Call-note discriminations in black-capped chickadees (Poecile atricapillus).

Bioacousticians (M. S. Ficken, S. R. Ficken, & S. R. Witken, 1978) classified black-capped chickadee call notes from the chick-a-dee call complex into 4 note types (A, B, C, and D) identified from sound spectrograms. In Experiment 1, chickadees (Poecile atricapillus) learned operant auditory discriminations both within and between the 4 note types but learned the between note-type discrimination significantly faster. In Experiment 2, when the original, unrewarded between-category exemplars were replaced with novel, rewarded exemplars of these same categories, chickadees showed transfer of inhibitory stimulus control to the novel exemplars. In Experiment 3, when novel exemplars were replaced by the original exemplars, chickadees showed propagation of positive stimulus control back to the original exemplars. This evidence suggests that chickadees and bioacousticians accurately sort conspecific call notes into the same open-ended categories (R. J. Hernstein, 1990). (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Discrimination of auditory distance cues by black-capped chickadees (Poecile atricapillus) and zebra finches (Taeniopygia guttata).

The authors examined auditory distance perception using a go/no-go operant discrimination task in the laboratory. They taught male black-capped chickadees (Poecile atricapillus) and male zebra finches (Taeniopygia guttata) to discriminate degraded (far) from undegraded (near) versions of male chickadee songs and female zebra finch calls, showing for the 1st time that males can discriminate distance cues in heterospecific vocalizations and in female calls. Chickadees learned faster than zebra finches, and both species learned to discriminate chickadee songs faster than zebra finch calls. Chickadees more than zebra finches attended to amplitude in tests pitting it against other distance cues, demonstrating that amplitude is a potentially useful cue for estimating distance from vocalizations. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Mechanisms of call note-type perception in black-capped chickadees (Poecile atricapillus): Peak shift in a note-type continuum.

We report on operant conditioning and artificial neural network (ANN) simulations aimed at further elucidating mechanisms of black-capped chickadee chick-a-dee call note category perception. Specifically, we tested for differences in the speed of acquisition among different discrimination tasks and, in two selected discrimination groups, searched for evidence of peak shift. Earlier, unreported ANN data were instrumental in providing the motivation for the current set of studies with chickadees and are provided here. The ANNs revealed differences in the speed of learning among note-type discrimination groups that is related to the degree of perceptual similarity among the three note types tested (i.e., A, B, and C notes). In many respects, bird and network results were in agreement (i.e., in the observation of peak shift in the same group), but they also differed in important ways (i.e., all discrimination groups showed differences in speed of learning in simulations but not in chickadees). We suggest that the start, peak and end frequency of the chick-a portion of chick-a-dee call notes, which form a graded but overlapping continuum, may drive the peak shift observed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Black-capped chickadees (Poecile atricapillus) anticipate future outcomes of foraging choices.

In 2 experiments we investigated the cognitive abilities of wild-caught black-capped chickadees (Poecile atricapillus) in future anticipation tasks. Chickadees were sensitive to anticipatory contrast effects over time horizons of 5, 10, and 30 min (Experiment 1). Chickadees also learned the order of events and anticipated that the quality of future foraging outcomes was contingent on current foraging choices. This behavior was demonstrated while foraging in a naturalistic aviary environment with a 30-min delay between the initial choice and the future outcome (Experiment 2). These results support the hypothesis that black-capped chickadees can cognitively travel in time both retrospectively and prospectively using episodic memory. This result shows the occurrence of anticipatory cognition in a noncorvid species of food-storing bird and supports the idea that cognitive time travel may have evolved in nonhuman animals in response to specific ecological selection pressures. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Neuron Production, Neuron Number, and Structure Size Are Seasonally Stable in the Hippocampus of the Food-Storing Black-Capped Chickadee (Poecile atricapillus).

Neuronal recruitment in the black-capped chickadee (Poecile atricapillus) hippocampus occurs at a higher rate in the fall than at other times of the year. As a means of determining whether this increase in recruitment results from greater neuron production, chickadees were caught in the wild between October and March and injected with the cell-birth marker 5-bromo-2'-deoxyuridine. Two weeks later, birds were killed by overdose, and hippocampal neuron production, apoptosis, neuron number, and hippocampal volume were determined. Chickadees collected in October, November, January, February, and March did not differ in neuron production, apoptosis, hippocampal volume, or neuron number. These findings indicate that increases in neuronal recruitment in the chickadee hippocampus in the fall do not result from increased neuron production, but instead, enhanced survival of new neurons. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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

14 captive black-capped chickadees were presented with normal and altered versions of their species-specific "fee bee" song to determine how note type, number, and sequence affect recognition. Perch changes and vocalizations given in response to playback did not differ reliably as a function of song type, whereas latency to 1st vocalization after playback did. In Exp I, using 2-note songs, Ss vocalized sooner to songs beginning with fee than with bee and to fee bee than to fee fee. In Exp II, Ss were presented with single-note, normal, and 3-note songs each consisting of a single-note type. Habituation slowed responding to altered songs but not to fee bee over 3 test sessions. Results suggest that Ss distinguished (a) single fees and 3-note songs from normal song, (b) single fees from single bees, (c) 2-note songs from 3-note songs, and (d) normal song from altered songs. It is concluded that the internal representation of conspecific song in the chickadee 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. (34 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Mechanisms of what-where-when memory in black-capped chickadees (Poecile atricapillus): Do chickadees remember “when”?

Integrated what-where-when memory, sometimes called 'episodic-like' memory, has been shown to occur in a number of animals, including food-storing birds and rodents. It is not always clear in these studies, however, what aspect of “when” is remembered. We examined memory for what, where, and when in black-capped chickadees Poecile atricapillus in a procedure designed to dissociate memory for elapsed time (how long ago) from memory for a point in time, the sense of “when” implied in most discussions of episodic or episodic-like memory. Chickadees searched for food in multiple trials that each involved a two-phase procedure. In Phase 1, birds found preferred mealworms in half the sites on one side of an aviary and less preferred sunflower seeds in half the sites on the other side of the aviary. In Phase 2, following a retention interval, birds searched in the aviary with fresh sunflower seeds placed in the sites that held seeds in Phase 1 but either fresh or degraded mealworms in the sites that held mealworms in Phase 1. Whether a site held fresh or degraded worms depended on either when during the day Phase 1 had occurred (Group When), how long ago Phase 1 had occurred (Group HLA), or both (Group When + HLA). Chickadees in all three groups were able to discriminate where and when palatable worms were to be found, but there were temporal limitations on the use of HLA and When + HLA cues that were not found for the use of When cues. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

A test of the adaptive specialization hypothesis: Population differences in caching, memory, and the hippocampus in black-capped chickadees (Poecile atricapilla).

To test the hypothesis that accurate cache recovery is more critical for birds that live in harsh conditions where the food supply is limited and unpredictable, the authors compared food caching, memory, and the hippocampus of black-capped chickadees (Poecile atricapilla) from Alaska and Colorado. Under identical laboratory conditions, Alaska chickadees (a) cached significantly more food; (b) were more efficient at cache recovery; (c) performed more accurately on one-trial associative learning tasks in which birds had to rely on spatial memory, but did not differ when tested on a nonspatial version of this task; and (d) had significantly larger hippocampal volumes containing more neurons compared with Colorado chickadees. The results support the hypothesis that these population differences may reflect adaptations to a harsh environment. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hippocampal growth and maintenance depend on food-caching experience in juvenile mountain chickadees (Poecile gambeli).

This experiment investigated the development of caching behavior and the hippocampus (HF) in postfledging mountain chickadees (Poecile gambeli). From Days 35 to 53, the number of seeds stored increased but the proportion recovered did not. Birds that stored and recovered during 3 or more trials had significantly enlarged HF but not telencephalon volumes (experienced) compared with those that stored but did not recover (store only) and those deprived of caching experience altogether (deprived). HF size did not increase linearly with the number of experience trials. Birds that received less than 3 experience trials did not differ from deprived birds in HF size, suggesting a threshold effect. Experienced birds prevented from caching for 1 month had significantly smaller HF volumes than those examined immediately after caching experience and did not differ from deprived birds. Experience of both storing and recovery is required to initiate growth and maintain HF size. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Pitch discrimination in field- and isolation-reared black-capped chickadees (Parus atricapillus).

Prior research has shown that male black-capped chickadees (Parus atricapillus) raised in the field produce constant relative pitch cues (frequency ratios) in their songs, whereas males reared in isolation from adult song do not. In this study, the authors found that field-reared male chickadees needed fewer than half as many sessions to learn an operant (go/no-go) auditory discrimination that linked S+ note pairs with a constant-frequency ratio than a discrimination that varied the frequency ratio of S+ pairs randomly. Most important, isolation-reared males needed over 5 times as many sessions to learn the constant-ratio discrimination compared with field-reared males. This is, to the authors' knowledge, the first report of impaired pitch discrimination in isolate songbirds. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Long-Term Memory for Spatial Locations in a Food-Storing Bird (Poecile atricapilla) Requires Activation of NMDA Receptors in the Hippocampal Formation During Learning.

Food-storing birds use a form of long-term memory to recover their hidden food caches that depends on the hippocampal formation (HF). The authors assessed whether food-storing birds' long-term memory for spatial locations requires N-methyl-D-aspartate receptor (NMDA-R)-dependent synaptic plasticity. Black-capped chickadees (Poecile atricapilla) were given bilateral infusions of the NMDA-R antagonist AP5 into the hippocampus, and their memory on a spatial reference memory task was assessed. NMDA-R inactivation during learning prevented formation of long-term spatial memories but did not affect short-term memory and retrieval processes. NMDA-R inactivation immediately following learning did not disrupt long-term memory formation. NMDA-R inactivation disrupted the learning of multiple serially encoded reward locations when a 180-min delay separated successive learning episodes, suggesting that NMDA-R activity has a role in the incorporation of new information into existing long-term memory, as well as in forming unitary long-term memories. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Alarm call responsivity of mallard ducklings (Anas platyrhynchos): IV. Effects of social experience.

In previous experiments on alarm call responsivity, mallard ducklings have been reared in a partially isolated condition in individual boxes that facilitate individual identification. The present study was designed to determine whether a more natural social rearing condition affects freezing behavior. In 3 experiments with mallard ducklings, rearing Ss socially in a 12-bird brood without a hen resulted in a significant reduction in alarm call responsivity. It is suggested that this reduction was caused by a decrease in the level of vocalizations uttered by socially reared Ss. Stimulating socially reared Ss with distress notes, contentment notes, or pre-exodus assembly calls reinstated high levels of freezing. (16 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Delayed figure reconstruction by a chimpanzee (Pan troglodytes) and humans (Homo sapiens).

A chimpanzee (Pan troglodytes) was trained to construct a copy of 3-element compound figures from a set of 9 elements. Delay intervals between sample offset and element presentation varied. The chimpanzee maintained accuracy at about 80% correct for a delay of 32 s, which was slightly higher than the mean of 4 human (Homo sapiens) Ss. Excellent visual reproductive memory in the chimpanzee as compared with that in humans was demonstrated. However, the nature of the reproductive memory was different in the 2 species in that humans better constructed meaningful figures, which represented food items, than meaningless ones, whereas the chimpanzee constructed these 2 types of figures with the same accuracy. This outcome suggests that the reproductive memory for meaningful figures of the chimpanzee may have been processed separately from symbolic processes. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Frequency ratio discrimination by zebra finches (Taeniopygia guttata) and humans (Homo sapiens).

We present the first evidence that songbirds can categorize stimulus sequences (note pairs) by frequency ratio (relative pitch) in an operant discrimination. Zebra finches (Taeniopygia guttata) and humans (Homo sapiens) were trained in a discrimination among 27 synthesized note pairs (9 S+ and 18 S– note pairs) with an operant go-no-go food-reward procedure. In the consistent-ratio discrimination groups, S+ note pairs had a common frequency ratio (1.12). In the jumbled-ratio discrimination group, S+ note pairs varied in frequency ratio (1.00, 1.12, and 1.26). Zebra finches and humans discriminated at least 8 of 9 consistent-ratio S+s, zebra finches discriminated 4 jumbled-ratio S+s, and humans discriminated few or no jumbled-ratio S+s. In this experiment, both species appeared to categorize note pairs by a consistent frequency ratio, but only zebra finches memorized several note pairs individually by rote. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Song-note discriminations in zebra finches (Taeniopygia guttata): Categories and pseudocategories.

Sturdy, Phillmore, and Weisman (1999) developed a simple, reliable procedure for classifying zebra finch (Taeniopygia guttata) song notes into types using exemplar notes from 4 captive North American laboratory colonies. In the present experiment, zebra finches learned true-category song-note discriminations based on Sturdy et al.'s note-type classification faster than pseudocategory discriminations based on random assortments of the same notes. Also, birds in the true-category discrimination were able to sort notes not heard during training into the same categories as the training notes. This evidence suggests that zebra finches accurately sort conspecific song notes into the same open-ended categories as Sturdy et al.'s human observers. (PsycINFO Database Record (c) 2010 APA, all rights reserved)