Rapidly learned song-discrimination without behavioral reinforcement in adult male zebra finches (Taeniopygia guttata)

Zebra finches communicate via several distinct vocalizations, of which song is the most studied. Behavioral observations indicate that adults are able to discriminate among the songs of different conspecific individuals. In the wild, zebra finches live in structured but mobile colonies, and encounter new individuals on a frequent basis. Thus it seems plausible that adult finches might have the capacity to recognize and remember new songs they encounter on a single day, but this has never been directly tested. Here we devised a simple observational assay to determine whether adult male zebra finches show recognition of a song they have heard repeatedly from taped playbacks, over a single three hour period the day before. We quantified the rate of production of six discrete behaviors (short calls, contact calls, singing, short hops, long hops, and beak swipes) made by adult male zebra finches as they listened to the playbacks. At the onset of song playback, all birds suspended these behaviors and sat silently-occasionally moving their heads. Then, after a measurable period ("response latency"), the birds resumed these activities. We observed that the response latency was long (approximately 10 min) when birds were hearing a particular song for the first time. The response latency was much shorter (approximately 1-2 min) when the birds had heard the same song the day before. Thus, functional song memories must result from as little as 3 h of passive song-exposure. These results suggest that ongoing song learning may play a natural role in the daily life of adult zebra finches, and provide a behavioral reference point for studies of molecular and physiological plasticity in the adult auditory system.     

Zebra finches (Taeniopygia guttata) demonstrate cognitive flexibility in using phonology and sequence of syllables in auditory discrimination

Animal Cognition - Zebra finches rely mainly on syllable phonology rather than on syllable sequence when they discriminate between two songs. However, they can also learn to discriminate two...     

Auditory scene analysis in estrildid finches (Taeniopygia guttata and Lonchura striata domestica): a species advantage for detection of conspecific song

Operant-conditioning techniques were used to investigate the ability of zebra finches (Taeniopygia guttata) and Bengalese finches (Lonchura striata domestica) to detect a zebra finch or a Bengalese finch target song intermixed with other birdsongs. Sixteen birds were trained to respond to the presence of a particular target song, either of their own species (n = 8) or of another species (n = 8). The birds were able to learn a discrimination between song mixtures that contained a target song and song mixtures that did not, and they were able to maintain their response to the target song when it was mixed with novel songs. Zebra finches, but not Bengalese finches, learned the discrimination with a conspecific target more quickly and were worse at detecting a Bengalese finch in the presence of a conspecific song. The results indicate that selective attention to birdsongs within an auditory scene is related to their biological relevance.     

Breeding cycle aggression in domesticated zebra finches (Poephila guttata)

Patterns of breeding cycle aggressive behavior were examined in colonies of domesticated zebra finches (Poephila guttata). Aggressive behaviors of zebra finch pairs were low during non-nesting periods but were maintained at high levels during the three nesting periods: the incubating, hatching, and nestling periods. Defense of the nest site was the prominent functional category of aggression, followed by defense of individual distance. Defense of mate, defense of young, and defense of nesting material rarely occurred. Male and female members of pair bonds shared the duties of defense of the nest site and parental care. During observation times males exhibited greater frequencies of aggressive actions throughout the breeding cycle and females spent a greater amount of time in the nest during the three nesting periods. The pattern and functional categories of aggressive behavior in the zebra finch colonies correspond most closely to the nest site defense hypothesis for the evolution of breeding season aggression.     

Relative salience of syllable structure and syllable order in zebra finch song

There is a rich history of behavioral and neurobiological research focused on the ‘syntax’ of birdsong as a model for human language and complex auditory perception. Zebra finches are one of the most widely studied songbird species in this area of investigation. As they produce song syllables in a fixed sequence, it is reasonable to assume that adult zebra finches are also sensitive to the order of syllables within their song; however, results from electrophysiological and behavioral studies provide somewhat mixed evidence on exactly how sensitive zebra finches are to syllable order as compared, say, to syllable structure. Here, we investigate how well adult zebra finches can discriminate changes in syllable order relative to changes in syllable structure in their natural song motifs. In addition, we identify a possible role for experience in enhancing sensitivity to syllable order. We found that both male and female adult zebra finches are surprisingly poor at discriminating changes to the order of syllables within their species-specific song motifs, but are extraordinarily good at discriminating changes to syllable structure (i.e., reversals) in specific syllables. Direct experience or familiarity with a song, either using the bird’s own song (BOS) or the song of a flock mate as the test stimulus, improved both male and female zebra finches’ sensitivity to syllable order. However, even with experience, birds remained much more sensitive to structural changes in syllables. These results help to clarify some of the ambiguities from the literature on the discriminability of changes in syllable order in zebra finches, provide potential insight on the ethological significance of zebra finch song features, and suggest new avenues of investigation in using zebra finches as animal models for sequential sound processing.     

Song recognition in zebra finches: Are there sensitive periods for song memorization?

Male zebra finches learn to sing songs that they hear between 25 and 65 days of age, the sensitive period for song learning. In this experiment, male and female zebra finches were exposed to zebra finch songs either before (n = 9) or during (n = 4) the sensitive period. Following song exposure, recognition memory for the songs was assessed with an operant discrimination between familiar and novel songs. Zebra finches that were exposed to songs between 22 and 30 days of age discriminated between familiar and novel songs; zebra finches exposed to songs from 9 to 17 days of age did not. Failure to memorize songs heard prior to the sensitive period may contribute to the exclusion of those songs from the repertoire of songbirds.     

Selective auditory grouping by zebra finches: testing the iambic–trochaic law

Humans have a strong tendency to spontaneously group visual or auditory stimuli together in larger patterns. One of these perceptual grouping biases is formulated as the iambic/trochaic law, where humans group successive tones alternating in pitch and intensity as trochees (high–low and loud–soft) and alternating in duration as iambs (short–long). The grouping of alternations in pitch and intensity into trochees is a human universal and is also present in one non-human animal species, rats. The perceptual grouping of sounds alternating in duration seems to be affected by native language in humans and has so far not been found among animals. In the current study, we explore to which extent these perceptual biases are present in a songbird, the zebra finch. Zebra finches were trained to discriminate between short strings of pure tones organized as iambs and as trochees. One group received tones that alternated in pitch, a second group heard tones alternating in duration, and for a third group, tones alternated in intensity. Those zebra finches that showed sustained correct discrimination were next tested with longer, ambiguous strings of alternating sounds. The zebra finches in the pitch condition categorized ambiguous strings of alternating tones as trochees, similar to humans. However, most of the zebra finches in the duration and intensity condition did not learn to discriminate between training stimuli organized as iambs and trochees. This study shows that the perceptual bias to group tones alternating in pitch as trochees is not specific to humans and rats, but may be more widespread among animals.     

Significance of visual cues in choice behavior in the female zebra finch (<Emphasis Type="Italic">Taeniopygia guttata</Emphasis><Emphasis Type="Italic">castanotis</Emphasis>)

Female zebra finches show a preference for male zebra finches over heterospecific males based solely on the auditory cues of males, such as songs. The present study was designed to investigate whether females show a similar preference for male zebra finches based solely on visual cues. Using a Y-maze apparatus, social preference of female zebra finches was studied between male zebra finches and male Bengalese finches in three experiments. In experiment 1, where female zebra finches could see and hear live male zebra finches and male Bengalese finches, the females preferred to associate with the male zebra finches. In experiment 2, using a sound-attenuated experimental apparatus, subjects could see, but not hear, male zebra finches and male Bengalese finches. The subjects did not show a significant preference for associating with zebra finches. In experiment 3, as in experiment 2, females could see live male zebra finches and male Bengalese finches in the sound-attenuated chambers. However, in experiment 3, the subjects also heard prerecorded auditory cues (i.e., songs and calls) of male zebra finches, which were presented simultaneously in both arms of the maze. Although the females could not use the auditory cues to identify the location of the male zebra finches, they preferred to associate with the male zebra finches rather than the male Bengalese finches. These results suggest that visual cues alone were effective in initiating choice behaviors by females and that auditory cues facilitate such visually based choice behaviors. Electronic Publication     

Experimental determination of a unit of song production in the zebra finch (Taeniopygia guttata)

Zebra finch (Taeniopygia guttata) song is composed of syllables delivered in a set order. Little is known about the program that controls this temporal delivery. A decision to sing or not to sing may or may not affect the entire song. Song, once commenced, may continue or may halt. If song is halted, stops may occur only at certain points. Seven zebra finches were presented with short bursts of strobe light while engaged in song. The variables of interest were whether the birds stopped and where they stopped. The results can be summarized as follows: Ongoing zebra finch song can be interrupted, interruptions occur at discrete locations in song, and the locations almost always fall between song syllables. These results reveal a functional representation of song production and place constraints on possible neural mechanisms that underlie song production in zebra finches and probably other oscine species. The results also raise hypotheses about the elements of song perception and memory.     

Timbre discrimination in zebra finch (Taeniopygia guttata) song syllables

Zebra finch (Taeniopygia guttata) songs include syllables of a fundamental frequency and harmonics. Individual harmonics in 1 syllable can be more or less emphasized. The functional role of this variability is unknown. These experiments provide evidence of how the phenomenon is perceived. We trained 12 male and female zebra finches on a go-no-go operant procedure to discriminate between 2 song syllables that varied only in the absence of the 2nd or 5th harmonic. Training involved many thousands of trials. Both sexes used the presence or absence of the 2nd harmonic as the sole discriminative cue. Females had more difficulty learning to perform the task when the presence of the 2nd harmonic was the go stimulus, which indicates that their use of the information was biased by stimulus-response contingencies. The results are discussed in terms of a broad strategy to understand how animals perceive sounds used in communication.     

Motivation and memory in zebra finch (<Emphasis Type="Italic">Taeniopygia guttata</Emphasis>) foraging behavior

Remembering combinations of information such as what resources have been seen in which locations could play an important role in enhancing individual survival through increased foraging success. To date, there have been relatively few investigations of avian memory involving more than one category of information. This study explored zebra finches’ (Taeniopygia guttata) capacity to recall two categories in combination, namely food-type and spatial location. Birds were trained to remove variously weighted flaps to find two types of food hidden beneath. Memory for food-types and locations was assessed by pre-feeding the birds to satiety on and devaluing one food-type, and then testing the birds’ efficiency at finding the non-devalued food. When allowed one trial to learn locations of two food rewards that were hidden beneath lightly weighted flaps, birds performed better than chance at locating a food reward. However, they did not preferentially search for the non pre-fed food, suggesting that they were unable to recall both food type and location in combination. Zebra finches made fewer errors when tested on the one-trial task using more heavily weighted flaps than with lightly weighted flaps; there was equivocal evidence that they remembered which food type was hidden where on this task. When given repeated exposures to the locations of the two food rewards, finches located a food reward more accurately than on the one-trial tasks, and were also more likely to recall the locations of the different food types. In this foraging paradigm, experience and motivation may have influenced the birds’ performance.     

Open-field nidus and pullusindices in Mus Musculus L.

Swiss albino mice ( Mus Musculus L. ) were studied in an open field (OF). The question whether laboratory rodents were more attracted to a nest equipped with familiar odoriferous cues from the home-cage bedding or to signals of an unfamiliar newborn pup inserted in the OF was addressed. The results of the three quantitatively recorded OF behaviors, ambulation, retrieval and radial latency , indicated that in females both ambulation and retrieval were enhanced by the presence of a pup in a nestless field; in males locomotion was suppressed by a nest in a pupless OF. In the case of both sexes radial latency was determined by the OF position of a nest. A comparison between the two qualitative assessments, the nidus and pullus indices, yielded that the sex of the adult and the OF positions of both targets equally impacted attraction towards both nest and pup.     

Dominance interactions in young adult paper wasp (Polistes dominulus) foundresses: a playlike behavior?

The idea that insects play has often aroused skepticism. Nevertheless, the authors investigated the occurrence of a playlike behavior in young individuals of a paper wasp. Polistes dominulus foundresses hibernate in aggregations and found associative nests in the spring. In aggregations, wasps exhibit most of the rank-dependent behaviors that they will use in the nest 6 months later. In aggregations, precocious dominance interactions are performed with exaggerated frequencies, whereas aggression and trophallaxis are rare. Dominance behavior in aggregations is apparently useless, but it probably allows wasps to assess their dominance potential without any apparent reproductive competition. Surprisingly, these interactions may be best interpreted as play because dominance behavior in aggregations shares several features with mammalian play fighting.     

Defensive aggression in terns: Discrimination and response to individual researchers

The defensive responses of roseate terns (Sterna dougallii) were studied during the middle of the breeding season when all reproductive stages (incubation, chick-hatching, chick-rearing) were represented, to determine the factors affecting levels of nest defense and whether terns responded differently towards a familiar versus an unfamiliar researcher. Defensive aggression generally varied as a function of researcher identity, time of day, section or location in the colony and nest density. These variables explained from 60 to 66% of the variation in defensive aggression. More terns mobbed one researcher with more dive attacks than the other researcher. Terns were more aggressive in areas where the chicks had just hatched and were most vulnerable. © 1993 Wiley-Liss, Inc.     

Hippocampal activation of immediate early genes Zenk and c-Fos in zebra finches (Taeniopygia guttata) during learning and recall of a spatial memory task

Zebra finches (Taeniopygia guttata) are able to learn the position of food by orienting on spatial cues in a ‘dry water maze’. In the course of spatial learning, the hippocampus shows high expression of the immediate early genes (IEGs) Zenk and c-Fos, indicating high activation of this area during learning. In contrast, the IEG activity is nearly absent if the birds do not have to rely on spatial cues. In the present experiment it was investigated whether hippocampal activation can also be observed if the learned spatial task is recalled. For this purpose, the hippocampal Zenk and c-Fos activation of birds in an early learning stage was compared with that of others having well reached their maximal performance. The results show that the avian hippocampus is also active during recall of a learned spatial task, but the activation is significantly lower than in animals learning actually. As in previous experiments, hippocampal IEG expression showed strong variation not only in the position of the active patches of neurons, but also in size and cell density. The observed difference contributes to the view that immediate early genes may not be indicators of activation alone, but may be due to a combination of activation and plastic changes.     

The striped mouse (Rhabdomys pumilio) from the succulent karoo, South Africa: a territorial group-living solitary forager with communal breeding and helpers at the nest

The authors studied the striped mouse (Rhabdomys pumilio) in the semiarid succulent karoo of South Africa. Mice forage alone, but they live in groups that share a common nest. Groups consist of 1 to 4 breeding females, 1 to 2 breeding males, and their offspring of both sexes, which remain in their natal group even after reaching adulthood, participating in territorial defense and nest building without showing signs of reproductive activity. Interactions are typically amicable and take place inside or in front of the nest. In contrast, encounters with mice from other groups are aggressive. Group living in the succulent karoo is possibly due to ecological constraints imposed by habitat saturation because of a year-round stable food supply as well as associated benefits of philopatry.     

Effects of food and mates on time budget in Aplysia fasciata: integration of feeding, reproduction, and locomotion

This study examines the time budgeted to different behaviors in Aplysia fasciata under three conditions: (1) animals have constant access to food and mates: (2) there is access to food, but not to mates; (3) neither food nor mates are present. The data suggest a number of rules underlying behavioral integration: (1) Feeding, reproductive behaviors, and activity seem to be natural categories for behavioral choice. Feeding and reproductive behaviors are controlled in tandem by a common arousal mechanism, while time left over after animals feed and reproduce is distributed in a fixed ratio between locomotion (crawling and swimming) and inactivity (immobility and movement in place). (2) Relative distribution between different forms of locomotion and inactivity is modified by changes in motivational state. More time is spent swimming than crawling when feeding and/or mating is prevented, while more time is spent moving in place than immobile when the arousal level is increased. (3) Feedback control of feeding and reproduction is asymmetric. Satiation of feeding inhibits the common arousal. In the absence of food, time spent on reproductive behaviors increases due to disinhibition of the common arousal. By contrast, positive feedback arising from sexual behavior excites the common arousal. When mating is prevented by removing potential mates, time spent feeding decreases. (4) Generally, animals choose between performing the three main categories of behavior. Although Aplysia simultaneously can feed and mate, or locomote and mate, they do so infrequently. By contrast, different types of reproductive behaviors (male mating, female mating, egg-laying) are commonly performed simultaneously.     

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.     

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.