Adaptation 1. Adaptation – examples and definitions 2. Adaptation as efficient coding 3. A tradeoff between sensitivity and efficiency 4. Adaptation and perception 5. Adaptation occurs over multiple timescales 6. Adaptation involves multiple mechanisms Adaptation with 302 neurons C. elegans Mori, Sasakura, & Kuhara, 2007 Adaptation with thousands (?) of neurons Adaptation with millions (?) of neurons exposed to compressive distortion human psychophysics exposed to expansive distortion Rhodes, Jeffery, Watson, Clifford, & Nakayama 2003 How does this explain that? Why might this be useful? stimulus Smirnakis & Meister, 1997 Adaptation 1. Adaptation – examples and definitions 2. Adaptation as efficient coding 3. A tradeoff between sensitivity and efficiency 4. Adaptation and perception 5. Adaptation occurs over multiple timescales 6. Adaptation involves multiple mechanisms The idea of efficient coding 1. Each neuron should use all parts of its dynamic range with roughly equal frequency. 2. Information encoded in the spike train of one neuron should not be duplicated in the spike train of another neuron. The idea of efficient coding 1. Each neuron should use all parts of its dynamic range with roughly equal frequency. Encoding a natural distribution efficiently Dunn and Rieke, 2006 Encoding a natural distribution efficiently Equal areas of probability density should correspond to equal segments of dynamic range. Laughlin 1981 redrawn in Wark & Fairhall 2007 Changing stimulus statistics Laughlin 1981 redrawn in Wark & Fairhall 2007 Changing stimulus statistics Laughlin 1981 redrawn in Wark & Fairhall 2007 Efficient coding of a changing distribution Laughlin 1981 redrawn in Wark & Fairhall 2007 Adaptation Adaptation 1. Adaptation – examples and definitions 2. Adaptation as efficient coding 3. A tradeoff between sensitivity and efficiency 4. Adaptation and perception 5. Adaptation occurs over multiple timescales 6. Adaptation involves multiple mechanisms An example: light adaptation in the retina cat RGC Enroth-Cudgell & Lennie, 1975 An adaptive change in receptive field low luminance (low signal-to-noise) high luminance (high signal-to-noise) spatial integrator high redundancy spatial edge detector low redundancy An adaptive change in receptive field low luminance (low signal-to-noise) high luminance (high signal-to-noise) temporal integrator high redundancy temporal edge detector low redundancy Adaptation 1. Adaptation – examples and definitions 2. Adaptation as efficient coding 3. A tradeoff between sensitivity and efficiency 4. Adaptation and perception 5. Adaptation occurs over multiple timescales 6. Adaptation involves multiple mechanisms Repulsive perceptual adaptation Repulsive perceptual adaptation http://homepages.inf.ed.ac.uk/jbednar/tae.html The cricket cercal system see http://www.biol.sc.edu/~vogt/courses/neuro/neurolabs.html Bacon & Murphey 1984 cell 1 cell 2 cell 3 cell 4 r / rmax wind direction (degrees) v Theunissen & Miller 1991 Salinas & Abbott 1994 Attractive neural adaptation ○ before ● after adapting stimulus adapting stimulus macaque MT Kohn and Movshon, 2004 Attractive neural adaptation can explain repulsive perceptual adaptation Kohn and Movshon, 2004 Adaptation 1. Adaptation – examples and definitions 2. Adaptation as efficient coding 3. A tradeoff between sensitivity and efficiency 4. Adaptation and perception 5. Adaptation occurs over multiple timescales 6. Adaptation involves multiple mechanisms Adaptation on many timescales in the same neuron salamander RGC Smirnakis et al. 1997 Adaptation on many timescales in the same neuron velocity blowfly H1 Fairhall et al. 2001 Adaptation on many timescales in the same neuron cascade of exponential processes single exponential process model Drew and Abbott, 2006 Adaptation Adaptation 1. Adaptation – examples and definitions 2. Adaptation as efficient coding 3. A tradeoff between sensitivity and efficiency 4. Adaptation and perception 5. Adaptation occurs over multiple timescales 6. Adaptation involves multiple mechanisms Mechanisms of adaptation: intrinsic mechanisms Vm Iinj Lancaster and Nicoll, 1987 Mechanisms of adaptation: synaptic mechanisms Tsodyks & Markram 1997 Mechanisms of adaptation: circuit mechanisms Hosoya et al., 2005 Costs of adaptation Adaptation worsens the ability of a system to encode the absolute magnitude of a stimulus. Noise in the signals controlling gain can produce noisy fluctuations in gain. input output