Left-side dominance for song discrimination in Bengalese finches (Lonchura striata var. domestica)

Male Bengalese finches are left-side dominant for the motor control of song in the sensorimotor nucleus (the high vocal center, or HVc) of the telencephalon. We examined whether perceptual discrimination of songs might also be lateralized in this species. Twelve male Bengalese finches were trained by operant conditioning to discriminate between a Bengalese finch song and a zebra finch song. Before training, the left HVc was lesioned in four birds and the right HVc was lesioned in four other birds. The remaining four birds were used as controls without surgery. Birds with a left HVc lesion required significantly more time to learn to discriminate between the two songs than did birds with a right HVc lesion or intact control birds. These results suggest that the left HVc is not only dominant for the motor control of song, but also for the perceptual discrimination of song. Accepted after revision: 11 September 2001 Electronic Publication     

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.     

Cross-fostering diminishes song discrimination in zebra finches (Taeniopygia guttata)

Song-production, -discrimination, and -preferences in oscine birds are dually influenced by species identity and the ontogenetic environment. The cross-fostering of a model species for recognition research, the zebra finch (Taeniopygia guttata) into heterospecific nests of the Bengalese finch (Lonchura striata vars. domestica) allows an exploration of the sensory limits of early development and the effects of species-specific acoustic cues upon song discrimination in adulthood. To quantify the song preferences of female and male normal-reared (control) and Bengalese finch fostered zebra finches, we recorded multiple behavioral measures, including spatial proximity, vocalization rates and response latency, during sequential song-playback choice-trials using both tutor species’ songs and the songs of two other ecologically relevant Australian species, the owl finch (Taeniopygia bichenovii) and the star finch (Neochmia ruficauda). Response strength was variable between the different measures, but no differences were detected within the specific behavioral responses towards the song playbacks of the two sexes. Control subjects strongly preferred their own species’ songs while Bengalese-fostered zebra finches exhibited reduced song discrimination between con-, tutor-, and heterospecific songs. Overall behavioral responsiveness was also modulated by social ontogeny. These results indicate a difference in the strength of preference for song that is dependent on the species identity of the rearing environment in oscine birds and illustrate the role of multiple behavioral measures and ecologically relevant stimulus species selection in behavioral research using zebra finches.     

Rule learning by zebra finches in an artificial grammar learning task: which rule?

A hallmark of the human language faculty is the use of syntactic rules. The natural vocalizations of animals are syntactically simple, but several studies indicate that animals can detect and discriminate more complex structures in acoustic stimuli. However, how they discriminate such structures is often not clear. Using an artificial grammar learning paradigm, zebra finches were tested in a Go/No-go experiment for their ability to distinguish structurally different three-element sound sequences. In Experiment 1, zebra finches learned to discriminate ABA and BAB from ABB, AAB, BBA, and ABB sequences. Tests with probe sounds consisting of four elements suggested that the discrimination was based on attending to the presence or absence of repeated A- and B-elements. One bird generalized the discrimination to a new element type. In Experiment 2, we continued the training by adding four-element songs following a ‘first and last identical versus different’ rule that could not be solved by attending to repetitions. Only two out of five birds learned the overall discrimination. Testing with novel probes demonstrated that discrimination was not based on using the ‘first and last identical’ rule, but on attending to the presence or absence of the individual training stimuli. The two birds differed in the strategies used. Our results thus demonstrate only a limited degree of abstract rule learning but highlight the need for extensive and critical probe testing to examine the rules that animals (and humans) use to solve artificial grammar learning tasks. They also underline that rule learning strategies may differ between individuals.     

Perception of temporal properties in self-generated songs by Bengalese finches (Lonchura striata var. domestica)

In the songbird forebrain, neuronal selectivity for temporal properties of each bird's self-generated song has been well described, but the behavioral and perceptual correlates of this selectivity are not known. By operant procedures, the authors trained Bengalese finches (Lonchura striata var. domestica) to discriminate between songs that were played normally and in reverse. Male Bengalese finches learned the discrimination quicker when their self-generated song was used as a stimulus than when a song of another conspecific bird was used. When the global note order was retained but each note was locally reversed, the song was more likely to be regarded as a forward song by the singer himself, but not by other birds. These results provide psychophysical evidence that the special processing of the self-generated song observed at the neural level might reflect an individual's perception of his self-produced song.     

Event-related potentials reveal the relations between feature representations at different levels of abstraction

In this paper, we use behavioural methods and event-related potentials (ERPs) to explore the relations between informational and instantiated features, as well as the relation between feature abstraction and rule type. Participants are trained to categorize two species of fictitious animals and then identify perceptually novel exemplars. Critically, two groups are given a perfectly predictive counting rule that, according to Hannah and Brooks (2009. Featuring familiarity: How a familiar feature instantiation influences categorization. Canadian Journal of Experimental Psychology/Revue Canadienne de Psychologie Expérimentale, 63, 263–275. Retrieved from http://doi.org/10.1037/a0017919), should orient them to using abstract informational features when categorizing the novel transfer items. A third group is taught a feature list rule, which should orient them to using detailed instantiated features. One counting-rule group were taught their rule before any exposure to the actual stimuli, and the other immediately after training, having learned the instantiations first. The feature-list group were also taught their rule after training. The ERP results suggest that at test, the two counting-rule groups processed items differently, despite their identical rule. This not only supports the distinction that informational and instantiated features are qualitatively different feature representations, but also implies that rules can readily operate over concrete inputs, in contradiction to traditional approaches that assume that rules necessarily act on abstract inputs.     

Perceptual chunking in the self-produced songs of Bengalese finches (Lonchura striata var. domestica)

Like humans, songbirds, including Bengalese finches, have hierarchical structures in their vocalizations. When humans perceive a sentence, processing occurs in phrase units, not words. In this study, we investigated whether songbirds also perceive their songs by chunks (clusters of song notes) rather than single song notes. We trained male Bengalese finches to react to a short noise in a Go/NoGo task. We then superimposed the noise onto recordings of their own songs and examined whether the reaction time was affected by the location of the short noise, that is, whether the noise was placed between chunks or in the middle of a chunk. The subjects’ reaction times to the noise in the middle of a chunk were significantly longer than those to the noise placed between chunks. This result was not observed, however, when the songs were played in reverse. We thus concluded that Bengalese finches perceive their songs by chunks rather than single notes.     

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     

Factors influencing song development in the white-crowned sparrow (Zonotrichia leucophrys)

Nestling white-crowned sparrows (Zonotrichia leucophrys nuttalli) were hand-reared in sound-isolation chambers under a variety of conditions. The songs of total isolates were compared with songs of birds tutored with song, and the number of inputs sufficient for a bird to produce a normal song was explored. The flexibility of the song learning system was investigated with a range of tape-recorded tutor songs: alien dialects, alien subspecies, alien species, alternating alien dialects, and an aberrant song. Adult songs were obtained for 40 males and 7 testosterone-injected females. All of the tutor songs could be learned. Also, some birds learned elements of an alien species' song. Birds tutored with two songs copied one or the other, were bilingual, or sang a hybrid of the two. No bird presented with fewer than 120 songs learned the tutor song; 2 birds tutored with 252 songs copied the tutor song. It is concluded that the song learning system is quite flexible, hat the results obtained with tape-tutors are very different from those with social tutors, and that there may be an interaction between total number of song inputs and the number presented on a single day. Some implications of these data for physiological mechanisms and the possible functional significance of the acquisition system are discussed.     

内隐序列学习表征方式的实验研究

采用Lewicki等人创造的“矩阵扫描”任务,对内隐序列学习的表征方式进行了研究。结果发现：规则试验的反应时显著低于随机试验,说明被试能够内隐地掌握复杂规则,没有证据表明这与“运动”频率的大小和序列样例的记忆有关;此外,序列结构也是影响内隐序列学习的因素之一。     

Auditory temporal pattern learning by songbirds using maximal stimulus diversity and minimal repetition

The sequential patterning of complex acoustic elements is a salient feature of bird song and other forms of vocal communication. For European starlings (Sturnus vulgaris), a songbird species, individual vocal recognition is improved when the temporal organization of song components (called motifs) follows the normal patterns of each singer. This sensitivity to natural motif sequences may underlie observations that starlings can also learn more complex, unnatural motif patterns. Alternatively, it has been proposed that the apparent acquisition of abstract motif patterning rules instead reflects idiosyncrasies of the training conditions used in prior experiments. That is, that motif patterns are learned not by recognizing differences in temporal structures between patterns, but by identifying serendipitous features (e.g., acoustical cues) in the small sets of training and testing stimuli used. Here, we investigate this possibility, by asking whether starlings can learn to discriminate between two arbitrary motif patterns, when unique examples of each pattern are presented on every trial. Our results demonstrate that abstract motif patterning rules can be acquired from trial-unique stimuli and suggest that such training leads to better pattern generalization compared with training with much smaller stimulus subsets.     

Direct social contacts override auditory information in the song-learning process in starlings (Sturnus vulgaris)

Social influence on song acquisition was studied in 3 groups of young European starlings raised under different social conditions but with the same auditory experience of adult song. Attentional focusing on preferred partners appears the most likely explanation for differences found in song acquisition in relation to experience, sex, and song categories. Thus, pair-isolated birds learned from each other and not from broadcast live songs, females did not learn from the adult male tutors, and sharing occurred more between socially associated peers. On the contrary, single-isolated birds clearly copied the adult songs that may have been the only source of attention stimulation. Therefore, social preference appears as both a motor for song learning and a potential obstacle for acquisition from nonpreferred partners, including adults.     

Vocal syntax development in the white-crowned sparrow (Zonotrichia leucophrys).

Birdsong development exemplifies the interplay between experience and predisposition that occurs during behavioral ontogeny. Songbirds must hear song models to develop normal song, yet they preferentially learn conspecific song when given a choice in the laboratory. To the extent that features guiding this selective learning are pre-encoded in the brain, such features should also develop in the song of young birds not exposed to them in tutor models. To investigate whether song syntax-phrase number and order--is such a feature in the white-crowned sparrow (Zonotrichia leucophrys), the authors tutored males of this species with separate phrase models. Birds learned and assembled these into songs of species-typical sequence, suggesting that syntax is to some degree pre-encoded in white-crowned sparrows. Birds also learned heterospecific phrases, confirming previous evidence that note phonology is not the primary cue for selective song learning in this species.     

The effects of cumulative practice on mathematics problem solving

This study compared three different methods of teaching five basic algebra rules to college students. All methods used the same procedures to teach the rules and included four 50-question review sessions interspersed among the training of the individual rules. The differences among methods involved the kinds of practice provided during the four review sessions. Participants who received cumulative practice answered 50 questions covering a mix of the rules learned prior to each review session. Participants who received a simple review answered 50 questions on one previously trained rule. Participants who received extra practice answered 50 extra questions on the rule they had just learned. Tests administered after each review included new questions for applying each rule (application items) and problems that required novel combinations of the rules (problem-solving items). On the final test, the cumulative group outscored the other groups on application and problem-solving items. In addition, the cumulative group solved the problem-solving items significantly faster than the other groups. These results suggest that cumulative practice of component skills is an effective method of training problem solving.     

Experience with songs in adulthood reduces song-induced gene expression in songbird auditory forebrain

Male songbirds learn to produce song within a limited phase early in life; however they continue to learn to recognize songs in adulthood. Studies looking at Zenk activation after exposure to songs learned early in life for song production and songs learned in adulthood show opposite patterns of activation, suggesting distinct neural mechanisms may be involved in these two forms of learning. In this study, we look at IEG Zenk activation in auditory regions NCM and CMM of song sparrows (Melospiza melodia) to see whether recent exposure to song in adulthood leads to greater or decreased Zenk activation upon hearing that song versus a novel song. We found significantly lower activation in birds exposed to previously heard songs versus novel songs in vNCM but not dNCM, though further analysis suggest an overall trend in NCM. We found no significant difference in the amount of activation to previously heard songs vs. novel songs in CMM. These results support previous findings suggesting that activation is reduced to learned stimuli; we discuss possible implications of these findings in relation to song production learning early in life and song recognition learning in adulthood.     

Artificial grammar learning in children: abstraction of rules or sensitivity to perceptual features?

We examined sensitivity to grammatical sequences of colors in an artificial grammar learning task in a sample of 120 children aged between 5 and 8 years. The aim of the experiment was to test whether the children would preferentially learn the specific salient features of the items they were exposed to or the rules that generated these items. The children were divided into two experimental groups (identical grammar but training items differing in their surface features) and a control group (random items). The results showed that regardless of age, participants learned the most frequent salient features of the items, as well as some kind of abstract relational information. However, the 8-year-olds presented a more complex result profile, with one of the experimental groups apparently developing sensitivity to grammatical rules. These results are discussed with reference to the main current models of implicit learning. Overall, the results provided more support for stimulus-specific processing models than for rule-based models.     

Failure of hierarchical conditional rule learning in the pigeon (Columba livia)

Pigeons were trained with a conditional discrimination task in three-key operant chambers. Choosing either the left or right key was followed by food according to combinations of three preceding events: (a) a houselight illumination condition (dark or light), (b) presence or absence of green flashes on the three keys, (c) a color (amber or blue) of the center sample key. With these 2×2×2 event combinations, eight types of correct trials were prepared: (1) dark→no flash→amber→LEFT, (2) dark→no flash→blue→RIGHT, (3) dark→flash→amber→RIGHT, (4) dark→flash→blue→LEFT, (5) light→no flash→amber→RIGHT, (6) light→no flash→blue→LEFT, (7) light→flash→amber→LEFT, and (8) light→flash→blue→RIGHT. Seven of these eight types were used for training of a given bird, and then the remaining trial type was presented as the test. If the birds had learned the conditional structure of the events (the hierarchical switching rule), they would have responded correctly to the test type. However, they chose the opposite side key, suggesting that they had learned cue configuration or multiple rules to solve the task. Accepted after revision: 13 January 2001 Electronic Publication     

Matching and oddity relational learning by pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>): transfer from color to shape

Relational learning, as opposed to perceptual learning, is based on the abstract properties of the stimuli. Although at present there is no doubt that pigeons are capable of relational behavior, this study aims to further disclose the conditions under which it occurs. Pigeons were trained in an outdoor cage on a matching-to-sample or an oddity-from-sample task, with colored cardboard stimuli presented horizontally. The apparatus involved three sliding lids on which the stimuli were drawn and which, when displaced, revealed the reinforcement. The lids were either adjacent to each other or somewhat separated. Training sessions involved two colors, and test sessions six different colors (same dimension test), or six different shapes (different dimension test). One group of birds trained under the ‘adjacent’ condition failed when tested with new stimuli, but succeeded in both dimension tests after training under the ‘separate’ condition. Two other groups of birds succeeded in all tests after training under the latter condition. These results show that depending on procedural details, pigeons are or are not able to transfer from one visual dimension to another, thus extending previous related findings.     

Processing abstract sequence structure: learning without knowing, or knowing without learning?

Constant interaction with a dynamic environment—from riding a bicycle to segmenting speech—makes sensitivity to the sequential structure of the world a fundamental dimension of information processing. Accounts of sequence learning vary widely, with some authors arguing that parsing and segmentation processes are central, and others proposing that sequence learning involves mere memorization. In this paper, we argue that sequence knowledge is essentially statistical in nature, and that sequence learning involves simple associative prediction mechanisms. We focus on a choice reaction situation introduced by Lee (1997), in which participants were exposed to material that follows a single abstract rule, namely that stimuli are selected randomly, but never appear more than once in a legal sequence. Perhaps surprisingly, people can learn this rule very well. Or can they? We offer a conceptual replication of the original finding, but a very different interpretation of the results, as well as simulation work that makes it clear how highly abstract dimensions of the stimulus material can in fact be learned based on elementary associative mechanisms. We conclude that, when relevant, memory is optimized to facilitate responding to events that have not occurred recently, and that sequence learning in general always involves sensitivity to repetition distance.     

Colour cues or spatial cues? Context-dependent preferences in the European greenfinch (<Emphasis Type="Italic">Carduelis chloris</Emphasis>)

Using featural cues such as colour to identify ephemeral food can increase foraging efficiency. Featural cues may change over time however; therefore, animals should use spatial cues to relocate food that occurs in a temporally stable position. We tested this hypothesis by measuring the cue preferences of captive greenfinches Carduelis chloris when relocating food hidden in a foraging tray. In these standardised associative learning trials, greenfinches favoured colour cues when returning to a foraging context that they had encountered before only once (“one-trial test”) but switched to spatial cues when they had encountered that scenario on ten previous occasions (“repeated-trial test”). We suggest that repeated encounters generated a context in which individuals had a prior expectation of temporal stability, and hence context-dependent cue selection. Next, we trained birds to find food in the absence of colour cues but tested them in the presence of visual distracters. Birds were able to learn spatial cues after one encounter, but only when visual distracters were identical in colouration. When a colourful distracter was present in the test phase, cue selection was random. Unlike the first one-trial test, birds were not biased towards this colourful visual distracter. Together, these results suggest that greenfinches are able to learn both cue types, colour cue biases represent learning, not simply distraction, and spatial cues are favoured over colour cues only in temporally stable contexts.