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Bird vocalization

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

Bird vocalization   includes both   bird   calls and bird

A male Blackbird  (Turdus merula) singing. Bogense havn, Fu-

nen, Denmark.  Blackbird song recorded at Lille, France

songs. In non-technical use, bird songs are the bird

sounds that are melodious to the human ear. Inornithology and  birding, (relatively complex) songs aredistinguished by function from (relatively simple) calls.

1 Definition

The distinction between  songs  and  calls  is based uponcomplexity, length, and context. Songs are longer andmore complex and are associated with courtship and mat-ing, while calls tend to serve such functions as alarmsor keeping members of a  flock in contact.[3] Other au-

thorities such as Howell and Webb (1995) make thedistinction based on function, so that short vocaliza-tions, such as those of pigeons, and even non-vocalsounds, such as the drumming of woodpeckers and theextquotedblwinnowing  extquotedbl of   snipes' wings indisplay flight, are considered songs.[4] Still others requiresong to have syllabic diversity and temporal regularityakin to the repetitive and transformative patterns that de-fine music. It is generally agreed upon in birding and or-nithology which sounds are songs and which are calls, anda good field guide will differentiate between the two.

Bird song is best developed in the order Passeriformes.

Some groups are nearly voiceless, producing onlypercussive and rhythmic sounds, such as the storks, whichclatter their bills. In some manakins (Pipridae), the males

Wing feathers of a male Club-winged Manakin , with the modifi-cations noted by P. L. Sclater in 1860[1] and discussed by Charles 

Darwin in 1871[2]

have evolved several mechanisms for mechanical soundproduction, including mechanisms for stridulation not un-like those found in some insects.[5]

Song is usually delivered from prominent perches, al-though some species may sing when flying. The produc-tion of sounds by mechanical means as opposed to the useof the syrinx has been termed variously instrumental mu-sic  by Charles Darwin, mechanical sounds [6] and more re-

cently sonation.[7] The term sonate has been defined as theact of producing non-vocal sounds that are intentionallymodulated communicative signals, produced using non-syringeal structures such as the bill, wings, tail, feet andbody feathers.[7]

In extratropical Eurasia and the Americas almost all songis produced by male birds; however in the tropics and toa greater extent the desert belts of Australia and Africait is more typical for females to sing as much as males.These differences have been known for a long time[8][9]

and are generally attributed to the much less regular andseasonal climate of Australian and African arid zones

requiring that birds breed at any time when conditionsare favourable, although they cannot breed in many yearsbecause food supply never increases above a minimum

1

https://en.wikipedia.org/wiki/Africahttps://en.wikipedia.org/wiki/Australiahttps://en.wikipedia.org/wiki/Deserthttps://en.wikipedia.org/wiki/The_Americashttps://en.wikipedia.org/wiki/Eurasiahttps://en.wikipedia.org/wiki/Sonationhttps://en.wikipedia.org/wiki/Charles_Darwinhttps://en.wikipedia.org/wiki/Syrinx_(biology)https://en.wikipedia.org/wiki/Stridulationhttps://en.wikipedia.org/wiki/Club-winged_Manakinhttps://en.wikipedia.org/wiki/Pipridaehttps://en.wikipedia.org/wiki/Storkhttps://en.wikipedia.org/wiki/Rhythmhttps://en.wikipedia.org/wiki/Percussivehttps://en.wikipedia.org/wiki/Passerinehttps://en.wikipedia.org/wiki/Order_(biology)https://en.wikipedia.org/wiki/Musichttp://-/?-https://en.wikipedia.org/wiki/Snipehttps://en.wikipedia.org/wiki/Drumming_(snipe)https://en.wikipedia.org/wiki/Woodpeckerhttps://en.wikipedia.org/wiki/Herdhttps://en.wikipedia.org/wiki/Bird#Breedinghttps://en.wikipedia.org/wiki/Bird#Breedinghttps://en.wikipedia.org/wiki/Birdinghttps://en.wikipedia.org/wiki/Ornithologyhttps://en.wikipedia.org/wiki/Media:Turdus_merula_male_song_at_dawn(20s).ogghttps://en.wikipedia.org/wiki/Common_Blackbirdhttps://en.wikipedia.org/wiki/Birdhttps://en.wikipedia.org/wiki/Birdsong_(disambiguation)https://en.wikipedia.org/wiki/Birdsong_(disambiguation)

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2   3 FUNCTION 

level.[8] With aseasonal irregular breeding, both sexesmust be brought into breeding condition and vocalisation,especially duetting, serves this purpose. The high fre-quency of female vocalisations in the tropics, Australiaand Southern Africa may also relate to very low mor-tality rates producing much stronger pair-bonding and

territoriality.[10]

2 Anatomy and physiology

The avian vocal organ is called the  syrinx; it is a bonystructure at the bottom of the trachea (unlike the larynxat the top of the  mammalian  trachea). The syrinx andsometimes a surrounding air sac resonate to sound wavesthat are made by membranes past which the bird forcesair. The bird controls the pitch by changing the tensionon the membranes and controls both pitch and volume bychanging the force of exhalation. It can control the twosides of the trachea independently, which is how somespecies can produce two notes at once.

3 Function

Scientists hypothesize that bird song has evolved throughsexual selection, and experiments suggest that the qualityof bird song may be a good indicator of fitness.[11] Ex-periments also suggest that parasites and diseases may di-

rectly affect song characteristics such as song rate, whichthereby act as reliable indicators of health.[12][13] Thesong repertoire also appears to indicate fitness in somespecies.[14][15] The ability of male birds to hold and ad-vertise territories using song also demonstrates their fit-ness.

Communication through bird calls can be between indi-viduals of the same species or even across species. Birdscommunicate alarm through vocalizations and move-ments that are specific to the threat, and bird alarms canbe understood by other animal species, including otherbirds, in order to identify and protect against the specific

threat.

[16]

Mobbing calls are used to recruit individuals inan area where an owl or other predator may be present.These calls are characterized by wide-frequency spectra,sharp onset and termination, and repetitiveness that arecommon across species and are believed to be helpful toother potential “mobbers” by being easy to locate. Thealarm calls of most species, on the other hand, are char-acteristically high-pitched, making the caller difficult tolocate.[17]

Individual birds may be sensitive enough to identify eachother through their calls. Many birds that nest in coloniescan locate their chicksusing their calls.[18] Calls are some-

times distinctive enough for individual identification evenby human researchers in ecological studies.[19]

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.[20]

Such duetting is noted in a wide range of families in-cluding quails,[21] bushshrikes,[22] babblers   such as thescimitar babblers, some owls[23] and parrots.[24] In terri-torial songbirds, birds are more likely to countersing when

they have been aroused by simulated intrusion into theirterritory.[25] This implies a role in intraspecies aggressivecompetition.

Sometimes, songs vocalized in post-breeding seasonact as a cue to conspecific eavesdroppers.[26] In black-throated blue warblers, males that have bred and repro-duced successfully singto their offspring to influence theirvocal development, while males that have failed to repro-duce usually abandon the nests and stay silent. The post-breeding song therefore inadvertently informs the unsuc-cessful males of particular habitats that have a higher like-lihood of reproductive success. The social communica-

tion by vocalization provides a shortcut to locating highquality habitats and saves the trouble of directly assess-ing various vegetation structures.

Some birds are excellent vocal mimics. In some tropicalspecies, mimics such as the drongos may have a role inthe formation of mixed-species foraging flocks.[27] Vocalmimicry can include conspecifics, other species or evenman-made sounds. Many hypotheses have been madeon the functions of vocal mimicry including suggestionsthat they may be involved in sexual selection by actingas an indicator of fitness, help brood parasites, or pro-tect against predation, but strong support is lacking for

any function.[28] Many birds, especially those that nest incavities, are known to produce a snakelike hissing soundthat may help deter predators at close range.[29]

Some cave-dwelling species, including the Oilbird[30] andswiftlets (Collocalia and  Aerodramus  spp.),[31] use audi-ble sound (with the majority of sonic location occurringbetween 2 and 5 kHz[32]) to echolocate in the darknessof caves. The only bird known to make use of infrasound(at about 20 Hz) is the western capercaillie.[33]

The hearing range of birds is from below 50 Hz (infra-sound) to around 12 kHz, with maximum sensitivity be-tween 1 and 5 kHz.[15][34]

The range of frequencies at which birds call in an envi-ronment varies with the quality of habitat and the ambientsounds. The acoustic adaptation hypothesis predicts thatnarrow bandwidths, low frequencies, and long elementsand inter-element intervals should be found in habitatswith complex vegetation structures (which would absorband muffle sounds), while high frequencies, broad band-width, high-frequency modulations (trills), and short ele-ments and inter-elements may be expected in open habi-tats, without obstructive vegetation.[35][36][37] Low fre-quency songs are optimal for obstructed, densely vege-tated habitats because low frequency, slowly modulatedsong elements are less susceptible to signal degradationby means of reverberations off of sound-reflecting veg-

http://-/?-https://en.wikipedia.org/wiki/Western_capercailliehttps://en.wikipedia.org/wiki/Infrasoundhttps://en.wikipedia.org/wiki/Animal_echolocationhttps://en.wikipedia.org/wiki/Aerodramushttps://en.wikipedia.org/wiki/Collocaliahttps://en.wikipedia.org/wiki/Oilbirdhttps://en.wikipedia.org/wiki/Mixed-species_feeding_flockhttps://en.wikipedia.org/wiki/Drongohttps://en.wikipedia.org/wiki/Mimicryhttps://en.wikipedia.org/wiki/Animal_communicationhttps://en.wikipedia.org/wiki/Animal_communicationhttps://en.wikipedia.org/wiki/Black-throated_blue_warblerhttps://en.wikipedia.org/wiki/Black-throated_blue_warblerhttps://en.wikipedia.org/wiki/Conspecificityhttps://en.wiktionary.org/wiki/countersinghttps://en.wikipedia.org/wiki/Scimitar_babblerhttps://en.wikipedia.org/wiki/Old_world_babblerhttps://en.wikipedia.org/wiki/Bushshrikehttps://en.wikipedia.org/wiki/Mobbing_behaviourhttps://en.wikipedia.org/wiki/Territory_(animal)http://-/?-https://en.wikipedia.org/wiki/Sexual_selectionhttps://en.wikipedia.org/wiki/Mammalhttps://en.wikipedia.org/wiki/Larynxhttps://en.wikipedia.org/wiki/Vertebrate_tracheahttps://en.wikipedia.org/wiki/Syrinx_(biology)

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3

etation. High frequency calls with rapid modulationsare optimal for open habitats because they degrade lessacross open space.[38][39] The acoustic adaptation hypoth-esis also states that song characteristics may take advan-tage of beneficial acoustic properties of the environment.Narrow-frequency bandwidth notes are increased in vol-

ume and length by reverberations in densely vegetatedhabitats.[40]

It has been hypothesized that the available frequencyrange is partitioned, and birds call so that overlap be-tween different species in frequency and time is re-duced. This idea has been termed the “acoustic niche”.[41]

Birds sing louder and at a higher pitch in urban areas,where there is ambient low-frequency noise.[42][43] Traf-fic noise was found to decrease reproductive success inthe Great Tit (Parsus major ) due to the overlap in acousticfrequency.[44] An increase in song volume restored fitnessto birds in urban areas, as did higher frequency songs.[45]

4 Learning

Seasonal Species

Age-limited

Learners

Open-ended

Learners

      H     a      t     c       h       i     n     g

      H     a      t     c       h       i     n     g

      H     a      t     c       h       i     n     g

Sensory Sensorimotor Crystallized

Spring Fall Spring

Sensorimotor Crystallized

20 Days 60 Days 90 Days

CrystallizedCrystallized SensorimotorSensorimotor

Sensory Sensory

Spring SpringSpringFall Fall

Timeline for song learning in different species. Diagram adapted 

 from Brainard & Doupe, 2002.[46]

The songs of different species of birds vary and are gen-erally typical of the species. Species vary greatly inthe complexity of their songs and in the number of dis-tinct kinds of song they sing (up to 3000 in the BrownThrasher); individuals within some species vary in thesame way. In a few species, such as   lyrebirds   andmockingbirds, songs imbed arbitrary elements learnedin the individual’s lifetime, a form of mimicry (thoughmaybe better called “appropriation” [Ehrlich et al.], asthe bird does not pass for another species). As early as1773, it was established that birds learned calls, andcross-fostering experiments succeeded in making linnet Acan-this cannabina   learn the song of a skylark,   Alauda ar-vensis .[47] 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 theirfathers, and these variations build up over generations to

form dialects.[48]

Song learning in juvenile birds occurs in two stages: sen-sory learning, which involves the juvenile listening tothe father or other conspecific bird and memorizing thespectral and temporal qualities of the song (song tem-

plate), and sensorimotor learning, which involves the ju-venile bird producing its own vocalizations and prac-ticing its song until it accurately matches the memo-rized song template.[49] During the sensorimotor learningphase, song production begins with highly variable sub-vocalizations called “sub-song”, which is akin to babblingin human infants.[50] Soon after, the juvenile song showscertain recognizable characteristics of the imitated adultsong, but still lacks the stereotypy of the crystallizedsong – this is called “plastic song”.[51] Finally, aftertwo or three months of song learning and rehearsal (de-pending on species), the juvenile produces a crystallizedsong, characterized by spectral and temporal stereotypy

(very low variability in syllable production and syllableorder).[52] Some birds, such as Zebra Finches, which arethe most popular species for birdsong research, have over-lapping sensory and sensorimotor learning stages.[46]

Research has indicated that birds’ acquisition of song isa form of   motor learning   that involves regions of thebasal ganglia. Further, the PDP (see Neuroanatomy be-low) has been considered  homologous to a mammalianmotor pathway originating in the cerebral cortex and de-scending through the brain stem, while the AFP has beenconsidered homologous to the mammalian cortical path-way through the basal ganglia and thalamus.[51] Models

of bird-song motor learning  can be useful in develop-ing models for how humans learn   speech.[53] In somespecies such as Zebra Finches, learning of song is limitedto the first year; they are termed “age-limited” or “close-ended” learners. Other species such as the canaries candevelop new songs even as sexually mature adults; theseare termed “open-ended” learners.[54][55]

Researchers have hypothesized that learned songs allowthe development of more complex songs through culturalinteraction, thus allowing intraspecies dialects that helpbirds to identify kin and to adapt their songs to differentacoustic environments.[56]

5 Neuroanatomy

The acquisition and learning of bird song involves a groupof distinct brain areas that are aligned in two connectingpathways:[51]

•  Anterior forebrain pathway (vocal learning): com-posed of Area X, which is a homologue tomammalian basal ganglia; the lateral part ofthe magnocellular  nucleus of anterior nidopallium

(LMAN), also considered a part of the avian basalganglia; and the dorso-lateral division of the medialthalamus (DLM).

https://en.wikipedia.org/wiki/Magnocellular_cellhttps://en.wikipedia.org/wiki/Magnocellular_cellhttps://en.wikipedia.org/wiki/Vocal_learninghttps://en.wikipedia.org/wiki/Speechhttps://en.wikipedia.org/wiki/Motor_learninghttps://en.wikipedia.org/wiki/Brain_stemhttps://en.wikipedia.org/wiki/Cerebral_cortexhttps://en.wikipedia.org/wiki/Homology_(biology)https://en.wikipedia.org/wiki/Basal_gangliahttps://en.wikipedia.org/wiki/Motor_learninghttps://en.wikipedia.org/wiki/Zebra_Finchhttps://en.wikipedia.org/wiki/Babblinghttps://en.wikipedia.org/wiki/Dialectshttps://en.wikipedia.org/wiki/Cross-fosteringhttps://en.wikipedia.org/wiki/Cross-fosteringhttps://en.wikipedia.org/wiki/Mockingbirdhttps://en.wikipedia.org/wiki/Lyrebird

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4   5 NEUROANATOMY 

HVC

RA

D  L M  

Area X

LMAN

  n   X   l   l  t  s

Song-learning pathway in birds (based on Nottebohm, 2005)

•   Posterior descending pathway (vocal production):composed of HVC (proper name, although some-times referred to as the  high vocal center); therobust nucleus of the arcopallium (RA); and thetracheosyringeal part of the   hypoglossal nucleus(nXIIts).[57][58]

The posterior descending pathway (PDP) is requiredthroughout a bird’s life for normal song production, whilethe anterior forebrain pathway (AFP) is necessary forsong learning in juvenilles and plasticity/maintenance in

adults, but not for adult song production.

[59]

Both neural pathways in the song system begin at thelevel of HVC, which projects information both to the RA(premotor nucleus) and to Area X of the anterior fore-brain. Information in the posterior descending pathway(also referred to as the vocal production or motor path-way) descends from HVC to RA, and then from RA tothe tracheosyringeal part of the hypoglossal nerve (nXI-Its), which then controls muscular contractions of thesyrinx.[51][60]

Information in the anterior forebrain pathway is projectedfrom HVC to Area X (basal ganglia), then from Area X to

the DLM (thalamus), and from DLM to LMAN, whichthen links the vocal learning and vocal production path-ways through connections back to the RA. Some investi-gators have posited a model in which the connection be-tween LMAN and RA carries an instructive signal basedon evaluation of auditory feedback (comparing the bird’sown song to the memorized song template), which adap-tively alters the motor program for song output.[59][61]

The generation of this instructive signal could be facili-tated by auditory neurons in Area X and LMAN that showselectivity for the temporal qualities of the bird’s ownsong (BOS) and its tutor song, providing a platform for

comparing the BOS and the memorized tutor song.

[61][62]

Models regarding the real-time error-correction interac-tions between the AFP and PDP will be considered in

the future. Other current research has begun to explorethe cellular mechanisms underlying HVC control of tem-poral patterns of song structure and RA control of sylla-ble production.[63] Brain structures involved in both path-ways show sexual dimorphism in many bird species, usu-ally causing males and females to sing differently. Some

of the known types of dimorphisms in the brain includethe size of nuclei, the number of neurons present, and thenumber of neurons connecting one nucleus to another.[64]

In the extremely dimorphic Zebra Finches (Taeniopygia guttata), a species in which only males typically sing, thesize of the HVC and RA are approximately three to sixtimes larger in males than in females, and Area X does notappear to be recognizable in females.[65] Research sug-gests that exposure to sex steroids during early develop-ment is partially responsible for these differences in thebrain. Female Zebra Finches treated with estradiol afterhatching followed by testosterone or dihydrotestosterone

(DHT) treatment in adulthood will develop an RA andHVC similar in size to males and will also display male-like singing behavior.[66] Hormone treatment alone doesnot seem to produce female finches with brain structuresor behavior exactly like males. Furthermore, other re-search has shown results that contradict what would beexpected based on our current knowledge of mammaliansexual differentiation. For example, male Zebra Finchescastrated or given sex steroid inhibitors as hatchlings stilldevelop normal masculine singing behavior.[64] This sug-gests that other factors, such as the activation of genes onthe z chromosome, might also play a role in normal malesong development.[67]

Hormones also have activational effects on singing andthe song nuclei in adult birds. In canaries (Serinus ca-naria), females normally sing less often and with lesscomplexity than males. However, when adult females aregiven androgen injections, their singing will increase toan almost male-like frequency.[68] Furthermore, adult fe-males injected with androgens also show an increased sizein the HVC and RA regions.[69] Melatonin is another hor-mone that is also believed to influence song behavior inadults, as many songbirds show melatonin receptors inneurons of the song nuclei.[70] Both the European Star-ling (Sturnus vulgaris ) and House Sparrow (Passer domes-

ticus ) have demonstrated changes in song nuclei corre-lated with differing exposures to darkness and secretionsof melatonin.[71][72] This suggests that melatonin mightplay a role in the seasonal changes of singing behavior insongbirds that live in areas where the amount of daylightvaries significantly throughout the year. Several otherstudies have looked at seasonal changes in the morphol-ogy of brain structures within the song system and havefound that these changes (adult neurogenesis, gene ex-pression) are dictated by photoperiod, hormonal changesand behavior.[73][74]

The gene FOXP2, defects of which affect both speech

production and comprehension of language in humans,becomes highly expressed in Area X during periods of

https://en.wikipedia.org/wiki/FOXP2https://en.wikipedia.org/wiki/Melatoninhttps://en.wikipedia.org/wiki/Dihydrotestosteronehttps://en.wikipedia.org/wiki/Sexual_dimorphismhttps://en.wikipedia.org/wiki/High_vocal_centerhttps://en.wikipedia.org/wiki/Vocal_learninghttps://en.wikipedia.org/wiki/High_vocal_centerhttps://en.wikipedia.org/wiki/Hypoglossal_nervehttps://en.wikipedia.org/wiki/High_vocal_centerhttps://en.wikipedia.org/wiki/High_vocal_centerhttps://en.wikipedia.org/wiki/Hypoglossal_nucleushttps://en.wikipedia.org/wiki/High_vocal_center

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6.1 Mirror neurons and vocal learning   5

vocal plasticity in both juvenile Zebra Finches and adultcanaries.[75]

6 Auditory feedback in birdsong

learning

Early experiments by Thorpe in 1954 showed the impor-tance of a bird being able to hear a tutor’s song. Whenbirds are raised in isolation, away from the influence ofconspecific males, they still sing. While the songthey pro-duce, called “isolate song”, resembles the song of a wildbird, it shows distinctly different characteristics from thewild song and lacks its complexity.[76] The importance ofthe bird being able to hear itself sing in the sensorimotorperiod was later discovered by Konishi. Birds deafenedbefore the song-crystallization period went on to producesongs that were distinctly different from the wild type andisolate song.[77][78] Since the emergence of these findings,investigators have been searching for the neural pathwaysthat facilitate sensory/sensorimotor learning and mediat-ing the matching of the bird’s own song with the memo-rized song template.

Several studies over recent decades have looked at theneural mechanisms underlying birdsong learning by per-forming lesions to relevant brain structures involved inthe production or maintenance of song or by deafeningbirds before and/or after song crystallization. Another re-cent experimental approach was recording the bird’s songand then playing it back while the bird is singing, causingperturbed auditory feedback (the bird hears the superpo-sition of its own song and a fragmented portion of a previ-ous song syllable).[52] After Nordeen & Nordeen[79] madea landmark discovery as they demonstrated that auditoryfeedback was necessary for the maintenance of song inadult birds with crystallized song, Leonardo & Konishi(1999) designed an auditory feedback perturbation pro-tocol in order to explore the role of auditory feedback inadult song maintenance further, to investigate how adultsongs deteriorate after extended exposure to perturbedauditory feedback, and to examine the degree to which

adult birds could recover crystallized song over time af-ter being removed from perturbed feedback exposure.This study offered further support for role of auditoryfeedback in maintaining adult song stability and demon-strated how adult maintenance of crystallized birdsong isdynamic rather than static.

Brainard & Doupe (2000) posit a model in which LMAN(of the anterior forebrain) plays a primary role in errorcorrection, as it detects differences between the song pro-duced by the bird and its memorized song template andthen sends an instructive error signal to structures in thevocal production pathway in order to correct or modify

the motor program for song production. In their study,Brainard & Doupe (2000) showed that while deafeningadult birds led to the loss of song stereotypy due to al-

tered auditory feedback and non-adaptive modificationof the motor program, lesioning LMAN in the anteriorforebrain pathway of adult birds that had been deafenedled to the stabilization of song (LMAN lesions in deaf-ened birds prevented any further deterioration in syllableproduction and song structure).

Currently, there are two competing models that elucidatethe role of LMAN in generating an instructive error signaland projecting it to the motor production pathway:

•   Bird’s Own Song (BOS)-tuned Error CorrectionModel

During singing, the activation of LMAN neu-rons will depend on the match between audi-tory feedback from the song produced by thebird and the stored song template. If this is

true, then the firing rates of LMAN neuronswill be sensitive to changes in auditory feed-back.

•  Efference Copy Model of Error Correction

An efference copy of the motor command forsong production is the basis of the real-timeerror-correction signal. During singing, acti-vation of LMAN neurons will depend on themotor signal used to generate the song, and thelearned prediction of expected auditory feed-

back based on that motor command. Error cor-rection would occur more rapidly in this model.

Leonardo   [80] tested these models directly by record-ing spike rates in single LMAN neurons of adult ZebraFinches during singing in conditions with normal and per-turbed auditory feedback. His results did not supportthe BOS-tuned error correction model, as the firing ratesof LMAN neurons were unaffected by changes in audi-tory feedback and therefore, the error signal generated byLMAN appeared unrelated to auditory feedback. More-over, the results from this study supported the predictions

of the efference copy model, in which LMAN neurons areactivated during singing by the efference copy of the mo-tor signal (and its predictions of expected auditory feed-back), allowing the neurons to be more precisely time-locked to changes in auditory feedback.

6.1 Mirror neurons and vocal learning

A mirror neuron is a neuron that discharges both when anindividual performs an action and when he/she perceivesthat same action being performed by another.[81] Theseneurons were first discovered in macaque monkeys, but

recent research suggests that mirror neuron systems maybe present in other animals including humans.[82]

Mirror neurons have the following characteristics:[81]

https://en.wikipedia.org/wiki/Macaquehttps://en.wikipedia.org/wiki/Neuronhttps://en.wikipedia.org/wiki/Mirror_neuronhttps://en.wikipedia.org/wiki/Efference_copy

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6   7 IDENTIFICATION AND SYSTEMATICS 

•  They are located in the premotor cortex.

•   They exhibit both sensory and motor properties.

•  They are action-specific – mirror neurons are onlyactive when an individual is performing or observing

a certain type of action (e.g., grasping an object).

Because mirror neurons exhibit both sensory and motoractivity, some researchers have suggested that mirror neu-rons may serve to map sensory experience onto mo-tor structures.[83] This has implications for birdsonglearning– many birds rely on auditory feedback to ac-quire and maintain their songs. Mirror neurons may bemediating this comparison of what the bird hears, howit compares to a memorized song template, and what heproduces.

 

primary song primary song

 

primary song other song

primary song

 

other song

Songfreq.(Hz)

Songfreq.(Hz)

Songfreq.(Hz)

a)

b)

c)

neuronactivity

neuronactivity

neuronactivity

time

time

time

Song selectivity in HVCx neurons:  neuron activity in response

to calls heard (green) and calls produced (red).   a.  Neurons firewhen the primary song type is either heard or sung.   b,c.   Neu-

rons 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, JonathanPrather and other researchers at Duke Universityrecorded the activity of single neurons in the  HVCs ofswamp sparrows.[84] They discovered that the neuronsthat project from the HVC to Area X (HVCX neurons)are highly responsive when the bird is hearing a playbackof his own song. These neurons also fire in similar pat-

terns when the bird is singing that same song. Swampsparrows employ 3-5 different song types, and the neuralactivity differs depending on which song is heard or sung.

The HVCX neurons only fire in response to the presen-tation (or singing) of one of the songs, the primary songtype. They are also temporally selective, firing at a pre-cise phase in the song syllable.

Prather, et al. found that during the short period of time

before and after the bird sings, his HVCX neurons be-come insensitive to auditory  input. In other words, thebird becomes “deaf” to his own song. This suggests thatthese neurons are producing a corollary discharge, whichwould allow for direct comparison of motor output andauditory input.[85] This may be the mechanism underly-ing learning via auditory feedback. These findings arealso in line with Leonardo’s (2004) efference copy modelof error correction in birdsong learning and production.

Overall, the HVCX auditory motor neurons in swampsparrows are very similar to the visual motor mirror neu-rons 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 bythe “primary song type”

The function of the mirror neuron system is still unclear.Some scientists speculate that mirror neurons may playa role in understanding the actions of others,  imitation,theory of mind and language acquisition, though there is

currently insufficient neurophysiological evidence in sup-port of these theories.[83] Specifically regarding birds, itis possible that the mirror neuron system serves as a gen-eral mechanism underlying vocal learning, but further re-search is needed. In addition to the implications for songlearning, the mirror neuron system could also play a rolein  territorial behaviors such as song-type matching andcountersinging.[86]

7 Identification and systematics

The specificity of bird calls has been used extensivelyfor species identification. The calls of birds have beendescribed using words or nonsense syllables or linediagrams.[87] Common terms in English include wordssuch as  quack ,  chirp  and  chirrup. These are subject toimagination and vary greatly; a well-known example isthe White-throated Sparrow's song, given in Canada asO sweet Canada Canada Canada   and in New Englandas  Old Sam Peabody Peabody Peabody  (also  Where areyou Frederick Frederick Frederick? ). In addition to non-sense words, grammatically correct phrases have beenconstructed as likenesses of the vocalizations of birds.

For example, the   Barred Owl   produces a motif whichsome bird guides describe as  Who cooks for you? Whocooks for you all?  with the emphasis placed on you.[88]

https://en.wikipedia.org/wiki/Barred_Owlhttps://en.wikipedia.org/wiki/New_Englandhttps://en.wikipedia.org/wiki/Canadahttps://en.wikipedia.org/wiki/White-throated_Sparrowhttps://en.wikipedia.org/wiki/Territorial_behaviorhttps://en.wikipedia.org/wiki/Vocal_learninghttps://en.wikipedia.org/wiki/Neurophysiologyhttps://en.wikipedia.org/wiki/Language_acquisitionhttps://en.wikipedia.org/wiki/Theory_of_mindhttps://en.wikipedia.org/wiki/Imitationhttps://en.wikipedia.org/wiki/Premotorhttps://en.wikipedia.org/wiki/Primateshttps://en.wikipedia.org/wiki/Corollary_dischargehttps://en.wikipedia.org/wiki/Soundhttps://en.wikipedia.org/wiki/Swamp_sparrowhttps://en.wikipedia.org/wiki/High_vocal_centerhttps://en.wikipedia.org/wiki/Motor_neuronhttps://en.wikipedia.org/wiki/Sensory_neuronhttps://en.wikipedia.org/wiki/Premotor_cortex

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Sonogram of the call of a  Laughing Dove.  Recorded in south

India

The use of spectrograms to visualize bird song was firstintroduced by W. H. Thorpe.[89][90] These visual repre-sentations are also called sonograms or sonagrams. Be-ginning in 1983, some field guides for birds use sono-grams to document the calls and songs of birds.[91] Thesonogram is objective, unlike descriptive phrases, butproper interpretation requires experience. Sonograms

can also be roughly converted back into sound.[92][93]

Bird song is an integral part of bird courtship and is apre-zygotic isolation mechanism involved in the processof speciation. Many allopatric sub-species show differ-ences in calls. These differences are sometimes minute,often detectable only in the sonograms. Song differencesin addition to other taxonomic attributes have been usedin the identification of new species.[94] The use of callshas led to proposals for splitting of species complexessuch as those of the Mirafra bushlarks.[95]

8 Recording

The first known recording of birdsong was made in 1889by Ludwig Koch,[96] who went on to become an eminentwildlife recordist and BBC natural history presenter.[96]

Other notable birdsong recordists include  Eric Simms,Chris Watson   and, in France, Jean-Claude Roché,François Charron, Fernand Deroussen.

9 Bird language

The language of the birds has long been a topic for anec-dote and speculation. That calls have meanings thatare interpreted by their listeners has been well demon-strated. Domestic  chickens have distinctive alarm callsfor aerial and ground predators, and they respond to thesealarm calls appropriately.[97][98] However, a language has,in addition to words,  grammar  (that is, structures andrules). Studies to demonstrate the existence of languagehave been difficult due to the range of possible inter-pretations. Research on parrots by Irene Pepperberg isclaimed to demonstrate the innate ability for grammati-

cal structures, including the existence of concepts such asnouns, adjectives and verbs.[99] Studies on starling vocal-izations have also suggested that they may have recursive

structures.[100]

The term  bird language  may also more informally referto patterns in bird vocalizations that communicate infor-mation to other birds or other animals in general.[101]

10 Bird song and music

Some musicologists believe that birdsong has had a largeinfluence on the development of music.[102] Although theextent of this influence is impossible to gauge,[103] it issometimes easy to see some of the specific ways com-posers have integrated birdsong with music.

There seem to be three general ways musicians or com-posers can be affected by birdsong: they can be influ-enced or inspired (consciously or unconsciously) by bird-song, they can include intentional imitations of bird song

in a composition, or they can incorporate recordings ofbirds into their works.

In his book  Why Birds Sing, David Rothernberg claimsthat birds vocalize traditional scales used in human mu-sic, such as the pentatonic scale (e.g., Hermit Thrush) anddiatonic scale (e.g., Wood Thrush), providing evidencethat birdsong not only sounds like music, but is music inthe human sense. This claim has been refuted by Sotorrio(Tone Spectra), who has shown that birds are not select-ing scale tones from a myriad of tonal possibilities, butare filtering out and reinforcing the available set of over-tones from the fundamental tones of their vocal cords.

This requires “far less musical intelligence and deliber-ate appropriation”, and in this regard, he suggests bird-song has something in common with Mongolian throat-singing and jaw-harp music. Sotorrio alsoclaimsthat mu-sicians like Rothernberg are deceived by “a peculiar formof Pareidolia extquotedbl whereby complex tonal infor-mation is reduced to human-scale concepts due to a “fix-ation on music as it is written rather than as it sounds”.Rothernberg’s claims were explored in the BBC docu-mentary Why Birds Sing.

10.1 Compositions that imitate or use

birdsong

One early example of a composition that imitates bird-song is  Janequin's “Le Chant Des Oiseaux”, written inthe 16th century.

Other composers who have quoted birds or haveused birdsong as a compositional springboard includeVivaldi (“Spring” from the Four Seasons ), Biber (SonataRepresentativa),   Beethoven   (Sixth Symphony),   Wagner(Siegfried ) and the jazz musicians Paul Winter (Flyway)and Jeff Silverbush (Grandma Mickey).[104]

The twentieth-century French composer Olivier Messi-aen composed with birdsong extensively. His Catalogued'Oiseaux  is a seven-book set of solo piano pieces based

https://en.wikipedia.org/wiki/Olivier_Messiaenhttps://en.wikipedia.org/wiki/Olivier_Messiaenhttps://en.wikipedia.org/wiki/Paul_Winterhttps://en.wikipedia.org/wiki/Wagnerhttps://en.wikipedia.org/wiki/Beethovenhttps://en.wikipedia.org/wiki/Heinrich_Ignaz_Biberhttps://en.wikipedia.org/wiki/Antonio_Vivaldihttps://en.wikipedia.org/wiki/Janequinhttps://en.wikipedia.org/wiki/Pareidoliahttps://en.wikipedia.org/wiki/Musicologistshttps://en.wikipedia.org/wiki/Starlinghttps://en.wikipedia.org/wiki/Parrothttps://en.wikipedia.org/wiki/Grammarhttps://en.wikipedia.org/wiki/Languagehttps://en.wikipedia.org/wiki/Chickenhttps://en.wikipedia.org/wiki/Language_of_the_birdshttps://en.wikipedia.org/wiki/Chris_Watson_(musician)https://en.wikipedia.org/wiki/Eric_Simms_(ornithologist)https://en.wikipedia.org/wiki/Ludwig_Karl_Kochhttps://en.wikipedia.org/wiki/Mirafrahttps://en.wikipedia.org/wiki/Allopatric_speciationhttps://en.wikipedia.org/wiki/Speciationhttp://-/?-https://en.wikipedia.org/wiki/Spectrogramhttps://en.wikipedia.org/wiki/Media:S-senegalensis.ogghttps://en.wikipedia.org/wiki/Media:S-senegalensis.ogghttps://en.wikipedia.org/wiki/Laughing_Dove

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upon birdsong. His orchestral piece Réveil des Oiseaux  iscomposed almost entirely of birdsong. Many of his othercompositions, including   Quatuor pour la fin du temps ,similarly integrate birdsong.[105]

The Italian composer  Ottorino Respighi, with his   The

Pines of Rome  (1923–1924), may have been the first tocompose a piece of music that calls for pre-recorded bird-song. A few years later, Respighi wrote Gli Uccelli  (“Thebirds”), based on Baroque pieces imitating birds.

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

The American jazz musician Eric Dolphy sometimes lis-tened to birds while he practiced the flute. He claimed tohave incorporated bird song into some of his improvisa-

tional music.

In the 1960s and 1970s, many rock bands included soundeffects in their recordings. Birds were a popular choice.The English band Pink Floyd included bird sound effectsin many of the songs from their 1969 albums Soundtrack 

 from the Film More   and   Ummagumma   (for example,extquotedblGrantchester Meadows  extquotedbl). Sim-ilarly, the English singer  Kate Bush   incorporated birdsound effects into much of the music on her 2005 album,Aerial .

The Music hall artist Ronnie Ronalde has gained notori-ety for his whistling imitations of birds and for integratingbirdsong with human song. His songs “In A MonasteryGarden” and “If I Were A Blackbird” include imitationsof the blackbird, his “signature bird”.

The French composer François-Bernard Mâche has beencredited with the creation of zoomusicology, the study ofthe music of animals. His essay “Musique, mythe, na-ture, ou les Dauphins d'Arion” (1983) includes a studyof “ornitho-musicology”, in which he speaks of “animalmusics” and a longing to connect with nature.

The German DJ, techno music  producer and  naturalistDominik Eulberg   is an avid  bird watcher, and several

tracks by him prominently feature sampled bird soundsand even are titled after his favourite specimens.

The productions of The Jewelled Antler Collective oftenuse field recordings featuring birdsong.

In 2007, the CT Collective issued two free albums de-voted to music made using bird songs (one with humaninteraction, one without). The project was co-ordinatedby looping musician Nick Robinson.

Other recent composers who have used birdsong in theirmusic include   R. Murray Schafer,  Michel Gonneville,Hollis Taylor,  Kyoko Kobayashi,  Rozalie Hirs,   Magnus

Robb,  Stephen Preston   and Emily Doolittle.

[106]

A. J.Mithra, India’s only know zoo-musicologist has com-posed music using natural birds, animals and frog sounds

since 2008.[107]

10.2 Bird song and poetry

Bird song is a popular subject in poetry. Famous poems

inspired by bird song include Percy Bysshe Shelley's “Toa Skylark” (“Hail to thee, blithe Spirit!/Bird thou neverwert”) and Gerard Manley Hopkins' “Sea and Skylark”.Bird songs and their relations to Middle-earth inhabitantsare a common motif in J. R. R. Tolkien's literary work.The Grateful Dead performed a song called “Bird Song”that Jerry Garcia and Robert Hunter wrote; Hunter dedi-cated the lyrics to Janis Joplin.

11 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

12 References

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14 Text and image sources, contributors, and licenses

14.1 Text

•   Bird vocalization Source:  http://en.wikipedia.org/wiki/Bird_vocalization?oldid=627092498  Contributors:  Zundark, William Avery, Cal-trop, Camembert, Shyamal, Julesd, Zimbres, Timwi, Fuzheado, Hyacinth, Samsara, Robbot, Pigsonthewing, Tlogmer, Psychonaut, Seglea,Fuelbottle, JerryFriedman, Cyberied, Andycjp, OverlordQ, DragonflySixtyseven, Sam Hocevar, Arcturus, Zondor, Rich Farmbrough,

Dbachmann, Evice, Brian0918, Dgorsline, Chibimagic, R. S. Shaw, ParticleMan, Jumbuck, Sabine’s Sunbird, Kurt Shaped Box, Ste-monitis, Joriki, Miss Madeline, Rjwilmsi, Nightscream, Dan Guan, Fragglet, Pete.Hurd, Bgwhite, Wavelength, Mushin, RussBot, Spuri-ousQ, Miskatonic, Nicke L, Mkbnett, Daniel Mietchen, Zwobot, Black Falcon, Mike Serfas, Closedmouth, Spondoolicks, Mike Selinker,Wbrameld, Stevouk, Kungfuadam, Sardanaphalus, SmackBot, CSZero, RedSpruce, Commander Keane bot, Mcld, Williamemersonwood,Ajsh, Chris the speller, Gil mo, Gracenotes, George Ho, Snowmanradio, Stevenmitchell, Richard001, Jidanni, JHunterJ, Luokehao, Learn-ingKnight, Wentu, Courcelles, Rouseaubade, ChrisCork, CmdrObot, Drinibot, ShelfSkewed, WeggeBot, Neelix, Veggie26, Jlperez11,Robertinventor, Thijs!bot, Barticus88, MichaelMaggs, Widefox, Jhsounds, Sreejithk2000, Natureguy1980, Mertseger, Maias, Tintin Mon-treal, Tomhannen, Think outside the box, Steven Walling, Rich257, Catgut, Richarddr, Human785, B9 hummingbird hovering, STBot,Keith D, R'n'B, CommonsDelinker, S.dedalus, J.delanoy, LordAnubisBOT, LittleHow, STBotD, Pnoble805, Squids and Chips, VolkovBot,Atzsch, Upupa epops, Cheesypea, 4444hhhh, Andreas Carter, Hyper oxtane, Hyper flyin', The high and mighty, Bunnyfinch, MeegsC,Mangostar, Jsfouche, Nuttycoconut, Callidior, DivineBurner, ClueBot, RitigalaJayasena, Mild Bill Hiccup, Cdh8, Jusdafax, PixelBot, SunCreator, Agutwin, Srw48, Prajnamarie, Ornisong, Dekisugi, SchreiberBike, A plague of rainbows, 3ICE, BOTarate, Grantus4504, Ost316,Addbot, DOI bot, Andrewtayeri, A Knight Who Says Ni, Bootboy41, Ajmithra, LinkFA-Bot, Numbo3-bot, Cesiumfrog, Zorrobot, Jarble,Luckas-bot, Yobot, Bunnyhop11, AnomieBOT, Rjanag, Galoubet, Flewis, Materialscientist, Citation bot, MidnightBlueMan, 4twenty42o,RibotBOT, Ckazilek, MichealH, Citation bot 1, Pinethicket, RedBot, Trappist the monk, Innotata, Clm82, Updatehelper, TjBot, An-

drostachys, John of Reading, Immunize, Joaosac, H3llBot, MaximeA, ةين ريد , ClueBot NG, Hypergraph, Wimpus, Helpful PixieBot, Gcc111, Voyal, BG19bot, Desmosa, Gpresearch, Weblars, MinonaLewis, Rynsaha, Glb58, BattyBot, Mediran, Dexbot, , Tianyiع دCai, Turn on a Dime, Ccevo2012, Thenatureexplorers, Anrnusna, Juhuyuta, Coyote Cósmico v Carlos Castañeda, Monkbot, Sometree,SyntaxSC and Anonymous: 112

14.2 Images

•   File:Bird_song_development_timeline.svg   Source:    http://upload.wikimedia.org/wikipedia/commons/f/f1/Bird_song_development_timeline.svg License:  CC-BY-3.0  Contributors:  Own work Original artist:   Hypergraph

•   File:Birdbrain.svg Source:   http://upload.wikimedia.org/wikipedia/commons/c/c8/Birdbrain.svg License:  CC-BY-2.5   Contributors:   ownwork based on  http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0030164   Original artist:    L.Shyamal based on Nottebohm F

•   File:Commons-logo.svg Source:  http://upload.wikimedia.org/wikipedia/en/4/4a/Commons-logo.svg License:  ?   Contributors:  ?  Original artist:  ?

•   File:Gnome-mime-sound-openclipart.svg   Source:    http://upload.wikimedia.org/wikipedia/commons/8/87/Gnome-mime-sound-openclipart.svg  License:  Public domain   Contributors:  Own work. Based on  File:Gnome-mime-audio-openclipart.svg, which is public domain. Original artist:  User:Eubulides

•  File:Little_brown_dove.png   Source:   http://upload.wikimedia.org/wikipedia/commons/8/8b/Little_brown_dove.png License:   CC-BY-2.5  Contributors:  Own work Original artist:  Shyamal

•   File:Loudspeaker.svg Source:   http://upload.wikimedia.org/wikipedia/commons/8/8a/Loudspeaker.svg License:   Public domain Contrib-utors:  ?  Original artist:  ?

•   File:PipraWing.jpg Source:  http://upload.wikimedia.org/wikipedia/commons/5/54/PipraWing.jpg License:  Public domain  Contributors: The Descent of Man. Volume 2 Original artist:  Charles Darwin, using the work of P L Sclater

•   File:Pixinguinha-_O_Urubu_e_o_Gaviao_1930.ogg Source:  ht tp://upload.wikimedia.org/wikipedia/commons/9/97/Pixinguinha-_O_Urubu_e_o_Gaviao_1930.ogg License:   Public domain  Contributors:  Public domain Original artist:  Pixinguinha, publicado pela gravadoraVictor em 1930

•   File:Solsort.jpg  Source:   http://upload.wikimedia.org/wikipedia/commons/b/ba/Solsort.jpg License:   CC-BY-SA-3.0   Contributors:   This

image was copied from da:Billede:Solsort.jpg on the Danske Wikipedia.  Original artist:  Malene Thyssen (User:Malene)•   File:Song_selective_HVCx_neurons.svg   Source:    http://upload.wikimedia.org/wikipedia/commons/2/29/Song_selective_HVCx_

neurons.svg License:  CC-BY-SA-3.0  Contributors:  Own work Original artist:  Clm82

•   File:Voice_of_Zonotrichia_albicollis.ogg   Source:    http://upload.wikimedia.org/wikipedia/commons/f/f6/Voice_of_Zonotrichia_albicollis.ogg License:  CC-BY-3.0  Contributors:  own recording in en:Algonquin Provincial Park  Original artist:  Liesel

•   File:Wiki_letter_w_cropped.svg   Source:    http://upload.wikimedia.org/wikipedia/commons/1/1c/Wiki_letter_w_cropped.svg   License: CC-BY-SA-3.0  Contributors: 

•  Wiki_letter_w.svg Original artist:  Wiki_letter_w.svg: Jarkko Piiroinen

14.3 Content license

•   Creative Commons Attribution-Share Alike 3.0

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