Song Control System PDF

The Song Control System Chapter 12 Communication Learned Vocalizations  Songbirds Most birds make noises (calls), but songbirds have a repertoire of up to 20 or so distinct vocal noises they use to compose distinct songs Order = Passeriformes Suborder = Passeri or Oscines...

The Song Control System Chapter 12 Communication Learned Vocalizations  Songbirds Most birds make noises (calls), but songbirds have a repertoire of up to 20 or so distinct vocal noises they use to compose distinct songs Order = Passeriformes Suborder = Passeri or Oscines Approximately 4000 species sing true songs about ½ of all bird species Song of Common Yellowthroat Songs are learned Birds are born being able to make noise, but the song is a learned sensorimotor skill Sonogram Frequency represented on Y axis Amplitude represented by intensity of marking Time represented on X axis Sonogram Birds learn songs, but there is a lot of variation in song acquisition across species Zebra finch (Taeniopygia guttata) Learn one song motif in the few weeks to a month or so after hatching and produce song consisting of repetitions of that motif without much variation throughout adulthood European starling (Sturnus vulgaris) Learn song in the sensitive period for song learning, but retain a high degree of vocal plasticity in learning and production throughout life, paralleling the open- endedness of vocal learning in humans Nowicki, Stephen and Peter R. Marler. “How Do Birds Sing ? Author ( s ) : Stephen Nowicki and.” (2008).  Zebra finch (Taeniopygia castanotis; previously T. guttata) Model Organism for studying bird song Found in Australia Also a common pet in USA Live for about 5 years Breeding occurs after periods of heavy rainfall Any time of year Sexually dimorphic Only males sing Sons learn from fathers Zebra Finch Sonogram Song Notes and Motif  Sensory phase (sensory learning phase) Young bird listens to tutor (usually father) and memorizes song Happens during a very short period early in life Memorized song (template) is highly stable Two phases of Sensorimotor phase (motor phase) song learning Young birds sing variations of songs (subsong), listening to their own song and trying to match it to the memorized template Song at end of this phase termed plastic song (resembles learned song, still variable) Both phases are dependent on auditory input Hearing the song from the “tutor” and then hearing own song  Once bird reaches sexual maturation, song is crystallized Crystallized Bird now sings song very similar to tutor Song Once crystalized, song is largely no longer dependent on auditory feedback  Territory defense Why sing? Song = deterrent WHITE THROATED SPARROW  Mate attraction Why sing? Numerous laboratory choice studies, field choice studies, and observational field studies Why sing? Might be used more generally for individual recognition Mates within monogamous pairs Parents or siblings Territorial neighbors How Do Birds Sing? Bird Brain Pallium – Cerebral Cortex Bird brains small compared to humans, but body/size ratio still relatively high Same basic structures shared Major difference is mammals have layered cortex Cell bodies in cortex (green) and axons (white matter) below in mammals Jarvis et al. 2005  Brain Nucleus A collection of neurons that are distinct in anatomy and are generally involved in same function  Pinaud, Raphael & Terleph, Thomas & Wynne, Ryan & The Song Control System Map Tremere, Liisa. (2008). Neural Correlates of Auditory Processing, Learning and Memory Formation in Songbirds. Progress of Theoretical Physics Supplement - PROG THEOR PHYS SUPPL. 173. 270-282.  HVC = High Vocal Center RA = Robust nucleus of the Arcopallium nXII = nucleus of the 12th cranial nerve LMAN = lateral magnocellular nucleus of the anterior nidopallium Anterior Forebrain DLM = dorsolateral Pathway thalamic nucleus Cortical-basal Song Motor Pathway Ganglion Pathway Song Control Nuclei Typically Larger in Males Early sex differences in zebra finch brains Neural Recordings in Awake Zebra Finches  HVC = High Vocal Center RA = Robust nucleus of the Arcopallium nXII = nucleus of the 12th cranial nerve LMAN = lateral magnocellular nucleus of the anterior nidopallium Anterior Forebrain DLM = dorsolateral Pathway thalamic nucleus Cortical-basal Song Motor Pathway Ganglion Pathway  HVC Neural Activity is Very Precise in Song Motor Pathway Singing-Related Activity of Identified HVC Neurons in the Zebra Finch Alexay A. Kozhevnikov and Michale S. Fee Journal of Neurophysiology 2007 97:6, 4271-4283 RA Neuronal Activity in Song Motor Pathway Ensemble Coding of Vocal Control in Birdsong. Anthony Leonardo, Michale S. Fee. Journal of Neuroscience 19 January 2005, 25 (3) 652-661; DOI: 10.1523/JNEUROSCI.3036-04.2005  HVC = High Vocal Center RA = Robust nucleus of the Arcopallium nXII = nucleus of the 12th cranial nerve LMAN = lateral magnocellular nucleus of the anterior nidopallium Anterior Forebrain DLM = dorsolateral Pathway thalamic nucleus Cortical-basal Song Motor Pathway Ganglion Pathway Area X Lesion has little effect on adult song production A comparative study of the behavioral deficits following lesions of various parts of the zebra finch song system: implications for vocal learning. C Scharff, F Nottebohm. Journal of Neuroscience 1 September 1991, 11 (9) 2896-2913 Area X Lesions between 30 and 40 days post-hatch disrupt song template A comparative study of the behavioral deficits following lesions of various parts of the zebra finch song system: implications for vocal learning. C Scharff, F Nottebohm. Journal of Neuroscience 1 September 1991, 11 (9) 2896-2913 LMAN Lesion Leads to Song Variability in Note Morphologies and Ordering A comparative study of the behavioral deficits following lesions of various parts of the zebra finch song system: implications for vocal learning. C Scharff, F Nottebohm. Journal of Neuroscience 1 September 1991, 11 (9) 2896-2913 Lesions to Song Nuclei Impact Song Production Lesions to HVC and RA disrupt song production, needed for song Lesions to LMAN lead to song variability Lesions to Area X show function in template of song Gregory F. Ball, Jacques Balthazart. Sex differences and similarities in the neural circuit regulating song and other reproductive behaviors in songbirds. Neuroscience & Biobehavioral Reviews. Volume 118. 2020. Pages 258-269  Anterior Song Forebrain Motor Pathway Pathway Learning Pathway Singing Pathway HVC HVC (pre-motor area) LMAN RA (motor cortex) Area X nXII (innervate muscles in DLM syrinx) RA Song Control System Uses Multiple Neurotransmitters Farries MA, Perkel DJ. A telencephalic nucleus essential for song learning contains neurons with physiological characteristics of both striatum and globus pallidus. J Neurosci. 2002 May 1;22(9):3776-87.  Pinaud, Raphael & Terleph, Thomas & Wynne, Ryan & The Song Control System Map Tremere, Liisa. (2008). Neural Correlates of Auditory Processing, Learning and Memory Formation in Songbirds. Progress of Theoretical Physics Supplement - PROG THEOR PHYS SUPPL. 173. 270-282.