Frequency-range discriminations and absolute pitch in black-capped chickadees (Poecile atricapillus), mountain chickadees (Poecile gambeli), and zebra finches (Taeniopygia guttata)

The acoustic frequency ranges in birdsongs provide important absolute pitch cues for the recognition of conspecifics. Black-capped chickadees (Poecile atricapillus), mountain chickadees (Poecile gambeli), and zebra finches (Taeniopygia guttata) were trained to sort tones contiguous in frequency into 8 ranges on the basis of associations between response to the tones in each range and reward. All 3 species acquired accurate frequency-range discriminations, but zebra finches acquired the discrimination in fewer trials and to a higher standard than black-capped or mountain chickadees, which did not differ appreciably in the discrimination. Chickadees' relatively poorer accuracy was traced to poorer discrimination of tones in the higher frequency ranges. During transfer tests, the discrimination generalized to novel tones when the training tones were included, but not when they were omitted.     

Comparing black-capped (Poecile atricapillus) and mountain chickadees (Poecile gambeli): use of geometric and featural information in a spatial orientation task

Since Cheng (Cognition 23:149–178, 1986) first proposed the “geometric module” in rats, a great deal of research has focused on how other species use geometric information and how geometric encoding may differ across species. Here, hand-reared and wild-caught black-capped chickadees and wild-caught mountain chickadees searched for food hidden in one corner in a rectangular environment. Previous research has shown that mountain chickadees do not spontaneously encode geometric information when a salient feature is present near the goal location. Using a slightly different training and testing procedure, we found that both hand-reared and wild-caught black-capped chickadees encoded geometric information, even in the presence of a salient landmark. Some, but not all, mountain chickadees also encoded geometric information. Overall, our results suggest that use of geometric information may be a less preferred strategy for mountain chickadees than for either wild-caught or hand-reared black-capped chickadees. To our knowledge, this is the first direct interspecies comparison of use of geometric information in a spatial orientation task.     

Discrimination of individual vocalizations by black-capped chickadees (Poecile atricapilla)

The auditory perceptual abilities of male black-capped chickadees (Poecile atricapilla) were examined using an operant go/no-go discrimination among 16 individual vocalizations recorded at 5 m. The birds learned to discriminate about equally well among eight male chickadee fee-bee songs and eight female zebra finch (Taeniopygia guttata) distance calls. These results do not indicate that chickadees have a species-specific advantage in individual recognition for conspecific over heterospecific vocalizations. We then transferred the chickadees to a discrimination of the same songs and calls rerecorded at a moderate distance. These results showed accurate transfer of discrimination from 16 vocalizations recorded at 5 m to novel versions of the same 16 songs and calls rerecorded at 25 m. That is, chickadees recognized individual songs and calls despite degradation produced by rerecording at 25 m. Identifying individual vocalizations despite their transformation by distance cues is here described as a biologically important example of perceptual constancy.     

Open-ended categorization of chick-a-dee calls by black-capped chickadees (Poecile atricapilla)

The authors trained black-capped chickadees (Poecile atricapilla) in an operant discrimination with exemplars of black-capped and Carolina chick-a-dee calls, with the goal of determining whether the birds memorized the calls of conspecifics and heterospecifics or classified the calls by species. Black-capped calls served as both rewarded (S+) and unrewarded (S-) stimuli (the within-category discrimination), whereas Carolina chick-a-dee calls served as S-s (the between-category discrimination) in the black-capped chick-a-dee call S+ group. The Carolina call S+ group had Carolina calls as S+s and S-s (within-category) and black-capped calls as S-s (between-category). Both groups discriminated between call categories faster than within a call category. In 2 subsequent experiments, both S+ groups showed transfer to novel calls and propagation back to between-category calls. The results favor the hypothesis that the acoustically similar social calls of the 2 species constitute separate open-ended categories.     

Statistical classification of black-capped (Poecile atricapillus) and mountain chickadee (Poecile gambeli) call notes

Both black-capped (Poecile atricapillus) and mountain chickadees (Poecile gambeli) produce a chick-a-dee call that consists of several distinct note types. In some regions, these 2 species live sympatrically, and it has been shown that 1 species will respond weakly to songs of the other. This suggests that chickadee song, and potentially other of their vocalizations, contains species-specific information. We tested the possibility that call notes were acoustically sufficient for species identification. Black-capped and mountain non-D notes were summarized as a set of 9 features and then analyzed by linear discriminant analysis. Linear discriminant analysis was able to use these notes to identify species with 100% accuracy. We repeated this approach, but with black-capped and mountain D notes that were summarized as a set of 4 features. Linear discriminant analysis was able to use these notes to identify species with 94% accuracy. This demonstrates that any of the note types in these chickadee calls possesses sufficient information for species classification.     

Acoustic mechanisms of note-type perception in black-capped chickadee (Poecile atricapillus) calls

Acoustic communication in black-capped chickadees (Poecile atricapillus) has been studied intensively, the "chick-a-dee" call being among the most well described. This call consists of 4 note types; chickadees perceive these notes as open-ended categories and do so in a continuous manner, with As more similar to Bs and Bs more similar to Cs. Acoustic features contributing to the note-type differentiation are unknown. Recent analyses suggested that certain acoustic features may play a role in note-type classification. Here, the authors tested black-capped chickadees in an operant-conditioning paradigm to determine which features were controlling note-type perception. The results suggest that the note pitch and the frequency modulation in the initial portion of the note control the perception of note types.     

Memory for what, where, and when in the black-capped chickadee (Poecile atricapillus)

Episodic memory is the ability to remember personally experienced past events (Tulving in Organization of memory. Academic Press, San Diego, pp. 381–403, 1972). In non-human animals, the behavioural criterion for episodic-like memory is remembering “what” occurred in conjunction with “when” and “where” (Clayton and Dickinson in Nature 395:272–274, 1998). We conducted tests for “what, where, and when” memory in a food-storing bird, the black-capped chickadee (Poecile atricapillus). In Experiment 1, chickadees found sunflower seeds and mealworms in concealed sites in their home cage. Birds later re-visited these sites after either a short (3 h) or long (123 h) retention interval. Chickadees normally prefer mealworms, but at the long retention interval mealworms were degraded in taste and appearance. Chickadees showed some memory for what kind of food they had encountered and where, but no memory for when food had previously been found. Experiment 2 followed a similar procedure, except that chickadees searched for hidden sunflower seeds and mealworms in trees in an indoor aviary. These more natural conditions increased both the spatial scale of the task and the effort required to find food. In this experiment, birds showed evidence for all three components of what–where–when memory. Unlike some previous studies of episodic-like memory, birds’ behaviour cannot be accounted for by differential memory strength for more recent events. The results show that memory for what, where, and when occurs in food-storing birds outside the corvid family, does not require food caching or retrieval, and that remembering “when” can depend on the nature of the task.     

Song sequence discrimination in the black-capped chickadee (Parus atricapillus)

Captive black-capped chickadees (Parus atricapillus) were presented with normal and altered versions of their species-specific "fee bee" song, to determine how note type, number, and sequence affect recognition. The number of perch changes and vocalizations (analyzed separately) given in response to playback did not differ reliably as a function of song type, whereas latency to first vocalization after playback did. In an initial experiment using two-note songs, birds vocalized sooner to songs beginning with fee than with bee and to fee bee than to fee fee. In a second experiment, birds were presented with shortened (single note), normal, and lengthened (three note) songs each consisting of a single-note type (either fee or bee). Habituation slowed responding to altered songs but not to fee bee over three test sessions. Results from the first session suggest that chickadees distinguished single fees and three-note songs from normal song, single fees from single bees, and two-note songs from three-note songs. Results from the third session suggested that chickadees distinguished normal song from any of the altered songs. The internal representation of conspecific song in the chickadee thus distinguishes between fee and bee notes, contains information about note order, and is sensitive to note number. The pattern of responses is consistent with a model of recognition based on note-by-note integration of individual decisions about song structure.     

Divided attention,memory, and spatial discrimination in food-storing and nonstoring birds,black-capped chickadees (Poecile atricapilla) and dark-eyed juncos (Junco hyemalis)

Food-storing birds, black-capped chickadees (Poecile atricapilla), and nonstoring birds, dark-eyed juncos (Junco hyemalis), matched color or location on a touch screen. Both species showed a divided attention effect for color but not for location (Experiment 1). Chickadees performed better on location than on color with retention intervals up to 40 s, but juncos did not (Experiment 2). Increasing sample-distractor distance improved performance similarly in both species. Multidimensional scaling revealed that both use a Euclidean metric of spatial similarity (Experiment 3). When choosing between the location and color of a remembered item, food storers choose location more than do nonstorers. These results explain this effect by differences in memory for location relative to color, not division of attention or spatial discrimination ability.     

Black-capped chickadee (Poecile atricapillus) and human (Homo sapiens) chord discrimination

Human music perception is related both to musical experience and the physical properties of sound. Examining the processing of music by nonhuman animals has been generally neglected. We tested both black-capped chickadees and humans in a chord discrimination task that replicates and extends prior research with pigeons. We found that chickadees and humans, in common with pigeons, showed similar patterns of discrimination across manipulations of the 3rd and 5th notes of the triadic chords. For all species (chickadee and humans here, pigeons previously), chords with half-step alterations in the 5th note were easier to discriminate than half-step manipulations of the 3rd note, which is likely due to the sensory consonance of these chords. There were differences among species in terms of the fine discrimination of the chords within this larger pattern of results. Further, the ability to relearn the chords when transposed to a new root differed across species. Our results provide new comparative data suggesting some similarities in chord perception that span a wide range of species, from pigeons (nonvocal learners) to songbirds and humans (vocal learners).     

Information theoretical approaches to chick-a-dee calls of Carolina chickadees (Poecile carolinensis)

One aim of this study was to apply information theoretical analyses to understanding the structural complexity of chick-a-dee calls of Carolina chickadees, Poecile carolinensis. A second aim of this study was to compare this structural complexity to that of the calls of black-capped chickadees, P. atricapillus, described in an earlier published report (Hailman, Ficken, & Ficken, 1985). Chick-a-dee calls were recorded from Carolina chickadees in a naturalistic observation study in eastern Tennessee. Calls were analyzed using approaches from information theory, including transition probability matrices, Zipf's rules, entropies, and information coding capacities of calls and notes of calls. As described for black-capped chickadees, calls of Carolina chickadees exhibited considerable structural complexity. Most results suggested that the call of Carolina chickadees is more structurally complex than that of black-capped chickadees. These findings add support to the growing literature on the complexity of this call system in Paridae species. Furthermore, these results point to the feasibility of detailed cross-species comparative analyses that may allow strong testing of hypotheses regarding signal evolution.     

Discrimination of acoustically similar conspecific and heterospecific vocalizations by black-capped chickadees (<Emphasis Type="Italic">Poecile atricapillus</Emphasis>)

Chickadees produce a multi-note chick-a-dee call in multiple socially relevant contexts. One component of this call is the D note, which is a low-frequency and acoustically complex note with a harmonic-like structure. In the current study, we tested black-capped chickadees on a between-category operant discrimination task using vocalizations with acoustic structures similar to black-capped chickadee D notes, but produced by various songbird species, in order to examine the role that phylogenetic distance plays in acoustic perception of vocal signals. We assessed the extent to which discrimination performance was influenced by the phylogenetic relatedness among the species producing the vocalizations and by the phylogenetic relatedness between the subjects’ species (black-capped chickadees) and the vocalizers’ species. We also conducted a bioacoustic analysis and discriminant function analysis in order to examine the acoustic similarities among the discrimination stimuli. A previous study has shown that neural activation in black-capped chickadee auditory and perceptual brain regions is similar following the presentation of these vocalization categories. However, we found that chickadees had difficulty discriminating between forward and reversed black-capped chickadee D notes, a result that directly corresponded to the bioacoustic analysis indicating that these stimulus categories were acoustically similar. In addition, our results suggest that the discrimination between vocalizations produced by two parid species (chestnut-backed chickadees and tufted titmice) is perceptually difficult for black-capped chickadees, a finding that is likely in part because these vocalizations contain acoustic similarities. Overall, our results provide evidence that black-capped chickadees’ perceptual abilities are influenced by both phylogenetic relatedness and acoustic structure.     

Group size and social interactions are associated with calling behavior in Carolina chickadees (Poecile carolinensis)

The complexity of a social group may influence the vocal behavior of group members. Recent evidence in Carolina chickadees, Poecile carolinensis, indicated that one component of social complexity, group size, influenced the complexity of the "chick-a-dee" call, a vocalization functioning in social cohesion. Individuals in larger social groups used calls with greater information than did individuals in smaller social groups. Here, the authors review this earlier work, and describe a recent study indicating that social interactions between females and males within female-male pairs of chickadees were associated with rates of chick-a-dee call production in the males. Together, these studies suggest that the nature and complexity of social interactions among members of chickadee social groups influence chick-a-dee calling behavior.     

Imitation of conditional discriminations in pigeons (Columba livia)

In the present experiments, the 2-action method was used to determine whether pigeons could learn to imitate a conditional discrimination. Demonstrator pigeons (Columba livia) stepped on a treadle in the presence of 1 light and pecked at the treadle in the presence of another light. Demonstration did not seem to affect acquisition of the conditional discrimination (Experiment 1) but did facilitate its reversal of the conditional discrimination (Experiments 2 and 3). The results suggest that pigeons are not only able to learn a specific behavior by observing another pigeon, but they can also learn under which circumstances to perform that behavior. The results have implications for proposed mechanisms of imitation in animals.     

Perceptual learning in maze discriminations

In Experiment 1, rats were trained on a discrimination between rubber- and sandpaper-covered arms of a maze after one group had been pre-exposed to these intra-maze cues. Pre-exposure facilitated subsequent discrimination learning, unless the discrimination was made easier by adding further discriminative stimuli, when it now significantly retarded learning. In Experiment 2, rats were trained on an extra-maze spatial discrimination, again after one group, but not another, had been pre-exposed to the extra-maze landmarks. Here too, pre-exposure facilitated subsequent discrimination learning, unless the discrimination was made substantially easier by arranging that the two arms between which rats had to choose were always separated by 135°. The results of both experiments can be explained by supposing that perceptual learning depends on the presence of features common to S+ and S-.     

Within-category discriminations in speech perception

The assumption that listeners are unaware of the highly encoded acoustic properties which lead to phoneme identification is questioned in the present study. It was found that some subjects can make use of small differences in voice onset time when making within-category discriminations. Subjects who can use these auditory features do so both implicitly (in a phonetic match task) and deliberately (in a physical match task). Results also indicate that some type of parallel process model is needed to account for the processing of auditory and phonetic information.     

Absolute pitch: frequency-range discriminations in pigeons (Columba livia): comparisons with zebra finches (Taeniopygia guttata) and humans (Homo sapiens)

Absolute pitch (AP) is the ability to classify individual pitches without an external referent. The authors compared results from pigeons (Columba livia, a nonsongbird species) with results (R. Weisman, M. Njegovan, C. Sturdy, L. Phillmore, J. Coyle, & D. Mewhort, 1998) from zebra finches (Taeniopygia guttata, a songbird species) and humans (Homo sapiens) in AP tests that required classification of contiguous tones into 3 or 8 frequency ranges on the basis of correlations between the tones in each frequency range and reward. Pigeons' 3-range discriminations were similar in accuracy to those of zebra finches and humans. In the more challenging 8-range task, pigeons, like zebra finches, discriminated shifts from reward to nonreward from range to range across all 8 ranges, whereas humans discriminated only the 1st and last ranges. Taken together with previous research, the present experiments suggest that birds may have more accurate AP than mammals.     

Successive reversal of concurrent discriminations by macaques (Macaca mulatta): proactive interference effects

Rhesus monkeys received concurrent within-session training on eight, two-choice object pairs and then underwent successive reversals of these problems. Initially, reversals required about six times more training than acquisition with no improvement over seven successive reversals. Surprisingly, performance on these eight problems was unimpaired if they were embedded in different eight-problem tasks, thereby indicating a release from proactive interference. When the original eight problems again underwent successive reversal, no improvement was seen over seven reversals, although there was significantly less error-per-reversal than in the initial test. Subsequently, monkeys appeared to be developing a learning set for successive reversal because performance on successive reversal of eight novel problems was not different from that seen with the old familiar task. Set acquisition was confirmed when proficient reversal was eventually achieved on both old and new concurrent tasks. Thus, concurrent reversal set did develop, but it required arduous training to overcome proactive interference effects on memory. The ubiquitous influence of measurement context on organization of monkey memory was noted.     

Transfer of control in ambiguous discriminations

Rats' acquisition and transfer of performance in ambiguous discriminations was examined using discrete-trial operant lever-press procedures. Rats learned serial ambiguous discriminations (X----A+, A-; B+, X----B-) by acquiring both positive and negative occasion setting functions to the X feature. Mutual transfer occurred among cues from serial ambiguous, feature positive (Y----C+, C-), and feature negative (Y----C-, C+) discriminations, but that transfer did not extend to cues not trained within one of those serial discriminations. The ambiguous feature's positive occasion setting powers were unaffected by nonreinforced presentation of that feature alone, and posttraining counterconditioning did not eliminate the feature's ability to serve as a negative occasion setter. Occasion setting was not acquired with simultaneous ambiguous discriminations (XA+, A-; XB-, B+), which apparently were solved with a configural strategy.