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Birds’ song variability is
modulated by LMAN
in anterior forebrain pathway
Hugo Trad & Stephen Larroque
19 Nov 2015
Learning to sing
• Birds’ singing is a complex, learnt task
• Uses cortical-basal ganglia circuit
• Different song characteristics between
directed (couple) and undirected (single)
• Song variability : how and why ?
(how to demonstrate direct contribution of AFP to adult
birdsong?)
2
(Kao et al, 2005)
AFP Pathway
3
(Kao et al, 2005)
Motor
Pathway
Basal ganglia-forebrain circuit
(anterior forebrain pathway)
What we know
• AFP activity correlated with singing
• LMAN lesion → no change in ongoing song
• HVC+RA (motor pathway) stimulation → vocalizations
• LMAN stimulation → no vocalization elicited and no gross
change on song
=> LMAN not required structure for song production
=> What is its purpose then ? Hypothesis :
AFP activity modulates vocal control via LMAN activity
patterns ?
4
(Kao et al, 2005)
Methodology: manipulating LMAN activity
5
(Kao et al, 2005)
Microstimulation of LMAN during precisely targeted parts
of a song using real-time template-matching algorithm
Results: pitch+loudness modulated by LMAN
6
(Kao et al, 2005)
(sd ; P < 0.0001) (sem ; P < 0.0001)
Frequency shift Amplitude shift
Using fixed current stimulation
Results: LMAN is functionally compartmentalized
7
(Kao et al, 2005)
And high current produce song suspension, like HVC stimulation !
Methodology: social behavioral analysis
8
(Kao et al, 2005)
Spectrogram + neural activity recordings when presented
with a female
Undirected singing Directed singing
Methodology: social behavioral analysis
9
(Kao et al, 2005)
Spectrogram + neural activity recordings when presented
with a female
Undirected singing Directed singing
Methodology: social behavioral analysis
10
(Kao et al, 2005)
Spectrogram + neural activity recordings when presented
with a female
Undirected singing Directed singing
After <= 2 min, hide female + play undirected songs
Results: Context modulates LMAN activity
11
(Kao et al, 2005)
Pitch mean is not changed (needs confirmation...), only variability !
(mean
P = 0.54)
(variance
P < 0.0001)
Correlation of LMAN activity and song variability
12
(Kao et al, 2005)
=> LMAN necessary for context-dependent variability
(confirmed by stereotaxic lesion)
(R² = 0.83 ; P < 0.05) (s.e.m. ; pre-P < 0.0001 ; post-P = 0.167)
Discussion
• What pathway(s) for context modulation ? (what modulates
LMAN difference in activity between directed and undirected
singing? Dopamine ?)
• Exploration-exploitation dilemma : what is the exploration
probability ? And how is this value chosen ? Is it dynamic ?
• How are those new explored syllabes evaluated ?
• Cerebellar implication ? (to bias variability towards the
correct pitch in juvenile learning ?)
• Different sample sizes for the two conditions (class skewing)
• Need extended theory about amplitude modulation (here the
article is mainly studying frequency!) 13
Take home message
• LMAN induces acute changes in individual song
elements (syllabes) frequency (pitch) and amplitude
(loudness), without altering order nor structure of
subsequent syllabes
• Noise and variability can be used advantageously to
explore new unknown solutions, but can become
pathological (Parkison?)
• Exploration behavior promoted in undirected singing
(« training mode »), while exploitation in directed
• Natural organisms exhibit a solution to the exploration-
exploitation dilemma close to the epsilon (noise) trick
→ possible predictive model ? 14
– Contributions of an avian basal ganglia–forebrain circuit to real-time
modulation of song, Kao, Mimi H., Allison J. Doupe, and Michael S.
Brainard, 2005 Nature 433.7026 (2005): 638-643.
– Behavioral and Neural Signatures of Readiness to Initiate a Learned
Motor Sequence, Rajan R. & Doupe A., 2013, Current Biology, 23 (1) 87-93.
– Interruption of a basal ganglia–forebrain circuit prevents plasticity of
learned vocalizations, Brainard, M. S., & Doupe, A. J., 2000, Nature,
404(6779), 762-766.
– Social context modulates singing-related neural activity in the songbird
forebrain, Hessler, N. A., & Doupe, A. J.,1999, Nature neuroscience, 2(3),
209-211.
References
slideshare.net/LRQ3000
Thank you!
slideshare.net/LRQ3000
Bonus Slides
THE END

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Birdsong variability is modulated by LMAN in anterior forebrain pathway

  • 1. Birds’ song variability is modulated by LMAN in anterior forebrain pathway Hugo Trad & Stephen Larroque 19 Nov 2015
  • 2. Learning to sing • Birds’ singing is a complex, learnt task • Uses cortical-basal ganglia circuit • Different song characteristics between directed (couple) and undirected (single) • Song variability : how and why ? (how to demonstrate direct contribution of AFP to adult birdsong?) 2 (Kao et al, 2005)
  • 3. AFP Pathway 3 (Kao et al, 2005) Motor Pathway Basal ganglia-forebrain circuit (anterior forebrain pathway)
  • 4. What we know • AFP activity correlated with singing • LMAN lesion → no change in ongoing song • HVC+RA (motor pathway) stimulation → vocalizations • LMAN stimulation → no vocalization elicited and no gross change on song => LMAN not required structure for song production => What is its purpose then ? Hypothesis : AFP activity modulates vocal control via LMAN activity patterns ? 4 (Kao et al, 2005)
  • 5. Methodology: manipulating LMAN activity 5 (Kao et al, 2005) Microstimulation of LMAN during precisely targeted parts of a song using real-time template-matching algorithm
  • 6. Results: pitch+loudness modulated by LMAN 6 (Kao et al, 2005) (sd ; P < 0.0001) (sem ; P < 0.0001) Frequency shift Amplitude shift Using fixed current stimulation
  • 7. Results: LMAN is functionally compartmentalized 7 (Kao et al, 2005) And high current produce song suspension, like HVC stimulation !
  • 8. Methodology: social behavioral analysis 8 (Kao et al, 2005) Spectrogram + neural activity recordings when presented with a female Undirected singing Directed singing
  • 9. Methodology: social behavioral analysis 9 (Kao et al, 2005) Spectrogram + neural activity recordings when presented with a female Undirected singing Directed singing
  • 10. Methodology: social behavioral analysis 10 (Kao et al, 2005) Spectrogram + neural activity recordings when presented with a female Undirected singing Directed singing After <= 2 min, hide female + play undirected songs
  • 11. Results: Context modulates LMAN activity 11 (Kao et al, 2005) Pitch mean is not changed (needs confirmation...), only variability ! (mean P = 0.54) (variance P < 0.0001)
  • 12. Correlation of LMAN activity and song variability 12 (Kao et al, 2005) => LMAN necessary for context-dependent variability (confirmed by stereotaxic lesion) (R² = 0.83 ; P < 0.05) (s.e.m. ; pre-P < 0.0001 ; post-P = 0.167)
  • 13. Discussion • What pathway(s) for context modulation ? (what modulates LMAN difference in activity between directed and undirected singing? Dopamine ?) • Exploration-exploitation dilemma : what is the exploration probability ? And how is this value chosen ? Is it dynamic ? • How are those new explored syllabes evaluated ? • Cerebellar implication ? (to bias variability towards the correct pitch in juvenile learning ?) • Different sample sizes for the two conditions (class skewing) • Need extended theory about amplitude modulation (here the article is mainly studying frequency!) 13
  • 14. Take home message • LMAN induces acute changes in individual song elements (syllabes) frequency (pitch) and amplitude (loudness), without altering order nor structure of subsequent syllabes • Noise and variability can be used advantageously to explore new unknown solutions, but can become pathological (Parkison?) • Exploration behavior promoted in undirected singing (« training mode »), while exploitation in directed • Natural organisms exhibit a solution to the exploration- exploitation dilemma close to the epsilon (noise) trick → possible predictive model ? 14
  • 15. – Contributions of an avian basal ganglia–forebrain circuit to real-time modulation of song, Kao, Mimi H., Allison J. Doupe, and Michael S. Brainard, 2005 Nature 433.7026 (2005): 638-643. – Behavioral and Neural Signatures of Readiness to Initiate a Learned Motor Sequence, Rajan R. & Doupe A., 2013, Current Biology, 23 (1) 87-93. – Interruption of a basal ganglia–forebrain circuit prevents plasticity of learned vocalizations, Brainard, M. S., & Doupe, A. J., 2000, Nature, 404(6779), 762-766. – Social context modulates singing-related neural activity in the songbird forebrain, Hessler, N. A., & Doupe, A. J.,1999, Nature neuroscience, 2(3), 209-211. References slideshare.net/LRQ3000