• Keine Ergebnisse gefunden

In conclusion, in this thesis, models for associative learning which may serve to asso-ciate meaning with sound were introduced as well as an conceptually complete model for vocal learning composed of a novel architecture and a learning mechanism that not only serves to form inverse models, but is also able to imprint spiking sequences onto recurrent networks, which may underlie the formation of sensory memories.

Throughout this thesis, all networks were numerically integrated using a simple Euler integration scheme. The simulations were written in Python and used a step size of 0.5ms for all leaky integrate-and-fire neurons, a step size of 0.01ms for the conductance-based integrate-and-fire neurons and a step size of 0.005msfor the Hodgkin-Huxley type neurons.

The values of neurons parameters can be found in the respective chapters.

Sascha Helduser, Maren Westkott, Klaus Pawelzik, and Onur G¨unt¨urk¨un. The putative pigeon homologue to song bird lman does not modulate behavioral variability. Behavioural brain research, 263:144148, 2014.

Christian Albers, Maren Westkott, and Klaus Pawelzik. Perfect associative learning with spike-timing-dependent plasticity. In C. J. C.Burges, L. Bottou, M. Welling, Z. Ghahra-mani, and K. Q. Weinberger, editors, Advances in Neural Information Processing Systems 26, pages 17091717. Curran Associates, Inc., 2013.

Christian Albers, Maren Westkott, and Klaus Pawelzik. Learning of Precise Spike Times with Homeostatic Membrane Potential Dependent Synaptic Plasticity. PLoS ONE 11(2):

e0148948. doi:10.1371/journal.pone.0148948, 2016.

Maren Westkott and Klaus Pawelzik. An Integrative Account of Acoustic Sequence Imita-tion Learning in the Songbird. Frontiers in Computational Neuroscience 2015under review.

I have a lot of people to thank without whom the completion of this work would have been impossible. First and foremost, I have to thank my supervisor Klaus Pawelzik for all the creative input and all the new ideas, as well as for the fruitful discussions. I would like to thank Agnes Janßen for the relentless administrative support, which makes life in the group so much easier, as well as for the enthusiasm, good mood and cookies she provides for the group. For great technical support, I would like to thank David Rothermund, who is always available for the stupidest of questions, which he answers with remarkable calmness. I would like to thank all members of the group for providing a friendly and stimulating work environment. In particular, I am grateful to Christian Albers for the work we did together, for long fruitful discussions and for advice. I would like to thank my proof readers Christian Albers, Nergis T¨omen, Federica Capparelli, Daniel Harnack and last but not least Axel Grzymisch, who greatly contributed to making this work more readable, understandable and overall better.

Finally, I would like to thank friends and family for the ongoing support over the course of the years. In particular, I am grateful to my partner M.L. for the ongoing support, feeding and bribing throughout the writing process.

[ABS90] A Artola, S Br¨ocher, and W Singer. Different voltage-dependent thresholds for inducing long-term depression and long-term potentiation in slices of rat visual cortex. Nature, 347(6288):69–72, 1990.

[AK90] LF Abbott and Thomas B Kepler. Model neurons: From hodgkin-huxley to hopfield. In Statistical mechanics of neural networks, pages 5–18. Springer, 1990.

[AL01] D Attwell and S B Laughlin. An energy budget for signaling in the grey matter of the brain.Journal of Cerebral Blood Flow & Metabolism, 21(10):1133–1145, 2001.

[APMM13] Ana Amador, Yonatan Sanz Perl, Gabriel B Mindlin, and Daniel Margoliash.

Elemental gesture dynamics are encoded by song premotor cortical neurons.

Nature, 495(7439):59–64, 2013.

[AWP13] Christian Albers, Maren Westkott, and Klaus Pawelzik. Perfect associative learning with spike-timing-dependent plasticity. In C. J. C. Burges, L. Bottou, M. Welling, Z. Ghahramani, and K. Q. Weinberger, editors,Advances in Neu-ral Information Processing Systems 26, pages 1709–1717. Curran Associates, Inc., 2013.

[AWP16] Christian Albers, Maren Westkott, and Klaus Pawelzik. Learning of precise spike times with membrane potential dependent synaptic plasticity. PLoS ONE, (11):(2): e0148948, 2016.

[BD02] Michael S Brainard and Allison J Doupe. What songbirds teach us about learning. 2002.

[BD13] Michael S Brainard and Allison J Doupe. Translating birdsong: songbirds as a model for basic and applied medical research. Annual review of neuroscience, 36:489, 2013.

[BKLP02] Sander M Bohte, Joost N Kok, and Han La Poutre. Error-backpropagation in temporally encoded networks of spiking neurons. Neurocomputing, 48:17–37, 2002.

[BMA84] Sarah W Bottjer, Elizabeth A Miesner, and Arthur P Arnold. Forebrain lesions disrupt development but not maintenance of song in passerine birds.

Science, 224(4651):901–903, 1984.

[BSP13] Johanni Brea, Walter Senn, and Jean-Pascal Pfister. Matching recall and storage in sequence learning with spiking neural networks. The Journal of Neuroscience, 33(23):9565–9575, 2013.

[CBVG10] Claudia Clopath, Lars B¨using, Eleni Vasilaki, and Wulfram Gerstner. Con-nectivity reflects coding: a model of voltage-based STDP with homeostasis.

Nature Neuroscience, 13(3):344–352, 2010.

[CD08] Natalia Caporale and Yang Dan. Spike timing-dependent plasticity: A heb-bian learning rule. Annu Rev Neurosci, 31:25–46, 2008.

[DA01] Peter Dayan and Larry F Abbott. Theoretical Neuroscience. The MIT Press, 2001.

[DK91] Allison J Doupe and Masakazu Konishi. Song-selective auditory circuits in the vocal control system of the zebra finch. Proceedings of the National Academy of Sciences, 88(24):11339–11343, 1991.

[DK99] Allison J Doupe and Patricia K Kuhl. Birdsong and human speech: Common themes and mechanisms. Annu. Rev. Neurosci, 22:567–631, 1999.

[DLH10] Prashanth D’Souza, Shih-Chii Liu, and Richard H R Hahnloser. Percep-tron learning rule derived from spike-frequency adaptation and spike-time-dependent plasticity. Proceedings of the National Academy of Sciences, 107(10):4722–4727, 2010.

[DM00] Amish S Dave and Daniel Margoliash. Song replay during sleep and com-putational rules for sensorimotor vocal learning. Science, 290(5492):812–816, 2000.

[DO87] Sigurd Diederich and Manfred Opper. Learning of correlated patterns in spin-glass networks by local learning rules. Physical review letters, 58(9):949, 1987.

[DP04] Yang Dan and Mu-Ming Poo. Spike timing-dependent plasticity of neural circuits. Neuron, 44(1):23–30, 2004.

[DP06] Yang Dan and Mu-Ming Poo. Spike timing-dependent plasticity: from synapse to perception. Physiological reviews, 86(3):1033–1048, 2006.

[DPFF+92] Giuseppe Di Pellegrino, Luciano Fadiga, Leonardo Fogassi, Vittorio Gallese, and Giacomo Rizzolatti. Understanding motor events: a neurophysiological study. Experimental brain research, 91(1):176–180, 1992.

[FD02] Robert C Froemke and Yang Dan. Spike-timing-dependent synaptic modifi-cation induced by natural spike trains. Nature, 416(6879):433–438, 2002.

[FDV09] Elodie Fino, Jean Michel Deniau, and Laurent Venance. Brief subthreshold events can act as Hebbian signals for long-term plasticity. PLoS One, 4(8), 2009.

[Fel00] Daniel E Feldman. Timing-based ltp and ltd at vertical inputs to layer ii/iii pyramidal cells in rat barrel cortex. Neuron, 27(1):45–56, 2000.

[FFS07] Ila R Fiete, Michale S Fee, and H Sebastian Seung. Model of birdsong learn-ing based on gradient estimation by dynamic perturbation of neural conduc-tances. Journal of neurophysiology, 98(4):2038–2057, 2007.

[Flo12a] Rzvan V. Florian. The chronotron: A neuron that learns to fire temporally precise spike patterns. PLoS One, 7(8), 2012.

[Flo12b] Rzvan V. Florian. The chronotron: A neuron that learns to fire temporally precise spike patterns. PLoS ONE, (7), 2012.

[FPD05] Robert C Froemke, Mu-Ming Poo, and Yang Dan. Spike-timing-dependent synaptic plasticity depends on dendritic location. Nature, 434(7030):221–225, 2005.

[FS06] Ila R Fiete and H Sebastian Seung. Gradient learning in spiking neural networks by dynamic perturbation of conductances. Physical review letters, 97(4):048104, 2006.

[FS10] Michale S Fee and Constance Scharff. The songbird as a model for the gener-ation and learning of complex sequential behaviors. ILAR journal, 51(4):362–

377, 2010.

[GK02] Wulfram Gerstner and Werner M Kistler. Spiking Neuron Models: Single Neurons, Populations, Plasticity. Cambridge university press, 2002.

[GKGH14] Nicolas Giret, Joergen Kornfeld, Surya Ganguli, and Richard HR Hahnloser.

Evidence for a causal inverse model in an avian cortico-basal ganglia circuit.

Proceedings of the National Academy of Sciences, 111(16):6063–6068, 2014.

[GM08] Tim Gollisch and Markus Meister. Rapid neural coding in the retina with relative spike latencies. Science, 319(5866):1108–1111, 2008.

[GS06] Robert G¨utig and Haim Sompolinsky. The tempotron: a neuron that learns spike timing–based decisions. Nature neuroscience, 9(3):420–428, 2006.

[Heb49] Donald Olding Hebb. The organization of behavior: A neuropsychological approach. John Wiley & Sons, 1949.

[HGH14] A Hanuschkin, S Ganguli, and RHR Hahnloser. A hebbian learning rule gives rise to mirror neurons and links them to control theoretic inverse models.

Closing the Loop Around Neural Systems, page 274, 2014.

[HKF02] Richard H R Hahnloser, Alexay A Kozhevnikov, and Michael S Fee. An ultra-sparse code underlies the generation of neural sequences in a songbird.

Nature, 419:65–70, 2002.

[HKP91] John Hertz, Anders Krogh, and Richard G Palmer.Introduction to the Theory of Neural Computation, volume 1. Santa Fe Institute Series (Addison-Wesley Longman, Boston), 1991.

[HNA06] Julie S Haas, Thomas Nowotny, and Henry DI Abarbanel. Spike-timing-dependent plasticity of inhibitory synapses in the entorhinal cortex. Journal of Neurophysiology, 96(6):3305–3313, 2006.

[Hop82] John J Hopfield. Neural networks and physical systems with emergent collec-tive computational abilities.Proceedings of the National Academy of Sciences, 79(8):2554–2558, 1982.

[Hop07] John J Hopfield. Hopfield network. Scholarpedia, 2(5), 2007. revision 91362.

[HTY+14] Kosuke Hamaguchi, Katherine A Tschida, Inho Yoon, Bruce R Donald, and Richard Mooney. Auditory synapses to song premotor neurons are gated off during vocalization in zebra finches. Elife, 3:e01833, 2014.

[HWPG14] Sascha Helduser, Maren Westkott, Klaus Pawelzik, and Onur G¨unt¨urk¨un. The putative pigeon homologue to song bird lman does not modulate behavioral variability. Behavioural brain research, 263:144–148, 2014.

[HZK10] Knut Holthoff, Dejan Zecevic, and Arthur Konnerth. Rapid time course of action potentials in spines and remote dendrites of mouse visual cortex neurons. The Journal of Physiology, 588(7):1085–1096, 2010.

[ID03] Eugene M Izhikevich and Niraj S Desai. Relating stdp to bcm. Neural com-putation, 15(7):1511–1523, 2003.

[KB06] Mimi H Kao and Michael S Brainard. Lesions of an avian basal ganglia circuit prevent context-dependent changes to song variability. Journal of neurophysiology, 96(3):1441–1455, 2006.

[KDB05] Mimi H Kao, Allison J Doupe, and Michael S Brainard. Contributions of an avian basal ganglia–forebrain circuit to real-time modulation of song. 2005.

[LB13] Rodrigo Laje and Dean V Buonomano. Robust timing and motor patterns by taming chaos in recurrent neural networks. Nature neuroscience, 16(7):925–

933, 2013.

[Leo04] Anthony Leonardo. Experimental test of the birdsong error-correction model.

Proceedings of the National Academy of Sciences of the United States of Amer-ica, 101(48):16935–16940, 2004.

[LHS+07] Karri P Lamsa, Joost H Heeroma, Peter Somogyi, Dmitri A Rusakov, and Dimitri M Kullmann. Anti-Hebbian long-term potentiation in the hippocam-pal feedback inhibitory circuit. Science, 315(5816):1262–1266, March 2007.

[LNM05] Robert Legenstein, Christian Naeger, and Wolfgang Maass. What can a neuron learn with spike-timing-dependent plasticity? Neural Computation, 17(11):2337–2382, 2005.

[Mar70] Peter Marler. Birdsong and speech development: Could there be parallels?

there may be basic rules governing vocal learning to which many species conform, including man. American scientist, 58(6):669–673, 1970.

[Mas13] Timothee Masquelier. Neural variablility, or lack thereof. Frontiers in com-putational neuroscience, 7, 2013.

[MLFS97] Henry Markram, Joachim L¨ubke, Michael Frotscher, and Bert Sakmann. Reg-ulation of synaptic efficacy by coincidence of postsynaptic APs and EPSPs.

Science, 275(5297):213–215, 1997.

[Moo09] Richard Mooney. Neural mechanisms for learned birdsong. Learning & Mem-ory, 16(11):655–669, 2009.

[Moo14] Richard Mooney. Auditoryvocal mirroring in songbirds. Phil. Trans. R. Soc.

B, 369:20130179, 2014.

[MP69] Marvin Minsky and Seymour Papert. Perceptrons. 1969.

[MR ¨OS14] Raoul-Martin Memmesheimer, Ran Rubin, Bence P ¨Olveczky, and Haim Som-polinsky. Learning precisely timed spikes. Neuron, 82(4):925–938, 2014.

[MS95] Zachary F Mainen and Terrence J Sejnowski. Reliability of spike timing in neocortical neurons. Science, 268(5216):1503–1506, 1995.

[NSL76] Fernando Nottebohm, Tegner M Stokes, and Christiana M Leonard. Cen-tral control of song in the canary, serinus canarius. Journal of Comparative Neurology, 165(4):457–486, 1976.

[ ¨OAF05] Bence P ¨Olveczky, Aaron S Andalman, and Michale S Fee. Vocal experimen-tation in the juvenile songbird requires a basal ganglia circuit. PLoS Biol, 3(5):e153, 2005.

[PGCZ14] Marko A Popovic, Xin Gao, Nicholas T Carnevale, and Dejan Zecevic. Cor-tical dendritic spine heads are not electrically isolated by the spine neck from membrane potential signals in parent dendrites. Cerebral Cortex, 24(2):385–

395, 2014.

[PK10] Filip Ponulak and Andrzej Kasinski. Supervised learning in spiking neural networks with ReSuMe: sequence learning, classification, and spike shifting.

Neural Computation, 22(2):467–510, 2010.

[PPNM08] Jonathan F Prather, S Peters, S Nowicki, and R Mooney. Precise auditory–

vocal mirroring in neurons for learned vocal communication. Nature, 451(7176):305–310, 2008.

[PS09] Lucy M Palmer and Greg J Stuart. Membrane potential changes in dendritic spines during action potentials and synaptic input. The Journal of Neuro-science, 29(21):6897–6903, 2009.

[Ros58] Frank Rosenblatt. The perceptron: a probabilistic model for information storage and organization in the brain. Psychological Review, 65(6):386, 1958.

[RPMJ04] Anton Reiner, David J Perkel, Claudio V Mello, and Erich D Jarvis. Songbirds and the revised avian brain nomenclature. Annals of the New York Academy of Sciences, 1016(1):77–108, 2004.

[SB06] Jon T Sakata and Michael S Brainard. Real-time contributions of audi-tory feedback to avian vocal motor control. The Journal of neuroscience, 26(38):9619–9628, 2006.

[SBC02] Harel Z Shouval, Mark F Bear, and Leon N Cooper. A unified model of NMDA receptor-dependent bidirectional synaptic plasticity. Proceedings of the National Academy of Sciences, 99(16):10831–10836, 2002.

[SH06] Per Jesper Sj¨ostr¨om and Michael H¨ausser. A cooperative switch determines the sign of synaptic plasticity in distal dendrites of neocortical pyramidal neurons. Neuron, 51(2):227–238, 2006.

[SMA00] Sen Song, Kenneth D Miller, and Larry F Abbott. Competitive hebbian learn-ing through spike-timlearn-ing-dependent synaptic plasticity. Nature Neuroscience, 3(9):919–926, 2000.

[SN91] Constance Scharff and Fernando Nottebohm. A comparative study of the be-havioral deficits following lesions of various parts of the zebra finch song sys-tem: implications for vocal learning.The Journal of neuroscience, 11(9):2896–

2913, 1991.

[STN01] Per Jesper Sj¨ostr¨om, Gina G Turrigiano, and Sacha B Nelson. Rate, tim-ing, and cooperativity jointly determine cortical synaptic plasticity. Neuron, 32(6):1149–1164, 2001.

[STN04] Per Jesper Sj¨ostr¨om, Gina G Turrigiano, and Sacha B Nelson.

Endocannabinoid-Dependent Neocortical Layer-5 LTD in the Absence of Postsynaptic Spiking. Journal of Neurophysiology, 92(6):3338–3343, August 2004.

[SWM03] Christopher B Sturdy, J Martin Wild, and Richard Mooney. Respiratory and telencephalic modulation of vocal motor neurons in the zebra finch. The Journal of neuroscience, 23(3):1072–1086, 2003.

[TBW+11] John A Thompson, Mark J Basista, Wei Wu, Richard Bertram, and Frank Johnson. Dual pre-motor contribution to songbird syllable variation. The Journal of Neuroscience, 31(1):322–330, 2011.

[US14] Robert Urbanczik and Walter Senn. Learning by the dendritic prediction of somatic spiking. Neuron, 81:521–528, 2014.

[VGO+13] Matthijs B Verhoog, Natalia A Goriounova, Joshua Obermayer, Jasper Stroeder, JJ Johannes Hjorth, Guilherme Testa-Silva, Johannes C Baayen, Christiaan PJ de Kock, Rhiannon M Meredith, and Huibert D Mansvelder.

Mechanisms underlying the rules for associative plasticity at adult human neo-cortical synapses. The Journal of Neuroscience, 33(43):17197–17208, 2013.

[VKLL16] Daniela Vallentin, Georg Kosche, Dina Lipkind, and Michael A Long. Inhibi-tion protects acquired song segments during vocal learning in zebra finches.

Science, 351(6270):267–271, 2016.

[VP96] Jonathan Victor and Keith P Purpura. Nature and precision of temporal coding in visual cortex: A metric-space analysis. Journal of Neurophysiology, 76(2):1310–1326, 1996.

[vR01] Mark CW van Rossum. A novel spike distance. Neural Computation, 13(4):751–763, 2001.

[VSZ+11] TP Vogels, Henning Sprekeler, Friedemann Zenke, Claudia Clopath, and Wul-fram Gerstner. Inhibitory plasticity balances excitation and inhibition in sen-sory pathways and memory networks. Science, 334(6062):1569–1573, 2011.

[VT01] Rufin Van Rullen and Simon J Thorpe. Rate coding versus temporal or-der coding: What the retional ganglion cells tell the visual cortex. Neural Computation, 13(6):1255–1283, 2001.

[WGNB05] Huai-Xing Wang, Richard C Gerkin, David W Nauen, and Guo-Qiang Bi.

Coactivation and timing-dependent integration of synaptic potentiation and depression. Nature Neuroscience, 8(2):187–193, 2005.

[XAS14] Mingshan Xue, Bassam V Atallah, and Massimo Scanziani. Equalizing excitation–inhibition ratios across visual cortical neurons. Nature, 511:596–

600, June 2014.

[XJPD12] Shengjin Xu, Wanchen Jiang, Mu-Ming Poo, and Yang Dan. Activity recall in a visual cortical ensemble. Nature Neuroscience, 15(3):449–455, 2012.

[XS04] Xiaohui Xie and H Sebastian Seung. Learning in neural networks by rein-forcement of irregular spiking. Physical Review E, 69(4):041909, 2004.

[XZZ13] Yan Xu, Xiaoqin Zeng, and Shuiming Zhong. A new supervised learning algorithm for spiking neurons. Neural computation, 25(6):1472–1511, 2013.

Fachbereich 1 Dr.- rer. - nat.

E r k l ä r u n g

Hiermit versichere ich, dass ich 1. die Arbeit ohne fremde

Hilfe angefertigt habe,

2. keine anderen als die von mir an- gegeben Quellen und Hilfsmittel

benutzt habe und

3. die den benutzten Werken wörtlich oder inhaltlich entnommenen Stellen als solche kenntlich gemacht habe.

,den

( Unterschrift )