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Social learning in birds

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Various studies of birds have positied animal social learning abilties. Bird song is a good example. Whilst bird song is affected by biological factors such as hormones and maturation, birds in different regions have different "dialects" which are explainable by social learning. Genetic explanations like differences caused by genetic drift or selection have been ruled out by studies which transfer young chicks to different regions, which then pick up the dialect of the environment. It has been shown that birds have a sensitive period (which varies across species) where the relocation of birds has an effect. [1] Some have argued that this represents a "special" form of social learning which is functionally and evolutionarily constricted to learning songs.  [2] Using the two-action paradigm, 

In Britian, a noticable trend was observed among blue tits, which began opening milk bottles left outside houses. The rapid spread of this behavior suggests a possible social learning explanation. However, naive chickadees (Parus atricapillus) have been shown to individual learn this ability relatively often (4/16 birds opened similar bottles without social learning) [3]. However, in the aforementioned experiment, milk bottles are put into small cages, which makes the liklihood of an individual opening the bottle a lot higher- it is possible that individual discoveries led to a local enhancement effect of birds drinking from milk bottles drawing the attention of others, which then performed the behavior. 

Stimulus learning and response learning by observation in the European starling, in a two-object/two-action test. Animal Behaviour, 58, 151-158.</ref> Another study conducted with pigeons assessed imitation of pecking a treadle for food compared to stepping on the treadle. In the stepping condition, 9 pigeons imitated the demonstrator, and one failed to complete the task. In the pecking condition, 5 stepped on the treadle, and 5 pecked at it. In Japanese quail, observers significantly pecked the treadle more in the treadle condition, however in the stepping condition, the individuals did not use either method significantly more often. Campbell has argued that these results could be explained by a mixture of contagion and local enhancement- that the birds were more likely to go near the treadle due to local enhancement, and pecking behaviors increased because of the well-known contagion effect of pecking in birds. [4]

File:Bird song development timeline.svg

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.[6] 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.[7]

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.[8] 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".[9] Finally, 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).[10] Some birds, such as Zebra Finches, which are the most popular species for birdsong research, have overlapping sensory and sensorimotor learning stages.[5]

Research has indicated that birds' acquisition of song is a form of motor learning that involves regions of the basal ganglia. Further, the PDP has been considered homologous to a mammalian motor pathway originating in the cerebral cortex and descending through the brain stem, while the AFP has been considered homologous to the mammalian cortical pathway through the basal ganglia and thalamus.[9] Models of bird-song motor learning can be useful in developing models for how humans learn speech.[11] 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.[12][13]

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.[14]


See alsoEdit

ReferencesEdit

  1. Soha, J.A. (2004) Genetic analysis of song dialect populations in Puget Sound white-crowned sparrows. Behavioral Ecology, 15, 4, 636-646.
  2. Zentall, T.R. (2006) Imitation: definition, evidnece and mechanisms. Animal Cognition, 9, p.342.
  3. Sherry, D.F & Galef, B.G. (1984) Cultural Transmission Without Imitation: Milk Bottle Opening by Birds. Animal Behavior, 32, 937.
  4. Campbell, F.M. et al. (1999) Stimulus learning and response learning by observation in the European starling, in a two-object/two-action test. Animal Behaviour, 58, 151-158.
  5. 5.0 5.1 Brainard, M. S. and Doupe, A. J. (2002). What songbirds teach us about learning. Nature 417 (6886): 351–358.
  6. Barrington, D. (1773). Experiments and observations on the singing of birds. Philosophical Transactions of the Royal Society of London 63: 249–291.
  7. Marler, P., & M. Tamura (1962). Song dialects in three populations of the white-crowned sparrow. Condor 64 (5): 368–377.
  8. Konishi, M. (2010). From central pattern generator to sensory template in the evolution of birdsong. Brain & Language 15: 18–20.
  9. 9.0 9.1 Nottebohm,F. (2005). The Neural Basis of Birdsong. PLoS Biol 3 (5).
  10. Leonardo, A., & Konishi, M. (1999). Decrystallization of adult birdsong by perturbation of auditory feedback. Nature 399 (6735): 466–470.
  11. Teramitsu I, Kudo LC, London SE, Geschwind DH, White SA (2004). Parallel FoxP1 and FoxP2 expression in songbird and human brain predicts functional interaction. J. Neurosci. 24 (13): 3152–63.
  12. Nottebohm, F. (2004). The road we travelled: discovery, choreography, and significance of brain replaceable neurons. Ann. N. Y. Acad. Sci. 1016: 628–658.
  13. Brenowitz, Eliot A. and Michael D. Beecher (2005). Song learning in birds: diversity and plasticity, opportunities and challenges. Trends in Neurosciences 28 (3): 127–132.
  14. Slater, P. J. B. (1989). Bird song learning: causes and consequences. Ethol. Ecol. Evol. 1: 19–46.

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