Chickadees fail standardized operant tests for octave equivalence.

Octave equivalence occurs when an observer judges notes separated by a doubling in frequency perceptually similar. The octave appears to form the basis of pitch change in all human cultures and thus may be of biological origin. Previously, we developed a nonverbal operant conditioning test of octave generalization and transfer in humans. The results of this testing showed that humans with and without musical training perceive the octave relationship between pitches. Our goal in the current study was to determine whether black-capped chickadees, a North American songbird, perceive octave equivalence. We chose these chickadees because of their reliance on pitch in assessing conspecific vocalizations, our strong background knowledge on their pitch height perception (log-linear perception of frequency), and the phylogenetic disparity between them and humans. Compared to humans, songbirds are highly skilled at using pitch height perception to classify pitches into ranges, independent of the octave. Our results suggest that chickadees used that skill, rather than octave equivalence, to transfer the note-range discrimination from one octave to the next. In contrast, there is evidence that at least some mammals, including humans, do perceive octave equivalence.

Pitch chroma information is processed in addition to pitch height information with more than two pitch-range categories.

Octave equivalence describes the perception that notes separated by a doubling in frequency sound similar. While the octave is used cross-culturally as a basis of pitch perception, experimental demonstration of the phenomenon has proved to be difficult. In past work, members of our group developed a three-range generalization paradigm that reliably demonstrated octave equivalence. In this study we replicate and expand on this previous work trying to answer three questions that help us understand the origins and potential cross-cultural significance of octave equivalence: (1) whether training with three ranges is strictly necessary or whether an easier-to-learn two-range task would be sufficient, (2) whether the task could demonstrate octave equivalence beyond neighbouring octaves, and (3) whether language skills and musical education impact the use of octave equivalence in this task. We conducted a large-sample study using variations of the original paradigm to answer these questions. Results found here suggest that the three-range discrimination task is indeed vital to demonstrating octave equivalence. In a two-range task, pitch height appears to be dominant over octave equivalence. Octave equivalence has an effect only when pitch height alone is not sufficient. Results also suggest that effects of octave equivalence are strongest between neighbouring octaves, and that tonal language and musical training have a positive effect on learning of discriminations but not on perception of octave equivalence during testing. We discuss these results considering their relevance to future research and to ongoing debates about the basis of octave equivalence perception.

Advice to young behavioral and cognitive scientists.

Modeled on Medawar's Advice to a Young Scientist [Medawar, P.B., 1979. Advice to a Young Scientist. Basic Books, New York], this article provides advice to behavioral and cognitive scientists. An important guiding principle is that the study of comparative cognition and behavior are natural sciences tasked with explaining nature. The author advises young scientists to begin with a natural phenomenon and then bring it into the laboratory, rather than beginning in the laboratory and hoping for an application in nature. He suggests collaboration as a way to include research outside the scientist's normal competence. He then discusses several guides to good science. These guides include Tinbergen's [Tinbergen, N., 1963. On aims and methods of ethology. Zeitschrift für Tierpsychologie, 20, 410-433. This journal was renamed Ethology in 1986. Also reprinted in Anim. Biol. 55, 297-321, 2005] four "why" questions, Platt's [Platt, J.R., 1964. Strong inference. Science 146, 347-353, (http://weber.ucsd.edu/~jmoore/courses/Platt1964.pdf)] notion of strong inference using multiple alternative hypotheses, and the idea that positive controls help scientists to follow Popper's [Popper, K.R., 1959. The Logic of Scientific Discovery. Basic Books, New York, p. 41] advice about disproving hypotheses. The author also recommends Strunk and White's [Strunk, W., White, E.B., 1979. The Elements of Style, third ed. Macmillan, New York] rules for sound writing, and he provides his personal advice on how to use the anticipation of peer review to improve research and how to decode editors' and reviewers' comments about submitted articles.

Rats' (Rattus norvegicus) temporal discrimination of brief auditory stimuli: how does it compare with that of humans (Homo sapiens) and zebra finches (Taeniopygia guttata)?

The present study with rats replicated an experiment on the ability of zebra finches and humans to discriminate among brief auditory stimuli (see Weisman et al., 1999, Experiment 2). We trained rats with 27 3-kHz tones that varied in duration from 10 ms to 1420 ms. Reinforcement was contingent on responding (approaching the food well) to the nine medium-durations range tones (56-255 ms) but not to the nine short-durations range (10-46 ms) or long-durations range tones (309-1420 ms). Rats also received post-discrimination transfer tests with 2 kHz and 4 kHz tones that varied over the same durations as the 3 kHz tones. Rats acquired the temporal discrimination to a slightly lower level of accuracy than seen in finches or humans by Weisman et al. (1999). We tested for transfer of the temporal discrimination to find that rats, similar to humans (data from Weisman et al., 1999), transferred to untrained 2-kHz and 4-kHz tones at levels approaching accuracy to that achieved to the trained 3-kHz tone. By contrast, zebra finches (data from Weisman et al., 1999) failed to transfer their discrimination to the trained tone. We conclude that (a) rats discriminate among tone durations at least as well as they do among auditory frequencies and (b) rats like humans, but unlike finches, are insensitive to absolute pitch in their temporal discrimination.

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.

Rationale and methodology for testing auditory cognition in songbirds.

Songbirds, and in particular zebra finches, present a wonderful opportunity to study cognition in species that have evolved specialized abilities and brain structures for auditory cognition. The authors describe the rationale, methods, and apparatus used to test the auditory perceptual and cognitive abilities of songbirds. They have developed an operant conditioning system for conducting discrimination experiments simultaneously with several songbirds. The system uses specialized single-board computers, standard personal computers, CD-ROMs, and custom-written software to present stimuli, control training, and record responses. Also, the authors describe software to produce high-quality synthesized and naturally occurring acoustic stimuli for use in studies of auditory cognition. Typical results from a challenging frequency-range discrimination are included.

Effects of songs and calls on ZENK expression in the auditory telencephalon of field- and isolate-reared black capped chickadees.

We examined the effects of hearing two different conspecific vocalizations on expression of the immediate-early gene ZENK in the caudomedial neostriatum (NCM) and the caudomedial portion of the ventral hyperstriatum (cmHV) in male and female black-capped chickadees (Poecile atricapilla). Both the fee-bee song and the chick-a-dee call induced Zenk protein expression in NCM and in cmHV, however, patterns of expression to songs and calls varied across brain region. In the dorsal region of NCM, fee-bee songs induced more Zenk expression than chick-a-dee calls. In ventral NCM and cmHV, Zenk expression did not differ between songs and calls. We found that sex of the listener also affected Zenk expression: there was more robust ZENK response in males than in females. Finally, we compared field- and isolate-reared chickadees and found similar Zenk expression to fee-bee song in each group. These findings indicated that the type of conspecific vocalization, as well as the sex of the listener, appear to modulate IEG expression in the songbird ascending auditory pathway.

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.

A behavior analysis of absolute pitch: sex, experience, and species.

Absolute pitch (AP) perception refers to the ability to identify, classify, and memorize pitches without use of an external reference pitch. In tests of AP, several species were trained to sort contiguous tones into three or eight frequency ranges, based on correlations between responding to tones in each frequency range and reinforcement. Two songbird species, zebra finches and white-throated sparrows, and a parrot species, budgerigars had highly accurate AP, they discriminated both three and eight ranges with precision. Relative to normally reared songbirds, isolate reared songbirds had impaired AP. Two mammalian species, humans and rats, had equivalent and weak AP, they discriminated three frequency ranges to a lackluster standard and they acquired only a crude discrimination of the lowest and highest of eight frequency ranges. In comparisons with mammals even isolate songbirds had more accurate AP than humans and rats.

Pitch chroma discrimination, generalization, and transfer tests of octave equivalence in humans.

Octave equivalence occurs when notes separated by an octave (a doubling in frequency) are judged as being perceptually similar. Considerable evidence points to the importance of the octave in music and speech. Yet, experimental demonstration of octave equivalence has been problematic. Using go/no-go operant discrimination and generalization, we studied octave equivalence in humans. In Experiment 1, we found that a procedure that failed to show octave equivalence in European starlings also failed in humans. In Experiment 2, we modified the procedure to control for the effects of pitch height perception by training participants in Octave 4 and testing in Octave 5. We found that the pattern of responding developed by discrimination training in Octave 4 generalized to Octave 5. We replicated and extended our findings in Experiment 3 by adding a transfer phase: Participants were trained with either the same or a reversed pattern of rewards in Octave 5. Participants transferred easily to the same pattern of reward in Octave 5 but struggled to learn the reversed pattern. We provided minimal instruction, presented no ordered sequences of notes, and used only sine-wave tones, but participants nonetheless constructed pitch chroma information from randomly ordered sequences of notes. Training in music weakly hindered octave generalization but moderately facilitated both positive and negative transfer.

Reinforcer devaluation by extinction depends on the food restriction protocol.

A common feature of reinforcer devaluation studies is that new learning induces the devaluation. The present study used extinction to induce new learning about the conditioned reinforcer in a heterogeneous chain schedule. Rats pressed a lever in a heterogeneous chain schedule to produce a conditioned reinforcer (light) associated with the opportunity to obtain an unconditioned reinforcer (food) by pulling a chain. The density of food reinforcement correlated with the conditioned reinforcer was varied in a comparison of continuous and variable-ratio reinforcement schedules of chain pulling; this had no noticeable effect on conditioned reinforcer devaluation produced by extinction of chain pulling. In contrast, how rats were deprived appeared to matter very much. Restricting meal duration to 1h daily produced more lever pressing during baseline training and a greater reductive effect of devaluation on lever pressing than restricting body weight to 80% of a control rat's weight, which eliminated the devaluation effect. Further analysis suggested that meal-duration restriction may have produced devaluation effects because it was more effective than weight restriction in reducing rats' body weights. Our results exposed an important limitation on the devaluation of conditioned reinforcers: slight differences in food restriction, using two commonly employed food-restriction procedures, can produce completely different interpretations of reinforcer devaluation while leaving reinforcer-based learning intact.

Using network models of absolute pitch to compare frequency-range discriminations across avian species.

The spectral frequency ranges of song notes are important for recognition in avian species tested in the field. Frequency-range discriminations in both the field and laboratory require absolute pitch (AP). AP is the ability to perceive pitches without an external referent. The authors provided a network model designed to account for differences in AP among avian species and evaluated it against discriminative performance in eight-frequency-range laboratory tests of AP for five species of songbirds and two species of nonsongbirds. The model's sensory component describes the neural substrate of avian auditory perception, and its associative component handles learning of the discrimination. Using only two free parameters to describe the selectivity and the sensitivity of each species' auditory sensory filters, the model provided highly accurate predictions of frequency-range discrimination in songbirds and in a parrot species, but performance and its prediction were less accurate in pigeons: the only species tested that does not learn its vocalizations. Here for the first time, the authors present a model that predicted individual species' performance in frequency-range discriminations and predicted differences in discrimination among avian species with high accuracy.

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.