Song is produced via vocal ‘membranes’ (labium), filtering of the vocal tract

1. Song is produced via vocal ‘membranes’ (labium), filtering of the vocal tract • (i.e., formants), and changes in the vocal apparatus; whereas mini-breaths are • an important component of the temporal pattern • Songbirds have dual voicedness, with independent nerve innervation • This feature is used for acoustic effect - add FM or syllable jumping; and these • effects may (likely) have an adaptive function – such as mate choice • Birds may also hit up against other performance constraints – rate of breathing; • bandwidth vrs trill rate. Singing at the physical limit may likewise have an adaptive • function • Performance constraints may also appear in non-vocal acoustic communication, • e.g., hummingbird’s dive @ 60mph, manakin’s wing-shrike

2. Order: Passeriformes (Passerines) Suborder: Tyranni (Suboscines) - Simple songs, simple syrinx musculature, do not learn songs; do not even need to hear

3. Eastern Phoebe

4. Eastern Phoebe

5. Order: Passeriformes Suborder: Passeri (Oscines) • Birds develop vocal patterns (song) guided by • information from sounds heard early in life, memorized, • and rehearsed many times • Very rare (cetaceans, several bats) • Opportunity to understand the pulls of Nature and • Nurture - Complex songs, syrinx musculature, learn songs; require a tutor

6. Learning can be adaptive in a changing world; However, how do you choose which of many stimuli to memorize? Auditory Template Model • Auditory Phase of memorization • Pattern selectivity for conspecific song even w/o prior experience • Motor phase of development

7. Pattern selectivity for conspecific song even w/o prior experience • learn only spp specific notes • regardless of syntax and tempo SWAMP • learn other spp if played in • song sparrow-like tempo SONG WHITE- CROWNED

8. “Learn what comes after the WCSP whistle” S_CD2_23 S_CD2_24_mod S_CD2_25 Belding’s Ground Squirrel Pupil Song

9. Here is the same with young bird tutored on the strawberry finch

10. What if there is NO tutor? Kroods_16 WHITE- CROWNED

11. Like humans, in birds there is an age-related pattern in learning – The first year is usually the most important – Sensitive phase # songs exposed to # songs learned MARSH WREN

12. There is both flexibility within a spp and • Considerable variation across spp • The timing provides an additional mechanism to avoid learning the wrong song - the • window coordinated to match the time adults of your own spp are actively singing • open-ended learners • close-ended learners

13. Social Factors: • Live tutors • Isolated birds with playback • Live = Iso/playback SWAMP/SONG Sparrows • but in WHITE-CROWNS live tutor is necessary after 50 days • Live > Iso/playback Starling/Canary • Live > Iso/playback = 0 Tree creepers • Nightingales actually do better learning with synchronize strobe light WHY the Variation ? Repertoire size or closed vs. open-ended learners

14. Sensorimotor phase of development • several weeks after memorization • 3 part process: subsong, plastic song, crystallized full song S_CD1_21ng S_CD1_22

15. Subsong • never the same moment to moment; great variability • no recognized spp specific elements • probably functions to train the vocal apparatus Plastic song • shift to more stereotyped vocalizations • first emergence of acquired imitations emerge • and recur in similar form

16. There is both flexibility within a spp and • Considerable variation across spp • The timing provides an additional mechanism to avoid learning the wrong song - the • window coordinated to match the time adults of your own spp are actively singing

17. Crystallized song • rapid progression from plastic to mature singing • apparently hastened by testosterone • vocalizations get ‘frozen’ • temporally segmented performance • Cull vocalizations (8% in nightingales to 80% in sparrows) …maybe neighborhood-based SWAMP SPARROW

18. AUDITORY TEMPLATE MODEL Crude template Day length increases II Own species song heard I Template matched to song heard Gonad size increases Requires Tutor MEMORIZATION PHASE Exact template This is a (brief) window of crystallization Testosterone III Requires hearing yourself Hears own song Song output Song matched to template Forgetting or culling MOTOR PHASE

19. Brown-headed cowbirds Brood parasites HOW DO THEY LEARN THEIR SONGS?

20. Action-based Learning • Eastern and Southern sub-spp • when transplanted chicks raised by the ‘other’ sub-spp they learn that sub-spp songs • males respond to female wing-flicks by repeating songs more when • females respond this way

22. Indirect pathway between HVC and RA – recursive loop Posterior Vocal pathway (production) Anterior Vocal pathway (learning) HVA=Higher vocal center RA=Robust nucleus of the Arcopallium

23. Many of the nuclei have neurons that recognize specific sounds in complex song

24. This is at the discretion of the researcher

28. Vocal Learning vs. Auditory learning Form memories of sounds you hear and associations with your external environment • Psittaciformes • Trochilidae • Oscine songbirds • Cetaceans • a few bats • Humans • Almost all land vertebrates • and many aquatic ones