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+ Synaptic Theory of Working Memory Caitlin Hof & Steven Larson + Spike Timing Measurements Measures the firing rate of neurons Is assumed to be a correlate to working memory Is done with single neuron electrodes (electrophysiology) and EEG Has allowed for the theoretical measurement and identification of working memory length and location + Synaptic Theory Memories are preserved in the short term by constant activation of neurons Measuring activity spikes shows what is thought to be memory Memories in working memory can be reactivated by a general stimulus Adjusting chemical balance within the synapses affects length of working memory + Synaptic Theory (Continued) Short term plasticity leads to the creation of Polychronous Neuronal Groups PNGs form and dissipate to hold temporary memories These are energy inefficient but hold memory in its purest form + Visual STM Where it occurs are Prefrontal Cortex and Inferior Temporal Regions Ablations to area TE affect STM while the Perirhinal cortex affects LTM Activity in the PFC takes place in fewer neurons but continues during distractors Suggests that a circuit like series of neurons stores the stimulus for later comparison + + Synaptic Theory of Working Memory Gianluigi Mongillo, Omri Barak, & Misha Tsodyks Science Volume 319 14 March 2008 + Working Memory Holds information temporarily Processing Crucial to cognitive tasks + Delayed response paradigm Stimulus kept until task execution Stimulus specific spiking activity – neuronal correlate of working memory + Theoretical Construct Delay activity – intrinsic cell properties OR Persistent reverberations in selective neural populations + Theoretical Construct Issues Both theories suggest working memory might not be entirely in spiking memory Energetically expensive to hold information in spiking form High metabolic cost of action potentials + Alternative Theory Presented alternative account Based on properties of excitatory synaptic transmission in prefrontal cortex + Prefrontal Cortex Higher order functions Cognitive behavior Personality Decisions Planning Thoughts and actions for internal goals Executive function https://www.google.com/search?hl=en&site=imghp&tbm=isch&source=hp&biw=1280&bih=666&q=pref ortex&oq=prefrontal+cortex&gs_l=img.3...0.0.0.147612.0.0.0.0.0.0.0.0..0.0...0.0...1ac..6.img.76JnFhhqhYc =1EeZTE69hjOEhM%3A%3BV6_SpVPUJJDzTM%3Bhttp%253A%252F%252Fwww.humansoul.com%252FP %252520cortex.jpg%3Bhttp%253A%252F%252Fwww.humansoul.com%252Fprefrontal_cortex1.html%3B 314 + Alternative Theory Prefrontal cortex implicated in working memory Excitatory synaptic transmission is facilitatory Sensory -> depression + Alternative Theory Proposed: An item is maintained in the working memory state by short-term synaptic facilitation mediated by increased residual calcium levels at the presynaptic terminal of the neurons that code for this item + A Little Biology https://www.google.com/search?hl=en&q=neurons%20with%20calcium&psj=1&bav=on.2,or.r_qf.&bvm=bv.43828540,d.b2U&biw 280&bih=666&um=1&ie=UTF-8&tbm=isch&source=og&sa=N&tab=wi&ei=fAVIUazmLMXT2QWslYDYBg#imgrc=egmQIIRhiuVYM%3A%3B6dqfSBdLjAvs0M%3Bhttp%253A%252F%252Fclick4biology.info%252Fc4b%252F6%252Fimages%252F6.5%252Fs apse.gif%3Bhttp%253A%252F%252Fclick4biology.info%252Fc4b%252F6%252Fhum6.5.htm%3B440%3B289 + Alternative Theory Removal of residual calcium from presynaptic terminals is relatively slow Memory can be transiently held for about 1 s without enhanced spike activated + Methods Recurrent network of integrate and fire neurons Network encodes a set of memories (items) Composed of randomly selective populations of excitatory neurons Connections between the neurons coding for the same memory stronger than connections between different populations + Figure 1B + Figure 1A + Figure 1 A + Memory Maintenance with Synaptic Facilitation (Figure 2A) Stimulation – loading one item into working memory by transient external excitation to the corresponding neuronal population + Memory Maintenance with Synaptic Facilitation (Figure 2B) + Memory Maintenance with Synaptic Facilitation (Figure 2C) + Memory Maintenance with Synaptic Facilitation (Figure 2) The use of residual calcium at synaptic terminals as a memory buffer requires low emmision rates Buffer content is not substantially affected by the neural activity of the rest of the cortex + Conclusions Working memory can be maintained by short-term synaptic facilitation Accumulation of residual calcium in the presynaptic terminals could carry the information about the recalled memory in a working form, reducing the need for metabolically costly action potentials Memories -> spiking memory A result of global reactivating input to the network or by virtue of the intrinsic network dynamics + Conclusions Not all stimuli enter working memory Decision to allow items into working memory is mediated by attention – represented by the global excitatory output Performance of human observers on memory tasks positively correlates with the level of neural activity during the presentation of the items Model predicts that residual calcium at the presynaptic terminals should be enhanced during working memory even if there is no increase in firing rate + Conclusions Suppressing the spiking activity for several hundred milliseconds should still allow for the memory to reactivate after the suppressing input is withdrawn Groups of neurons could exhibit brief epochs of coherent firing Model provides a possible target for a pharmacological interference with working memory Manipulations that modify the facilitation/depression balance in the memory related cortical areas are predicted to have a strong effect on the stability and duration of memory