Non-adjacent Dependency Learning in Humans and Other Animals.

Learning and processing natural language requires the ability to track syntactic relationships between words and phrases in a sentence, which are often separated by intervening material. These nonadjacent dependencies can be studied using artificial grammar learning paradigms and structured sequence processing tasks. These approaches have been used to demonstrate that human adults, infants and some nonhuman animals are able to detect and learn dependencies between nonadjacent elements within a sequence. However, learning nonadjacent dependencies appears to be more cognitively demanding than detecting dependencies between adjacent elements, and only occurs in certain circumstances. In this review, we discuss different types of nonadjacent dependencies in language and in artificial grammar learning experiments, and how these differences might impact learning. We summarize different types of perceptual cues that facilitate learning, by highlighting the relationship between dependent elements bringing them closer together either physically, attentionally, or perceptually. Finally, we review artificial grammar learning experiments in human adults, infants, and nonhuman animals, and discuss how similarities and differences observed across these groups can provide insights into how language is learned across development and how these language-related abilities might have evolved.

Temporal modulation in speech, music, and animal vocal communication: evidence of conserved function.

Speech is a distinctive feature of our species. It is the default channel for language and constitutes our primary mode of social communication. Determining the evolutionary origins of speech is a challenging prospect, in large part because it appears to be unique in the animal kingdom. However, direct comparisons between speech and other forms of acoustic communication, both in humans (music) and animals (vocalization), suggest that important components of speech are shared across domains and species. In this review, we focus on a single aspect of speech-temporal patterning-examining similarities and differences across speech, music, and animal vocalization. Additional structure is provided by focusing on three specific functions of temporal patterning across domains: (1) emotional expression, (2) social interaction, and (3) unit identification. We hypothesize an evolutionary trajectory wherein the ability to identify units within a continuous stream of vocal sounds derives from social vocal interaction, which, in turn, derives from vocal emotional communication. This hypothesis implies that unit identification has parallels in music and precursors in animal vocal communication. Accordingly, we demonstrate the potential of comparisons between fundamental domains of biological acoustic communication to provide insight into the evolution of language.

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.

Spontaneous generalization of abstract multimodal patterns in young domestic chicks.

From the early stages of life, learning the regularities associated with specific objects is crucial for making sense of experiences. Through filial imprinting, young precocial birds quickly learn the features of their social partners by mere exposure. It is not clear though to what extent chicks can extract abstract patterns of the visual and acoustic stimuli present in the imprinting object, and how they combine them. To investigate this issue, we exposed chicks (Gallus gallus) to three days of visual and acoustic imprinting, using either patterns with two identical items or patterns with two different items, presented visually, acoustically or in both modalities. Next, chicks were given a choice between the familiar and the unfamiliar pattern, present in either the multimodal, visual or acoustic modality. The responses to the novel stimuli were affected by their imprinting experience, and the effect was stronger for chicks imprinted with multimodal patterns than for the other groups. Interestingly, males and females adopted a different strategy, with males more attracted by unfamiliar patterns and females more attracted by familiar patterns. Our data show that chicks can generalize abstract patterns by mere exposure through filial imprinting and that multimodal stimulation is more effective than unimodal stimulation for pattern learning.

Can Birds Perceive Rhythmic Patterns? A Review and Experiments on a Songbird and a Parrot Species.

While humans can easily entrain their behavior with the beat in music, this ability is rare among animals. Yet, comparative studies in non-human species are needed if we want to understand how and why this ability evolved. Entrainment requires two abilities: (1) recognizing the regularity in the auditory stimulus and (2) the ability to adjust the own motor output to the perceived pattern. It has been suggested that beat perception and entrainment are linked to the ability for vocal learning. The presence of some bird species showing beat induction, and also the existence of vocal learning as well as vocal non-learning bird taxa, make them relevant models for comparative research on rhythm perception and its link to vocal learning. Also, some bird vocalizations show strong regularity in rhythmic structure, suggesting that birds might perceive rhythmic structures. In this paper we review the available experimental evidence for the perception of regularity and rhythms by birds, like the ability to distinguish regular from irregular stimuli over tempo transformations and report data from new experiments. While some species show a limited ability to detect regularity, most evidence suggests that birds attend primarily to absolute and not relative timing of patterns and to local features of stimuli. We conclude that, apart from some large parrot species, there is limited evidence for beat and regularity perception among birds and that the link to vocal learning is unclear. We next report the new experiments in which zebra finches and budgerigars (both vocal learners) were first trained to distinguish a regular from an irregular pattern of beats and then tested on various tempo transformations of these stimuli. The results showed that both species reduced the discrimination after tempo transformations. This suggests that, as was found in earlier studies, they attended mainly to local temporal features of the stimuli, and not to their overall regularity. However, some individuals of both species showed an additional sensitivity to the more global pattern if some local features were left unchanged. Altogether our study indicates both between and within species variation, in which birds attend to a mixture of local and to global rhythmic features.

Budgerigars and zebra finches differ in how they generalize in an artificial grammar learning experiment.

The ability to abstract a regularity that underlies strings of sounds is a core mechanism of the language faculty but might not be specific to language learning or even to humans. It is unclear whether and to what extent nonhuman animals possess the ability to abstract regularities defining the relation among arbitrary auditory items in a string and to generalize this abstraction to strings of acoustically novel items. In this study we tested these abilities in a songbird (zebra finch) and a parrot species (budgerigar). Subjects were trained in a go/no-go design to discriminate between two sets of sound strings arranged in an XYX or an XXY structure. After this discrimination was acquired, each subject was tested with test strings that were structurally identical to the training strings but consisted of either new combinations of known elements or of novel elements belonging to other element categories. Both species learned to discriminate between the two stimulus sets. However, their responses to the test strings were strikingly different. Zebra finches categorized test stimuli with previously heard elements by the ordinal position that these elements occupied in the training strings, independent of string structure. In contrast, the budgerigars categorized both novel combinations of familiar elements as well as strings consisting of novel element types by their underlying structure. They thus abstracted the relation among items in the XYX and XXY structures, an ability similar to that shown by human infants and indicating a level of abstraction comparable to analogical reasoning.

Pauses enhance chunk recognition in song element strings by zebra finches.

When learning a language, it is crucial to know which syllables of a continuous sound string belong together as words. Human infants achieve this by attending to pauses between words or to the co-occurrence of syllables. It is not only humans that can segment a continuous string. Songbirds learning their song tend to copy 'chunks' from one or more tutors' songs and combine these into their own song. In the tutor songs, these chunks are often separated by pauses and a high co-occurrence of elements, suggesting that these features affect chunking and song learning. We examined experimentally whether the presence of pauses and element co-occurrence affect the ability of adult zebra finches to discriminate strings of song elements. Using a go/no-go design, two groups of birds were trained to discriminate between two strings. In one group (Pause-group), pauses were inserted between co-occurring element triplets in the strings, and in the other group (No-pause group), both strings were continuous. After making a correct discrimination, an individual proceeded to a reversal training using string segments. Segments were element triplets consistent in co-occurrence, triplets that were partly consistent in composition and triplets consisting of elements that did not co-occur in the strings. The Pause-group was faster in discriminating between the two strings. This group also responded differently to consistent triplets in the reversal training, compared to inconsistent triplets. The No-pause group did not differentiate among the triplet types. These results indicate that pauses in strings of song elements aid song discrimination and memorization of co-occurring element groups.

Zebra finches are sensitive to prosodic features of human speech.

Variation in pitch, amplitude and rhythm adds crucial paralinguistic information to human speech. Such prosodic cues can reveal information about the meaning or emphasis of a sentence or the emotional state of the speaker. To examine the hypothesis that sensitivity to prosodic cues is language independent and not human specific, we tested prosody perception in a controlled experiment with zebra finches. Using a go/no-go procedure, subjects were trained to discriminate between speech syllables arranged in XYXY patterns with prosodic stress on the first syllable and XXYY patterns with prosodic stress on the final syllable. To systematically determine the salience of the various prosodic cues (pitch, duration and amplitude) to the zebra finches, they were subjected to five tests with different combinations of these cues. The zebra finches generalized the prosodic pattern to sequences that consisted of new syllables and used prosodic features over structural ones to discriminate between stimuli. This strong sensitivity to the prosodic pattern was maintained when only a single prosodic cue was available. The change in pitch was treated as more salient than changes in the other prosodic features. These results show that zebra finches are sensitive to the same prosodic cues known to affect human speech perception.