mimicking several different native Australian bird calls The songs of different species of birds vary and are generally typical of the species. Species vary greatly in the complexity of their songs and in the number of distinct kinds of song they sing (up to 3000 in the
brown thrasher); individuals within some species vary in the same way. In a few species, such as
lyrebirds and
mockingbirds, songs imbed arbitrary elements learned in the individual's lifetime, a form of mimicry (though maybe better called "appropriation" (Ehrlich et al.), as the bird does not pass for another species). As early as 1773, it was established that birds learned calls, and
cross-fostering experiments succeeded in making linnet
Acanthis cannabina learn the song of a skylark,
Alauda arvensis. In many species, it appears that although the basic song is the same for all members of the species, young birds learn some details of their songs from their fathers, and these variations build up over generations to form
dialects. Song learning in juvenile birds occurs in two stages: sensory learning, which involves the juvenile listening to the father or other conspecific bird and memorizing the spectral and temporal qualities of the song (song template), and sensorimotor learning, which involves the juvenile bird producing its own vocalizations and practicing its song until it accurately matches the memorized song template. During the sensorimotor learning phase, song production begins with highly variable sub-vocalizations called "sub-song", which is akin to
babbling in human infants. Soon after, the juvenile song shows certain recognizable characteristics of the imitated adult song, but still lacks the stereotypy of the crystallized song – this is called "plastic song". After two or three months of song learning and rehearsal (depending on species), the juvenile produces a crystallized song, characterized by spectral and temporal stereotypy (very low variability in syllable production and syllable order). Some birds, such as
zebra finches, which are the most popular species for birdsong research, have overlapping sensory and sensorimotor learning stages. In some species such as zebra finches, learning of song is limited to the first year; they are termed "age-limited" or "close-ended" learners. Other species such as the canaries can develop new songs even as sexually mature adults; these are termed "open-ended" learners. Researchers have hypothesized that learned songs allow the development of more complex songs through cultural interaction, thus allowing intraspecies dialects that help birds to identify kin and to adapt their songs to different acoustic environments.
Auditory feedback in birdsong learning Early experiments by Thorpe in 1954 showed the importance of a bird being able to hear a tutor's song. When birds are raised in isolation, away from the influence of conspecific males, they still sing. While the song they produce, called "isolate song", resembles the song of a wild bird, it shows distinctly different characteristics from the wild song and lacks its complexity. The importance of the bird being able to hear itself sing in the sensorimotor period was later discovered by Konishi. Birds deafened before the song-crystallization period went on to produce songs that were distinctly different from the wild type and isolate song. Since the emergence of these findings, investigators have been searching for the neural pathways that facilitate sensory/sensorimotor learning and mediating the matching of the bird's own song with the memorized song template. Several studies in the 1990s have looked at the neural mechanisms underlying birdsong learning by performing lesions to relevant brain structures involved in the production or maintenance of song or by deafening birds before and/or after song crystallization. Another experimental approach was recording the bird's song and then playing it back while the bird is singing, causing perturbed auditory feedback (the bird hears the superposition of its own song and a fragmented portion of a previous song syllable). made a landmark discovery as they demonstrated that auditory feedback was necessary for the maintenance of song in adult birds with crystallized song, Leonardo & Konishi (1999) designed an auditory feedback perturbation protocol in order to explore the role of auditory feedback in adult song maintenance further, to investigate how adult songs deteriorate after extended exposure to perturbed auditory feedback, and to examine the degree to which adult birds could recover crystallized song over time after being removed from perturbed feedback exposure. This study offered further support for role of auditory feedback in maintaining adult song stability and demonstrated how adult maintenance of crystallized birdsong is dynamic rather than static. Brainard & Doupe (2000) posit a model in which LMAN (of the anterior forebrain) plays a primary role in error correction, as it detects differences between the song produced by the bird and its memorized song template and then sends an instructive error signal to structures in the vocal production pathway in order to correct or modify the motor program for song production. In their study, Brainard & Doupe (2000) showed that while deafening adult birds led to the loss of song stereotypy due to altered auditory feedback and non-adaptive modification of the motor program, lesioning LMAN in the anterior forebrain pathway of adult birds that had been deafened led to the stabilization of song (LMAN lesions in deafened birds prevented any further deterioration in syllable production and song structure). Currently, there are two competing models that elucidate the role of LMAN in generating an instructive error signal and projecting it to the motor production pathway: ''Bird's own song (BOS)-tuned error correction model'' : During singing, the activation of LMAN neurons will depend on the match between auditory feedback from the song produced by the bird and the stored song template. If this is true, then the firing rates of LMAN neurons will be sensitive to changes in auditory feedback.
Efference copy model of error correction : An
efference copy of the motor command for song production is the basis of the real-time error-correction signal. During singing, activation of LMAN neurons will depend on the motor signal used to generate the song, and the learned prediction of expected auditory feedback based on that motor command. Error correction would occur more rapidly in this model. Leonardo tested these models directly by recording spike rates in single LMAN neurons of adult zebra finches during singing in conditions with normal and perturbed auditory feedback. His results did not support the BOS-tuned error correction model, as the firing rates of LMAN neurons were unaffected by changes in auditory feedback and therefore, the error signal generated by LMAN appeared unrelated to auditory feedback. Moreover, the results from this study supported the predictions of the efference copy model, in which LMAN neurons are activated during singing by the efference copy of the motor signal (and its predictions of expected auditory feedback), allowing the neurons to be more precisely time-locked to changes in auditory feedback.
Mirror neurons and vocal learning A
mirror neuron is a
neuron that discharges both when an individual performs an action and when he/she perceives that same action being performed by another. These neurons were first discovered in
macaque monkeys, but recent research suggests that mirror neuron systems may be present in other animals including humans. Mirror neurons have the following characteristics: This has implications for birdsong learning– many birds rely on auditory feedback to acquire and maintain their songs. Mirror neurons may be mediating this comparison of what the bird hears, how it compares to a memorized song template, and what he produces. In search of these auditory-motor neurons, Jonathan Prather and other researchers at Duke University recorded the activity of single neurons in the
HVCs of
swamp sparrows. This may be the mechanism underlying learning via auditory feedback. These findings are also in line with Leonardo's (2004) efference copy model of error correction in birdsong learning and production. Overall, the HVCX auditory motor neurons in swamp sparrows are very similar to the visual motor mirror neurons discovered in
primates. Like mirror neurons, the HVCX neurons: • Are located in a
premotor brain area • Exhibit both sensory and motor properties • Are action-specific – a response is only triggered by the "primary song type" The function of the mirror neuron system is still unclear. Some scientists speculate that mirror neurons may play a role in understanding the actions of others,
imitation,
theory of mind and
language acquisition, though there is currently insufficient
neurophysiological evidence in support of these theories.
Learning through cultural transmission Culture in animals is usually defined to consist of socially transmitted behavior patterns ("traditions") that are characteristic of certain populations. The learned nature of bird song as well as evidence of "dialect"-like local variations have support theories about the existence of
avian culture. This suggested that many aspects of song development in songbirds depends on tutoring by older members of the same species. Later studies observed canary-like elements in the song of a chaffinch raised by
canaries, evidencing the strong role of tutors in the learning of song by juvenile birds. Similar chaffinch song types (categorized based on their distinct elements and their order) were observed to cluster in similar geographic areas, and this discovery led to hypotheses about "dialects" in birdsong. It has since been postulated that these song type variations are not
dialects like those we found in human language. This is because not all members of a given geographic area will conform to the same song type, and also because there is no singular characteristic of a song type that differentiates it from all other types (unlike human dialects where certain words are unique to certain dialects). Their data shows that certain brain areas in juvenile zebra finches are excited by the singing of conspecific tutors and not by loudspeakers playing zebra finch song. Additionally, they show that dopamine released into the HVC aids in the encoding of song.
Evolutionary preservation of bird vocal learning The cultural trap hypothesis Although a significant amount of research was done on bird song during the 20th century, none was able to elucidate the evolutionary "use" behind birdsong, especially with regards to large vocal repertoires. In response, Lachlan and Slater proposed a "cultural trap" model to explain persistence of wide varieties of song. This model is based on a concept of "filters", in which: • a male songbird's (i.e. singer's) filter contains the range of songs that it can develop • a female songbird's (i.e. receiver's) filter contains the range of songs that it finds acceptable for
mate choice In one possible situation, the population consists mainly of birds with wide filters. In this population, a male songbird with a wide filter will rarely be chosen by the few females with narrow filters (as the male's song is unlikely to fall within a narrower filter). Such females will have a relatively small choice of males to mate with, so the genetic basis of the females' narrow filter does not persist. Another possible situation deals with a population with mostly narrow filters. In the latter population, wide-filter males can feasibly avoid mate choice rejection by learning from older, narrow-filter males. Therefore, the average reproductive success of wide-filter birds is enhanced by the possibility of learning, and vocal learning and large song repertoires (i.e. wide filters) go hand-in-hand. This has led some researchers to hypothesize that sexual selection for more complex songs indirectly selects for stronger cognitive ability in males. Further investigation showed that male
song sparrows with larger vocal repertoires required less time to solve detour-reaching cognitive tasks. Some have proposed that bird song (among other sexually selected traits such as flashy coloring, body symmetry, and elaborate courtship) allow female songbirds to quickly assess the cognitive skills and development of multiple males. ==Identification and systematics==