with intensity for the budgerigar. For tones that varied only in frequency, budgerigars showed evidence of an "acoustic fovea" for frequency change in the spectral region of 2-4 kHz. Budgerigars and humans also differed in their perceptual grouping of tone sequences that rise, fall, or remain constant in pitch. Surprisingly, budgerigars were much less responsive to pitch contour than were humans.     

Speech perception by budgerigars (Melopsittacus undulatus): spoken vowels

Discrimination of natural, sustained vowels was studied in 5 budgerigars. The birds were trained using operant conditioning procedures on a same-different task, which was structured so that response latencies would provide a measure of stimulus similarity. These response latencies were used to construct similarity matrices, which were then analyzed by multidimensional scaling (MDS) procedures. MDS produced spatial maps of these speech sounds where perceptual similarity was represented by spatial proximity. The results of the three experiments suggest that budgerigars perceive natural, spoken vowels according to phonetic categories, find the acoustic differences among different talkers less salient than the acoustic differences among vowel categories, and use formant frequencies in making these complex discriminations.     

Perception of distance calls by budgerigars (Melopsittacus undulatus) and zebra finches (Poephila guttata): assessing species-specific advantages

Budgerigars (Melopsittacus undulatus) and zebra finches (Poephila guttata) share a common functional class of vocalizations called distance calls. The perception of species-specific distance calls by both species was measured with a habituation-dishabituation operant paradigm. Changes in discrimination performance were noted as listening conditions were modified or stimulus properties were altered. Both species showed better performance for calls of their own species. For zebra finches this tendency increased slightly when a background noise was added to the testing environment. Shifting the frequency region of the calls affected the discrimination performance of male budgerigars but not females or zebra finches. Reversing the temporal order of the calls affected the perceptual advantage for conspecific vocalizations in zebra finches but not in budgerigars. These results highlight species differences in perceiving acoustic communication signals.     

Auditory perception of conspecific and heterospecific vocalizations in birds: evidence for special processes.

Budgerigars (Melopsittacus undulatus), canaries (Serinus canaria), and zebra finches (Poephila guttata castanotis) were tested for their ability to discriminate among distance calls of each species. For comparison, starlings (Sturnus vulgaris) were tested on the same sounds. Response latencies to detect a change in a repeating background of sound were taken as a measure of the perceptual similarity among calls. All 4 species showed clear evidence of 3 perceptual categories corresponding to the calls of the 3 species. Also, budgerigars, canaries, and zebra finches showed an enhanced ability to discriminate among calls of their own species over the calls of the others. Starlings discriminated more efficiently among canary calls than among budgerigar or zebra finch calls. The results show species differences in discrimination of species-specific acoustic communication signals and provide insight into the nature of specialized perceptual processes.     

Acoustic and perceptual categories of vocal elements in the warble song of budgerigars (Melopsittacus undulatus)

The warble songs of budgerigars (Melopsittacus undulatus) are composed of a number of complex, variable acoustic elements that are sung by male birds in intimate courtship contexts for periods lasting up to several minutes. If these variable acoustic elements can be assigned to distinct acoustic-perceptual categories, it provides the opportunity to explore whether birds are perceptually sensitive to the proportion or sequential combination of warble elements belonging to different categories. By the inspection of spectrograms and by listening to recordings, humans assigned the acoustic elements in budgerigar warble from several birds to eight broad, overlapping categories. A neural-network program was developed and trained on these warble elements to simulate human categorization. The classification reliability between human raters and between human raters and the neural network classifier was better than 80% both within and across birds. Using operant conditioning and a psychophysical task, budgerigars were tested on large sets of these elements from different acoustic categories and different individuals. The birds consistently showed high discriminability for pairs of warble elements drawn from between acoustic categories and low discriminability for pairs drawn from within acoustic categories. With warble elements reliably assigned to different acoustic categories by humans and birds, it affords the opportunity to ask questions about the ordering of elements in natural warble streams and the perceptual significance of this ordering.     

Perception of warble song in budgerigars (Melopsittacus undulatus): evidence for special processing

The long, rambling warble song of male budgerigars is composed of a large number of acoustically complex elements uttered in streams lasting minutes a time and accompanied by various courtship behaviors. Warble song has no obvious sequential structure or patterned repetition of elements, raising questions as to which aspects of it are perceptually salient, whether budgerigars can detect changes in natural warble streams, and to what extent these capabilities are species-specific. Using operant conditioning and a psychophysical paradigm, we examined the sensitivity of budgerigars, canaries, and zebra finches to changes in long (>6?min) natural warble sequences of a male budgerigar. All three species could detect a single insertion of pure tones, zebra finch song syllables, budgerigar contact calls, or warble elements from another budgerigar’s warble. In each case, budgerigars were more sensitive to these changes than were canaries or finches. When warble elements from the ongoing warble stream were used as targets and inserted, out of order, into the natural warble stream so that the only cue available was the violation of the natural ordering of warble elements, only budgerigars performed above chance. When the experiment was repeated with all the ongoing warble stream elements presented in random order, the performance of budgerigars fell to chance. These results show species-specific advantages in budgerigars for detecting acoustic changes in natural warble sequences and indicate at least a limited sensitivity to sequential rules governing the structure of their species-specific warble songs.     

Effects of lesions of the central nucleus of the anterior archistriatum on contact call and warble song production in the budgerigar (Melopsittacus undulatus)

We studied the effects of both unilateral and bilateral lesions of the central nucleus of the anterior archistriatum (AAc) on the production of contact calls and warble song in adult male and female budgerigars. Birds were sorted into three experimental groups based on the percentage of AAc destroyed and whether lesions were unilateral or bilateral. The experimental groups were Unilateral Lesion (N = 8), Partial Bilateral Lesion (N = 5), and Bilateral Lesion birds (N = 12). Each group contained both sexes. Unilateral lesions had no demonstrable effects on contact call or warble song production. Bilateral lesions resulted in immediate and permanent disruption of all learned temporal and spectral characteristics of contact calls, although call initiation was not dependent on the AAc. Partial bilateral lesion effects varied with lesion size and location. At least 20-30% sparing of the AAc, including sparing portions of both the dorsal (AAcd) and ventral (AAcv) subdivisions on the same side of the brain, is necessary for production of prelesion contact call patterns. Warble song was absent in birds with complete bilateral destruction. Two birds with large yet incomplete lesions of the AAc sang after surgery, although the warble song of these birds was extremely impoverished and contained only a few of the typical warble song elements. Lesion results indicate that the AAc mediates the production of learned vocal features in male and female budgerigars, with each hemisphere capable of supporting a normal vocal repertoire.     

Competition between color salience and perceptual load during visual selection can be biased by top-down set

Visual salience and perceptual load may both influence the efficiency of visual selection. Recently, Gibson and Bryant (2008) showed that perceptual load can dominate color salience in a distractor interference paradigm. The purpose of the present study was to investigate the possibility that knowledge (of color or load) may modulate the relative operation of these two mechanisms. Consistent with previous findings, perceptual load dominated color salience, but only in certain contexts in which display load was mixed and high-load displays preceded other high-load displays. More important, color salience dominated perceptual load in other contexts in which display load was mixed and low-load displays preceded high-load displays. In addition, color salience also dominated perceptual load in contexts in which display load was fixed and advance knowledge of load was available. Altogether, the present findings suggest that the competition between color salience and perceptual load can vary as a function of task context, thereby supporting top-down accounts, although the precise aspect of task context remains to be identified.     

Spectral versus temporal features in dichotic listening

Ear advantage for the processing of dichotic speech sounds can be separated into two components. One of these components is an ear advantage for those phonetic features that are based on spectral acoustic cues. This ear advantage follows the direction of a given individual's ear dominance for the processing of spectral information in dichotic sounds, whether speech or nonspeech. The other factor represents a right-ear advantage for the processing of temporal information in dichotic sounds, whether speech or nonspeech. The present experiments were successful in dissociating these two factors. Since the results clearly show that ear advantage for speech is influenced by ear dominance for spectral information, a full understanding of the asymmetry in the perceptual salience of speech sounds in any individual will not be possible without knowing his ear dominance.     

Effects of deafening on the development of nestling and juvenile vocalizations in budgerigars (Melopsittacus undulatus).

The effects of complete and partial cochlear extirpation at ages 9-11 days posthatch were assessed in 5 nestling budgerigars (Melopsittacus undulatus) to determine if auditory feedback is necessary for the production of nestling vocalizations. Although early deafening had no effect on the production of food-begging calls produced during the first 2 weeks posthatch, deafening did disrupt the expected transition from these early calls to the longer and more complex frequency-modulated, patterned food-begging calls normally appearing 3-4 weeks posthatch. All birds sustaining either complete or partial cochlear extirpation failed to develop stereotyped contact calls around the time of fledging at 5 weeks. These results are consistent with previous research showing that deafened nestlings do not develop normal contact calls (R. J. Dooling, B. F. Gephart, P. H. Price, C. McHale, & S. E. Brauth, 1987) and also indicate that a form of sensorimotor learning is involved in the production of mature, patterned food-begging calls in budgerigars.     

Symmetry perception in the blind

Bilateral mirror symmetry, especially vertical symmetry, is a powerful phenomenon in spatial organization of visual shapes. However, the causes of vertical symmetry salience in visual perception are not completely clear. Here we investigated whether the perceptual salience of vertical symmetry depends on visual experience by testing a group of congenitally blind individuals in a memory task in which either horizontal or vertical symmetry was used as an incidental feature. Both blind and sighted subjects remembered more accurately configurations that were symmetrical compared to those that were not. Critically, whereas sighted subjects displayed a higher level of facilitation by vertical than horizontal symmetry, no such difference was found in the blind. This suggests that the perceptual salience of the vertical dimension is visually based.     

Trading relations in the perception of speech by 5-year-old children

Five-year-old children were tested for perceptual trading relations between a temporal cue (silence duration) and a spectral cue (F₁ onset frequency) for the “say-stay” distinction. Identification functions were obtained for two synthetic “say-stay” continua, each containing systematic variations in the amount of silence following the /s/ noise. In one continuum, the vocalic portion had a lower F₁ onset than in the other continuum. Children showed a smaller trading relation than has been found with adults. They did not differ from adults, however, in their perception of an “ay-day” continuum formed by varying F₁ onset frequency only. The results of a discrimination task in which the two acoustic cues were made to “cooperate” or “conflict” phonetically supported the notion of perceptual equivalence of the temporal and spectral cues along a single phonetic dimension. The results indicate that young children, like adults, perceptually integrate multiple cues to a speech contrast in a phonetically relevant manner, but that they may not give the same perceptual weights to the various cues as do adults.     

Assessing clinically relevant perceptual organization with multidimensional scaling techniques

Multidimensional scaling (MDS) techniques provide a promising measurement strategy for characterizing individual differences in cognitive processing, which many clinical theories associate with the development, maintenance, and treatment of psychopathology. The authors describe the use of deterministic and probabilistic MDS techniques for investigating numerous aspects of perceptual organization, such as dimensional attention, perceptual correlation, within-attribute organization, and perceptual variability. Additionally, they discuss how formal quantitative models can be used, in conjunction with MDS-derived representations of individual differences in perceptual organization, to test theories about the role of cognitive processing in clinically relevant phenomena. They include applied examples from their work in the areas of eating disorders and sexual coercion.     

Differential vocalization in budgerigars: towards an experimental analysis of naming.

In Experiment 1, 3 budgerigars (Melopsittacus undulatus) were trained with food reinforcement to make low- or high-frequency calls in response to different color stimuli, C1 and C2 (a color-naming task), using a gradual response-differentiation procedure and an automatic call-recognition system. Thus, a call within a certain frequency band was reinforced in the presence of C1 ("C1 call"), and a call within a different band was reinforced in the presence of C2 ("C2 call"). In Experiment 2, all 3 budgerigars were trained in a form-to-color matching-to-sample task, alternating trial by trial with either the color-naming task (2 birds) or an identity color matching-to-sample task (1 bird). Sample stimuli for the new matching-to-sample task were forms (F1 or F2) and comparisons were the same two colors (C1 and C2). Given Sample F1 or F2, birds had to make a call to produce Comparison Pair C1 and C2. With F1 as the sample, a peck on C1 was reinforced; with F2 as the sample, a peck on C2 was reinforced. Although no particular call was specified in the presence of F1 and F2, 2 birds made the C1 call in the presence of F1 and the C2 call in the presence of F2. In Experiment 3, the bird that failed to match form and color calls in Experiment 2 and another bird were first trained in a color-to-form matching-to-sample task: C1 to F3 and C2 to F4. In this task, to produce the comparison pair of forms, a high call (or low for the other bird) was required in the presence of C1, and a low call (or high) was required in the presence of C2. Both birds were then trained with an identity matching-to-sample task in which sample and comparison stimuli were the same two forms, F3 and F4. Trials on the identity task alternated with the color-to-form trials. Although no particular call was required in the presence of Samples F3 and F4, both birds came to make the C1 call in the presence of F3 and the C2 call in the presence of F4. Our technique promises to be useful for the study of emergent vocal relations in budgerigars and other animals.     

Auditory grouping based on fundamental frequency and formant peak frequency

The perceptual grouping of a four-tone cycle was studied as a function of differences in fundamental frequencies and the frequencies of spectral peaks. Each tone had a single formant and at least 13 harmonics. In Experiment 1 the formant was created by filtering a flat spectrum and in Experiment 2 by adding harmonics. Fundamental frequency was found to be capable of controlling grouping even when the spectra spanned exactly the same frequency range. Formant peak separation became more effective as the sharpness (amplitude of the peak relative to a spectral pedestal) increased. The effect of each type of acoustic difference depended on the task. Listeners could group the tones by either sort of difference but were also capable of resisting the disruptive effect of the other one. This was taken as evidence for the presence of a schema-based process of perceptual grouping and the relative weakness of primitive segregation.     

Why does a red snake in the grass capture your attention?

It has been suggested that affective stimuli automatically capture attention; this preferential processing is thought to be related to the evolutionary significance of affective stimuli. However, recent evidence suggests that perceptual salience alone might explain why some affective stimuli are more likely to influence attentional processes in certain contexts. In this study, we manipulated affective and perceptual salience to better understand how affective information is processed and how it impacts attentional processes in different contexts. We used stimuli that are both affectively and perceptually salient, while varying the task requirement to encourage the processing of perceptually salient (Experiment 1) or affectively salient (Experiment 2) information. This design made it possible to observe independent and interdependent relationships between perceptual and affective salience. The results showed that when the task encouraged the processing of perceptually salient information, affective salience did not influence task performance. In contrast, when the task encouraged the processing of affectively salient information, affectively salient information impaired task performance. The findings suggest that the affective nature of the stimuli does not always influence attentional processes.     

Pitch and loudness interact in auditory displays: can the data get lost in the map?

Many auditory displays use acoustic attributes such as frequency, intensity, and spectral content to represent different characteristics of multidimensional data. This study demonstrated a perceptual interaction between dynamic changes in pitch and loudness, as well as perceived asymmetries in directional acoustic change, that distorted the data relations represented in an auditory display. Three experiments showed that changes in loudness can influence pitch change, that changes in pitch can influence loudness change, and that increases in acoustic intensity are judged to change more than equivalent decreases. Within a sonification of stock market data, these characteristics created perceptual distortions in the data set. The results suggest that great care should be exercised when using lower level acoustic dimensions to represent multidimensional data.     

The prioritization of perceptual processing in categorization

In three experiments, the effects of selective attention on perceptual processes in a complex multidimensional object categorization task were investigated. In each experiment, participants completed a perceptual-matching task to gain estimates of the perceptual salience of each stimulus dimension, then a categorization task using the same stimuli. In Experiments 1 and 2, the perceptual processing of stimulus dimensions was faster when dimensions were more diagnostic of category membership, regardless of their perceptual salience. Experiment 3 demonstrated that this prioritization of perceptual processing was evident even when stimuli were presented in unpredictable locations during categorization, indicating that the physical characteristics of the stimulus guide selective attention to diagnostic stimulus dimensions.     

Frequency-shift detectors bind binaural as well as monaural frequency representations

Previous psychophysical work provided evidence for the existence of automatic frequency-shift detectors (FSDs) that establish perceptual links between successive sounds. In this study, we investigated the characteristics of the FSDs with respect to the binaural system. Listeners were presented with sound sequences consisting of a chord of pure tones followed by a single test tone. Two tasks were performed. In the "present/absent" task, the test tone was either identical to one of the chord components or positioned halfway in frequency between two components, and listeners had to discriminate between these two possibilities. In the "up/down" task, the test tone was slightly different in frequency from one of the chord components and listeners had to identify the direction (up or down) of the corresponding shift. When the test tone was a pure tone presented monaurally, either to the same ear as the chord or to the opposite ear, listeners performed the up/down task better than the present/absent task. This paradoxical advantage for directional frequency shifts, providing evidence for FSDs, persisted when the test tone was replaced by a dichotic stimulus consisting of noise but evoking a pitch sensation as a consequence of binaural processing. Performance in the up/down task was similar for the dichotic stimulus and for a monaural narrow-band noise matched in pitch salience to it. Our results indicate that the FSDs are insensitive to sound localization mechanisms and operate on central frequency representations, at or above the level of convergence of the monaural auditory pathways.