Bird vocalization

"Bird song" redirects here. For other uses, see Birdsong (disambiguation).

A male Blackbird (Turdus merula) singing. Bogense havn, Funen, Denmark.  Blackbird song recorded at Lille, France (help·info)

Bird vocalization includes both bird calls and bird songs. In non-technical use, bird songs are the bird sounds that are melodious to the human ear. In ornithology and birding, (relatively complex) songs are distinguished by function from (relatively simple) calls.

Definition

Wing feathers of a male Club-winged Manakin, with the modifications noted by P L Sclater in 1860^[1] and discussed by Charles Darwin in 1871^[2]

The distinction between songs and calls is based upon complexity, length, and context. Songs are longer and more complex and are associated with courtship and mating, while calls tend to serve such functions as alarms or keeping members of a flock in contact.^[3] Other authorities such as Howell and Webb (1995) make the distinction based on function, so that short vocalizations such as those of pigeons and even non-vocal sounds such as the drumming of woodpeckers and the "winnowing" of snipes' wings in display flight are considered songs.^[4] Still others require song to have syllabic diversity and temporal regularity akin to the repetitive and transformative patterns which define music. It is generally agreed upon in birding and ornithology which sounds are songs and which are calls, and a good field guide will differentiate between the two.

Bird song is best developed in the order Passeriformes. Most song is emitted by male rather than female birds. Song is usually delivered from prominent perches although some species may sing when flying. Some groups are nearly voiceless, producing only percussive and rhythmic sounds, such as the storks, which clatter their bills. In some manakins (Pipridae), the males have evolved several mechanisms for mechanical sound production, including mechanisms for stridulation not unlike those found in some insects.^[5]

The production of sounds by mechanical means as opposed to the use of the syrinx has been termed variously instrumental music by Charles Darwin, mechanical sounds^[6] and more recently sonation.^[7] The term sonate has been defined as the act of producing non-vocal sounds that are intentionally modulated communicative signals, produced using non-syringeal structures such as the bill, wings, tail, feet and body feathers.^[7]

Anatomy

The avian vocal organ is called the syrinx; it is a bony structure at the bottom of the trachea (unlike the larynx at the top of the mammalian trachea). The syrinx and sometimes a surrounding air sac resonate to sound waves that are made by membranes past which the bird forces air. The bird controls the pitch by changing the tension on the membranes and controls both pitch and volume by changing the force of exhalation. It can control the two sides of the trachea independently, which is how some species can produce two notes at once.

Function

Scientists hypothesize that bird song has evolved through sexual selection, and experiments suggest that the quality of bird song may be a good indicator of fitness.^[8] Experiments also suggest that parasites and diseases may directly affect song characteristics such as song rate, which thereby act as reliable indicators of health.^[9][10] The song repertoire also appears to indicate fitness in some species.^[11][12] The ability of male birds to hold and advertise territories using song also demonstrates their fitness.

Communication through bird calls can be between individuals of the same species or even across species. Birds communicate alarm through vocalizations and movements that are specific to the threat, and bird alarms can be understood by other animal species, including other birds, in order to identify and protect against the specific threat.^[13] Mobbing calls are used to recruit individuals in an area where an owl or other predator may be present. These calls are characterized by wide frequency spectra, sharp onset and termination, and repetitiveness which are common across species and are believed to be helpful to other potential "mobbers" by being easy to locate. The alarm calls of most species, on the other hand, are characteristically high-pitched making the caller difficult to locate.^[14]

Individual birds may be sensitive enough to identify each other through their calls. Many birds that nest in colonies can locate their chicks using their calls.^[15] Calls are sometimes distinctive enough for individual identification even by human researchers in ecological studies.^[16]

Many birds engage in duet calls. In some cases the duets are so perfectly timed as to appear almost as one call. This kind of calling is termed antiphonal duetting.^[17] Such duetting is noted in a wide range of families including quails,^[18] bushshrikes,^[19] babblers such as the scimitar babblers, some owls^[20] and parrots.^[21] In territorial songbirds, birds are more likely to countersing when they have been aroused by simulated intrusion into their territory.^[22] This implies a role in intraspecies aggressive competition.

Some birds are excellent vocal mimics. In some tropical species, mimics such as the drongos may have a role in the formation of mixed-species foraging flocks.^[23] Vocal mimicry can include conspecifics, other species or even man-made sounds. Many hypotheses have been made on the functions of vocal mimicry including suggestions that they may be involved in sexual selection by acting as an indicator of fitness, help brood parasites, protect against predation but strong support is lacking for any function.^[24] Many birds, and especially those that nest in cavities, are known to produce a snake like hissing sound that may help deter predators at close range.^[25]

Some cave-dwelling species, including Oilbird^[26] and Swiftlets (Collocalia and Aerodramus spp.),^[27] use audible sound (with the majority of sonic location occurring between 2 and 5 kHz^[28]) to echolocate in the darkness of caves. The only bird known to make use of infrasound (at about 20 Hz) is the western capercaillie.^[29]

The hearing range of birds is from below 50 Hz (infrasound) to above 20 kHz (ultrasound) with maximum sensitivity between 1 and 5 kHz.^[12][30] The range of frequencies at which birds call in an environment varies with the quality of habitat and the ambient sounds. It has been suggested that narrow bandwidths, low frequencies, low-frequency modulations, and long elements and inter-element intervals should be found in habitats with complex vegetation structures (which would absorb and muffle sounds) while high frequencies, broad bandwidth, high-frequency modulations (trills), and short elements and inter-elements may be expected in habitats with herbaceous cover.^{[vague] [31][32]} It has been hypothesized that the available frequency range is partitioned and birds call so that overlap between different species in frequency and time is reduced. This idea has been termed the "acoustic niche".^[33] Birds sing louder and at a higher pitch in urban areas, where there is ambient low-frequency noise.^[34][35]

Bird Language

The language of the birds has long been a topic for anecdote and speculation. That calls have meanings that are interpreted by their listeners has been well demonstrated. Domestic chickens have distinctive alarm calls for aerial and ground predators, and they respond to these alarm calls appropriately.^[36][37] However a language has, in addition to words, structures and rules. Studies to demonstrate the existence of language have been difficult due to the range of possible interpretations. Research on parrots by Irene Pepperberg is claimed to demonstrate the innate ability for grammatical structures, including the existence of concepts such as nouns, adjectives and verbs.^[38] Studies on starling vocalizations have also suggested that they may have recursive structures.^[39]

The term "bird language" may also more informally refer to patterns in bird vocalizations that communicate information to other birds or other animals in general.^[40] Wilderness Awareness School groups bird vocalizations into five different classes, sometimes called "voices," each of which communicates different information.^[41] Song has been discussed at length in this article. Companion calling is a short vocalization made between mates, parent and young, or members of a flock to maintain contact when out of visual range. Juvenile begging is a strident, loud vocalization often made by young to a parent when begging for food. Intraspecific aggression can consist of loud, alarmed-sounding vocalizations or of energetic song, and may be heard when members of the same species behave aggressively toward each other. Alarm may be heard when birds are startled, frightened, or terrified for their lives, and can take many forms. Mobbing is one example of alarm, while a high-pitched alarm call is another.

Of the five voices of the birds, four of them communicate the message that the bird feels safe^{[citation needed]}. Birds that engage in song, companion calling, juvenile begging, and intraspecific aggression all display what Jon Young calls "baseline" behavior, or a relaxed state free of the fear of predation.^[42] Alarm communicates the presence of a predator, or an influence that the bird may see as predatory such as a human hiker. Alarms have distinct sounds and shapes, each of which is specific to the source of the disturbance.^[43] For example, ravens mobbing a hawk or owl in a tree will clump around the predator in a loose ball, calling and diving. If the ravens rise off the tree and fly higher, the predator was a hawk and has flown up to escape, as is typical of hawks. If the ravens drop out of the tree and fly low and away, the predator was an owl and has dropped low off its perch to escape, as is typical of owls.^[44]

Neurophysiology

The main brain areas involved in bird song are:

• Anterior forebrain pathway (vocal learning): composed of the lateral part of the magnocellular nucleus of anterior neostriatum (LMAN), which is a homologue to mammalian basal ganglia); Area X, which is part of the basal ganglia; and the Dorso-Lateral division of the Medial thalamus (DLM).
• Song production pathway: composed of the HVC (sometimes, inaccurately, called the hyperstriatum ventrale, pars caudalis); robust nucleus of the arcopallium (RA); and the tracheosyringeal part of the hypoglossal nucleus (nXIIts).^[45][46]

Both pathways show sexual dimorphism, with the male producing song most of the time.^[47] It has been noted that injecting testosterone in non-singing female birds can induce growth of the HVC and thus production of song.

Birdsong production is generally thought to start at the nucleus uvaeformis of the thalamus with signals emanating along a pathway that terminates at the syrinx. The pathway from the thalamus leads to the interfacial nucleus of the nidopallium to the HVC, and then to RA, the dorso-lateral division of the medial thalamus and to the tracheosyringeal nerve.

The gene FOXP2, defects of which affect both speech and comprehension of language in humans, becomes more active in the striatal region of songbirds during the time of song learning.^[48]

Recent research in birdsong learning has focused on the Ventral Tegmental Area (VTA), which sends a dopamine input to the para-olfactory lobe and Area X, LMAN and the ventrolateral medulla. Other researchers have explored the possibility that HVc is responsible for syllable production, while the robust nucleus of the arcopallium, the primary song output nucleus, may be responsible for syllable sequencing and production of notes within a syllable.^{[citation needed]}

Learning

The songs of different species of birds vary, and are more or less characteristic of the species. In modern-day biology, bird song is typically analysed using acoustic spectroscopy. 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); in some species, individuals vary in the same way. In a few species such as starlings 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 learnt calls and cross-fostering experiments were able to force a Linnet Acanthis cannabina to learn the song of a skylark Alauda arvensis.^[49] 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.^[50]

Birds learn songs early in life with sub-vocalizations that develop into renditions of adult songs. Zebra Finches, the most popular species for birdsong research, develop a version of a familiar adult's song after 20 or more days from hatch. By around 35 days, the chick will have learned the adult song. The early song is "plastic" or variable and it takes the young bird two or three months to perfect the "crystallized" song (which is less variable) of sexually mature birds.^[51]

Timeline for song learning in different species. Diagram adapted from Brainard & Doupe, 2002^[52]..

Research indicates birds' acquisition of song is a form of motor learning that involves regions of the basal ganglia. Models of bird-song motor learning are sometimes used as models for how humans learn speech.^[53] 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.^[54][55]

Researchers have hypothesized that learned songs allow the development of more complex songs through cultural interaction, thus allowing intraspecies dialects that help birds stay with their own kind within a species, and it allows birds to adapt their songs to different acoustic environments.^[56]

Auditory feedback in bird song 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 resembles the song of a wild bird, it lacks the complexity and sounds distinctly different.^[57] The importance of the bird being able to hear himself sing in the sensorimotor period was later discovered by Konishi. Birds deafened before the song crystallization period went on to produce very different songs from the wild type.^[58] These findings led scientists to believe there could be a specific part of the brain dedicated to this specific type of learning.

Song learning pathway in birds (Based on Nottebohm, 2005)

The main focus in the search for the neuronal aspect of bird song learning was guided by the song template hypothesis. This hypothesis is the idea that when a bird is young he memorizes the song of his tutor. Later, during the development phase as an adult, he matches his own trial vocalizations using auditory feedback to an acoustic template in the brain. Based on this information, he adjusts his song if needed. To find this "song template," experimenters lesioned certain parts of the brain and observed the effects.

• Lesioning the song production pathway (RA, xXII or HVc) in the brain creates serious effects on song production in all birds.^[59]
• Lesions parts of the anterior forebrain pathway, or vocal learning pathway, DLM and area X, result in deficits in learning in all birds.^[47]
• Lesioning LMAN, located in the anterior forebrain pathway in young birds disrupts song production.^[47]
• Lesioning LMAN on an adult bird shows no effect.^[47]
• Lesioning LMAN on an adult canary (an "open-ended learner" species, which can learn songs later in life) shows a progressive deterioration of song.^[47]

These results show that the area known as LMAN is the only brain area in the pathway that shows some plasticity and further studies have shown that this area of the brain responds best to the bird's own song.^[45][52][60] This neuroplasticity is vital for a bird being able to learn a song. The ability to make small adjustments based on auditory feedback is needed for the complexity of these beautiful songs. Just like any musician, birds need to practice and be able to evaluate what their song sounds like and what it's supposed to sound like in order to get it right.

To complete the picture on bird song learning, experimenters needed to discover the true plasticity of the brain. While deafening and creating auditory isolation were good techniques for discovering basic characteristics about the brain, a reversible procedure was needed to investigate further. The solution was found in disruption of the auditory feedback, or what a bird hears. A computer is able to capture the song of a singing bird and play back portions of its song, or selectively play back a certain syllable while the bird is singing. The computer is basically playing the age old trick of repeating whatever the bird sings, the "stop copying me" game. This creates such a disruption that an adult bird will start to decrystallize its song, which includes a loss of spectral and temporal rigidity characteristic of adult song. It reverts back to the song it started singing with, before any learning took place. Furthermore, when the feedback was stopped, the birds slowly recovered their original song, something that was unheard of. These results show that there is a fair amount of plasticity retained in the brain, even for close-ended learners.^[61] This new found plasticity in adult birds and the results on the plasticity of LMAN (shown above) combine into a model for bird song learning (diagram coming soon).^[62]

Mirror neurons and vocal learning

A mirror neuron is a neuron that discharges both when an individual performs an action, and when he perceives that same action being performed by another.^[63] These neurons were first discovered in macaque monkeys, but recent research suggests that mirror neuron systems may be present in other animals including humans.^[64]

Mirror neurons have the following characteristics:^[63]

• They are located the premotor cortex
• They exhibit both sensory and motor properties
• They are action-specific – mirror neurons are only active when an individual is performing or observing a certain type of action (e.g.: grasping an object).

Because mirror neurons exhibit both sensory and motor activity, some researchers have suggested that mirror neurons may serve to map sensory experience onto motor structures.^[65] 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 and what he produces.

Song selectivity in HVCx neurons: neuron activity in response to calls heard (green) and calls produced (red). a. Neurons fire when the primary song type is either heard or sung. b,c. Neurons do not fire in response to the other song type, regardless of whether it is heard or sung. Sketch based on figure from Prather et al. (2008)

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.^[66] They discovered that the neurons that project from the HVC to Area X (HVC_X neurons) are highly responsive when the bird is hearing a playback of his own song. These neurons also fire in similar patterns when the bird is singing that same song. Swamp sparrows employ 3-5 different song types, and the neural activity differs depending on which song is heard or sung. The HVC_X neurons only fire in response to the presentation (or singing) of one of the songs, the primary song type. They are also temporally selective, firing at a precise phase in the song syllable.

Because the timing of the neural response is identical regardless of whether the bird was listening or singing, how can we be sure that the bird isn’t just hearing himself? Prather et al. found that during the short period of time before and after the bird sings, his HVC_X neurons become insensitive to auditory input. In other words, the bird becomes "deaf" to his own song. This suggests that these neurons are producing a corollary discharge, which would allow for direct comparison of motor output and auditory input.^[67] This may be the mechanism underlying learning via auditory feedback.

Overall, the HVC_X auditory-motor neurons in swamp sparrows are very similar to the visual-motor mirror neurons discovered in primates. Like mirror neurons, the HVC_X 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.^[65] Specifically regarding birds, it is possible that the mirror neuron system serves as a general mechanism underlying vocal learning, but further research is needed. In addition to the implications for song learning, the mirror neuron system could also play a role in territorial behaviors such as song-type matching and countersinging.^[68]

Identification and systematics

Song of the White-throated Sparrow

The specificity of bird calls has been used extensively for species identification. The calls of birds have been described using words or nonsense syllables, or line diagrams.^[69] Common terms in English include words such as quack, chirp and chirrup. These are subject to imagination and vary greatly; a well-known example is the White-throated Sparrow's song, given in Canada as O sweet Canada Canada Canada and in New England as Old Sam Peabody Peabody Peabody (also Where are you Frederick Frederick Frederick?). In addition to nonsense words, grammatically correct phrases have been constructed as likenesses of the vocalizations of birds. For example, the Barred Owl produces a motif which some bird guides describe as Who cooks for you? Who cooks for you all? with the emphasis placed on you.^[70]

Sonogram of the call of a Laughing Dove.  Recorded in south India (help·info)

The use of spectrograms to visualize bird song was first introduced by W. H. Thorpe.^[71][72] These visual representations are also called sonograms or sonagrams. Some recent field guides for birds use sonograms to document the calls and songs of birds.^[73] The sonogram is objective, unlike descriptive phrases, but proper interpretation requires experience. Sonograms can also be roughly converted back into sound.^[74][75]

Bird song is an integral part of bird courtship and is a pre-zygotic isolation mechanism involved in the process of speciation. Many allopatric sub-species show differences in calls. These differences are sometimes minute, often detectable only in the sonograms. Song differences in addition to other taxonomic attributes have been used in the identification of new species.^[76] The use of calls has led to proposals for splitting of species complexes such as those of the Mirafra Bushlarks.^[77]

Recording

The first known recording of birdsong was made in 1889 by Ludwig Koch,^[78] who went on to become an eminent wildlife recordist and BBC natural history presenter.^[78]

Other notable birdsong recordists include Eric Simms and Chris Watson.

Bird song and music

Some musicologists believe that birdsong has had a large influence on the development of music.^[79] Although the extent of this influence is impossible to gauge,^[80] it is sometimes easy to see some of the specific ways composers have integrated birdsong with music.

There seem to be three general ways musicians or composers can be affected by birdsong: they can be influenced or inspired (consciously or unconsciously) by birdsong, they can include intentional imitations of bird song in a composition, or they can incorporate recordings of birds into their works.

In his book 'Why Birds Sing' David Rothernberg claims that birds vocalize traditional scales used in human music, such as the pentatonic scale (e.g. Hermit Thrush) and diatonic scale (e.g. Wood Thrush), providing evidence that birdsong not only sounds like music, but is music in the human sense. This claim has been refuted by Sotorrio ('Tone Spectra') who has shown that birds are not selecting 'scale tones' from a miriad of tonal possibilities, but are 'filtering out' and reinforcing the available set of overtones from the fundamental tones of their vocal chords. This requires "far less musical intelligence and deliberate appropriation" and in this regard, he suggests birdsong has something in common with Mongolian throat-singing and jaw-harp music. Sotorrio also claims that musicians like Rothernberg are deceived by "a perculiar form of Pareidolia" whereby complex tonal information is reduced to human scale concepts due to a "fixation on music as it is written rather than as it sounds". Rothernberg's claims were expored in the BBC documentary 'Why Birds Sing'.

Compositions that imitate or use birdsong

One early example of a composition that imitates birdsong is Janequin's "Le Chant Des Oiseaux", written in the 16th century.

Other composers who have quoted birds or have used birdsong as a compositional springboard include Vivaldi (Spring from the Four Seasons)), Biber (Sonata Representativa), Beethoven (Sixth Symphony), Wagner (Siegfried) and the jazz musicians Paul Winter (Flyway) and Jeff Silverbush (Grandma Mickey).^[81]

The twentieth-century French composer Olivier Messiaen composed with birdsong extensively. His Catalogue d'Oiseaux is a seven-book set of solo piano pieces based upon birdsong. His orchestral piece Réveil des Oiseaux is composed almost entirely of birdsong. Many of his other compositions, including Quatuor pour la fin du temps, similarly integrate birdsong.^[82]

The Italian composer Ottorino Respighi, with his The Pines of Rome (1923–1924), may have been the first to compose a piece of music that calls for pre-recorded birdsong. A few years later, Respighi wrote Gli Uccelli ("The birds"), based on Baroque pieces imitating birds.

The Finnish composer Einojuhani Rautavaara in 1972 wrote an orchestral piece of music called Cantus Arcticus (Opus 61, dubbed Concerto for Birds and Orchestra) making extensive use of pre-recorded birdsongs from Arctic regions, such as migrating swans.

The American jazz musician Eric Dolphy sometimes listened to birds while he practiced flute. He claimed to have incorporated bird song into some of his improvisational music.

In the 1960s and 1970s, many rock bands included sound effects in their recordings. Birds were a popular choice. The English band Pink Floyd included bird sound effects in many of the songs from their 1969 albums Soundtrack from the Film More and Ummagumma (for example, Grantchester Meadows). Similarly, the English singer Kate Bush incorporated bird sound effects into much of the music on her 2005 album, Aerial.

The Music hall artist Ronnie Ronalde has gained notoriety for his whistling imitations of birds and for integrating birdsong with human song. His songs 'In A Monastery Garden' and 'If I Were A Blackbird' include imitations of the blackbird, his "signature bird."^{[citation needed]}

The French composer François-Bernard Mâche has been credited with the creation of zoomusicology, the study of the music of animals. His essay Musique, mythe, nature, ou les Dauphins d'Arion (1983) includes a study of "ornitho-musicology", in which he speaks of "animal musics" and a longing to connect with nature.

The German DJ, techno music producer and naturalist Dominik Eulberg is an avid bird watcher, and several tracks by him prominently feature sampled bird sounds and even are titled after his favourite specimens.^{[citation needed]}

The productions of The Jewelled Antler Collective often use field recordings featuring birdsong.

In 2007, The CT Collective issed two free albums devoted to music made using bird songs (one with human interaction, one without). The project was co-ordinated by looping musician Nick Robinson

Bird song and poetry

Bird song is a popular subject in poetry. Famous poems inspired by bird song include Percy Bysshe Shelley's To a Skylark ("Hail to thee, blithe Spirit!/Bird thou never wert") and Gerard Manley Hopkins' Sea and Skylark. Birdsongs and their relations to Middle-earth inhabitants are a common motif in J. R. R. Tolkien's literary work.^{[citation needed]} The Grateful Dead performed a song called "Bird Song" that Jerry Garcia wrote and dedicated to Janis Joplin.

See also

• Animal communication
• Animal language
• Bioacoustics
• Biomusic
• Biophony
• Cock a doodle doo
• Dawn chorus
• Flight Call
• Language of the birds
• Lateralization of bird song
• Lombard effect
• Talking bird
• Vinkenzetting
• Vocal learning

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External links

• Bird Language: Exploring the Language of Nature with Jon Young A blog with stories and tips for learning the patterns in bird vocalizations.
• Large collection of audio bird calls collected in Arizona from Ask A Biologist.
• Community online database of downloadable bird sounds from around the globe ~50000 recordings of ~6700 species as of May 2010.
• British Library's archive of bird sounds representing more than 8,000 species.
• Listen to Nature includes article "The Language of Birds"
• Bird songs in movies: an unnatural history Humor piece on soundtrack errors
• How do Birds Sing? The mechanics and anatomy of bird song production
• Bioacoustic Research Program at the Cornell Lab of Ornithology distributes a number of different free bird song synthesis & analysis programs.
• Macaulay Library at the Cornell Lab of Ornithology is the world's largest collection of animal sounds and associated video.
• Male shama songs and mimic of sounds
• Audio Pitch Tracer Accurate transcription of clean recordings of bird vocalizations to midi