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Basics of Computational Neuroscience
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Lecture: Computational Neuroscience, Contents
1) Introduction The Basics – A reminder: 1) Brain, Maps, Areas, Networks, Neurons, and Synapses The tough stuff: 2,3) Membrane Models 3,4) Spiking Neuron Models 5) Calculating with Neurons I: adding, subtracting, multiplying, dividing 5,6) Calculating with Neurons II: Integration, differentiation 6) Calculating with Neurons III: networks, vector-/matrix- calculus, assoc. memory 6,7) Information processing in the cortex I: Neurons as filters 7) Information processing in the cortex II:Correlation analysis of neuronal connections 7,8) Information processing in the cortex III: Neural Codes and population responses 8) Information processing in the cortex IV: Neuronal maps Something interesting – the broader perspective 9) On Intelligence and Cognition – Computational Properties? Motor Function 10,11) Models of Motor Control Adaptive Mechanisms 11,12) Learning and plasticity I: Physiological mechanisms and formal learning rules 12,13) Learning and plasticity II: Developmental models of neuronal maps 13) Learning and plasticity III: Sequence learning, conditioning Higher functions 14) Memory: Models of the Hippocampus 15) Models of Attention, Sleep and Cognitive Processes Lecture: Computational Neuroscience, Contents
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The Interdisciplinary Nature of Computational Neuroscience
What is computational neuroscience ?
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Different Approaches towards Brain and Behavior
Neuroscience: Environment Stimulus Behavior Reaction
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Psychophysics (human behavioral studies):
Black Box Environment Stimulus Behavior Reaction
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Neurophysiology: Environment Stimulus Behavior Reaction
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Theoretical/Computational Neuroscience:
Environment Stimulus Behavior Reaction
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Levels of information processing in the nervous system
CNS 1m Sub-Systems 10cm Areas / „Maps“ 1cm Local Networks 1mm Neurons 100mm Synapses 1mm Molecules 0.1mm
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CNS (Central Nervous System):
Systems Areas Local Nets Neurons Synapses Molekules
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Cortex: CNS Systems Areas Local Nets Neurons Synapses Molekules
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Where are things happening in the brain.
The Phrenologists view at the brain (18th-19th centrury) Is the information represented locally ? Molekules Synapses Neurons Local Nets Areas Systems CNS
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Results from human surgery
CNS Systems Areas Local Nets Neurons Synapses Molekules
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Results from imaging techniques – There are maps in the brain
CNS Systems Areas Local Nets Neurons Synapses Molekules
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Visual System: More than 40 areas !
Parallel processing of „pixels“ and image parts Hierarchical Analysis of increasingly complex information Many lateral and feedback connections CNS Systems Areas Local Nets Neurons Synapses Molekules
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Primary visual Cortex:
CNS Systems Areas Local Nets Neurons Synapses Molekules
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Retinotopic Maps in V1: V1 contains a retinotopic map of the visual Field. Adjacent Neurons represent adjacent regions in the retina. That particular small retinal region from which a single neuron receives its input is called the receptive field of this neuron. V1 receives information from both eyes. Alternating regions in V1 (Ocular Dominanz Columns) receive (predominantely) Input from either the left or the right eye. Each location in the cortex represents a different part of the visual scene through the activity of many neurons. Different neurons encode different aspects of the image. For example, orientation of edges, color, motion speed and direction, etc. V1 decomposes an image into these components. Molekules Synapses Neurons Local Nets Areas Systems CNS
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Orientation selectivity in V1:
stimulus Orientation selective neurons in V1 change their activity (i.e., their frequency for generating action potentials) depending on the orientation of a light bar projected onto the receptive Field. These Neurons, thus, represent the orientation of lines oder edges in the image. Their receptive field looks like this: Molekules Synapses Neurons Local Nets Areas Systems CNS
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Superpositioning of maps in V1:
Thus, neurons in V1 are orientation selective. They are, however, also selective for retinal position and ocular dominance as well as for color and motion. These are called „features“. The neurons are therefore akin to „feature-detectors“. For each of these parameter there exists a topographic map. These maps co-exist and are superimposed onto each other. In this way at every location in the cortex one finds a neuron which encodes a certain „feature“. This principle is called „full coverage“. Molekules Synapses Neurons Local Nets Areas Systems CNS
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Local Circuits in V1: stimulus
Orientation selective cortical simple cell Local Circuits in V1: stimulus Selectivity is generated by specific connections Molekules Synapses Neurons Local Nets Areas Systems CNS
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Layers in the Cortex: CNS Systems Areas Local Nets Neurons Synapses
Molekules
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Local Circuits in V1: LGN inputs Cell types Circuit Spiny stellate
Molekules Synapses Neurons Local Nets Areas Systems CNS LGN inputs Cell types Circuit Spiny stellate cell Smooth stellate cell
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Structure of a Neuron: At the dendrite the incoming
signals arrive (incoming currents) At the soma current are finally integrated. At the axon hillock action potential are generated if the potential crosses the membrane threshold The axon transmits (transports) the action potential to distant sites Molekules Synapses Neurons Local Nets Areas Systems CNS At the synapses are the outgoing signals transmitted onto the dendrites of the target neurons
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Different Types of Neurons:
dendrite dendrite Bipolar cell Unipolar cell axon soma axon soma (Invertebrate N.) Retinal bipolar cell Different Types of Multi-polar Cells Hippocampal pyramidal cell Purkinje cell of the cerebellum Spinal motoneuron
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Cell membrane: The cell membrane separates intra- from extra-cellular spaces Na+ and Cl- ions are more concentrated outside, while negative ions (A-) and plenty of K+ are more concentrated inside. Due to differences in the ion-concenrations across the membrane a potential difference arises: In addition, the membrane acts like a capacitor: Cl- K+
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Ion channels: Ion channels consist of big (protein) molecules which are inserted into to the membrane and connect intra- and extracellular space. Channels act as a restistance against the free flow of ions.
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Membrane - Circuit diagram:
rest
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Membrane - Circuit Diagram (advanced version):
The whole thing gets more complicated due to the fact that there are many different ion channels all of which have their own characteristics depending on the momentarily existing state of the cell. The conducitvity of a channel depends on the membrane potential and on the concentration difference between intra- and extracellular space (and sometimes also on other parameters). One needs a computer simulation to describe this complex membrane behavior. The whole thing gets more complicated due to the fact that there are many different ion channels all of which have their own characteristics depending on the momentarily existing state of the cell. The conducitvity of a channel depends on the membrane potential and on the concentration difference between intra- and extracellular space (and sometimes also on other parameters). The whole thing gets more complicated due to the fact that there are many different ion channels all of which have their own characteristics depending on the momentarily existing state of the cell.
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Structure of a Neuron: At the dendrite the incoming
signals arrive (incoming currents). Signals propagate (normally) in a passive, electrotonic way towards the soma At the dendrite the incoming signals arrive (incoming currents). Signals propagate (normally) in a passive, electrotonic way towards the soma At the dendrite the incoming signals arrive (incoming currents). Signals propagate (normally) in a passive, electrotonic way towards the soma At the dendrite the incoming signals arrive (incoming currents). Signals propagate (normally) in a passive, electrotonic way towards the soma At the soma current are finally integrated. At the axon hillock action potential are generated if the potential crosses the membrane threshold The axon transmits (transports) the action potential to distant sites Molekules Synapses Neurons Local Nets Areas Systems CNS At the synapses are the outgoing signals transmitted onto the dendrites of the target neurons
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Electrotonic Signal Propagation:
Injected Current Membrane Potential Injected current flows out from the cell evenly across the membrane. The cell membrane has everywhere the same potential. The change in membrane potention follows an exponential with time constant: t = RC Injected current flows out from the cell evenly across the membrane. The cell membrane has everywhere the same potential. Injected current flows out from the cell evenly across the membrane.
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Electrotonic Signal Propagation:
The potential decays along a dendrite (or axon) according to the distance from the current injection site. The potential decays along a dendrite (or axon) according to the distance from the current injection site. At every location the temporal response follows an exponential but with ever decreasing amplitude. If plotting only the maxima against the distance then you will get another exponential. The potential decays along a dendrite (or axon) according to the distance from the current injection site. At every location the temporal response follows an exponential but with ever decreasing amplitude. The potential decays along a dendrite (or axon) according to the distance from the current injection site. At every location the temporal response follows an exponential but with ever decreasing amplitude. If plotting only the maxima against the distance then you will get another exponential. Different shape of the potentials in the dendrite and the soma of a motoneuron.
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Compartment-Model: One can model the electrotonic propagation of potentials in the complex dendritic tree by subdividing the tree into small (cyklindrical) compartments. For each compartment the membrane equations can then be solved and integrated. (All this is tedious and complicated.)
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Structure of a Neuron: At the axon hillock action potential
At the dendrite the incoming signals arrive (incoming currents) At the soma current are finally integrated. At the axon hillock action potential are generated if the potential crosses the membrane threshold. At the axon hillock action potential are generated if the potential crosses the membrane threshold. At the axon hillock action potential are generated if the potential crosses the membrane threshold. At the axon hillock action potential are generated if the potential crosses the membrane threshold. The axon transmits (transports) the action potential to distant sites Molekules Synapses Neurons Local Nets Areas Systems CNS At the synapses are the outgoing signals transmitted onto the dendrites of the target neurons
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Action potential
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Action Potential / Shapes:
Cat - Heart Rat - Muscle Squid Giant Axon
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Structure of a Neuron: The axon transmits (transports) the
At the dendrite the incoming signals arrive (incoming currents) At the soma current are finally integrated. At the axon hillock action potential are generated if the potential crosses the membrane threshold. The axon transmits (transports) the action potential to distant sites The axon transmits (transports) the action potential to distant sites The axon transmits (transports) the action potential to distant sites The axon transmits (transports) the action potential to distant sites Molekules Synapses Neurons Local Nets Areas Systems CNS At the synapses are the outgoing signals transmitted onto the dendrites of the target neurons
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Propagation of an Action Potential:
Distance Time Local current loops mm2 membrane area Open channels per Action potentials propagate without being diminished (active process). All sites along a nerve fiber will be depolarized until the potential passes threshold. As soon as this happens a new AP will be elicited at some distance to the old one. Main current flow is across the fiber. Action potentials propagate without being diminished (active process). All sites along a nerve fiber will be depolarized until the potential passes threshold. As soon as this happens a new AP will be elicited at some distance to the old one. Action potentials propagate without being diminished (active process).
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Structure of a Neuron: At the dendrite the incoming signals arrive (incoming currents) At the soma current are finally integrated. At the axon hillock action potential are generated if the potential crosses the membrane threshold The axon transmits (transports) the action potential to distant sites CNS At the synapses are the outgoing signals transmitted onto the dendrites of the target neurons At the synapses are the outgoing signals transmitted onto the dendrites of the target neurons At the synapses are the outgoing signals transmitted onto the dendrites of the target neurons At the synapses are the outgoing signals transmitted onto the dendrites of the target neurons Systems Areas Local Nets Neurons Synapses Molekules
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Chemical synapse Neurotransmitter Receptors
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Neurotransmitters Chemicals (amino acids, peptides, monoamines) that transmit, amplify and modulate signals between neuron and another cell. Cause either excitatory or inhibitory PSPs. Glutamate – excitatory transmitter GABA, glycine – inhibitory transmitter
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Synaptic Transmission:
Synapses are used to transmit signals from the axon of a source to the dendrite of a target neuron. There are electrical (rare) and chemical synapses (very common) At an electrical synapse we have direct electrical coupling (e.g., heart muscle cells). At a chemical synapse a chemical substance (transmitter) is used to transport the signal. Electrical synapses operate bi-directional and are extremely fast, chem. syn. operate uni-directional and are slower. Synapses are used to transmit signals from the axon of a source to the dendrite of a target neuron. There are electrical (rare) and chemical synapses (very common) At an electrical synapse we have direct electrical coupling (e.g., heart muscle cells). At a chemical synapse a chemical substance (transmitter) is used to transport the signal. Electrical synapses operate bi-directional and are extremely fast, chem. syn. operate uni-directional and are slower. Chemical synapses can be excitatory or inhibitory they can enhance or reduce the signal change their synaptic strength (this is what happens during learning). Synapses are used to transmit signals from the axon of a source to the dendrite of a target neuron. There are electrical (rare) and chemical synapses (very common) At an electrical synapse we have direct electrical coupling (e.g., heart muscle cells). At a chemical synapse a chemical substance (transmitter) is used to transport the signal. Synapses are used to transmit signals from the axon of a source to the dendrite of a target neuron. There are electrical (rare) and chemical synapses (very common) At an electrical synapse we have direct electrical coupling (e.g., heart muscle cells). Synapses are used to transmit signals from the axon of a source to the dendrite of a target neuron. Synapses are used to transmit signals from the axon of a source to the dendrite of a target neuron. There are electrical (rare) and chemical synapses (very common)
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Structure of a Chemical Synapse:
Axon Motor Endplate (Frog muscle) Synaptic cleft Active zone vesicles Muscle fiber Presynaptic membrane Postsynaptic membrane Synaptic cleft
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What happens at a chemical synapse during signal transmission:
Pre-synaptic action potential Concentration of transmitter in the synaptic cleft Post-synaptic The pre-synaptic action potential depolarises the axon terminals and Ca2+-channels open. Ca2+ enters the pre-synaptic cell by which the transmitter vesicles are forced to open and release the transmitter. Thereby the concentration of transmitter increases in the synaptic cleft and transmitter diffuses to the postsynaptic membrane. Transmitter sensitive channels at the postsyaptic membrane open. Na+ and Ca2+ enter, K+ leaves the cell. An excitatory postsynaptic current (EPSC) is thereby generated which leads to an excitatory postsynaptic potential (EPSP). The pre-synaptic action potential depolarises the axon terminals and Ca2+-channels open. Ca2+ enters the pre-synaptic cell by which the transmitter vesicles are forced to open and release the transmitter. Thereby the concentration of transmitter increases in the synaptic cleft and transmitter diffuses to the postsynaptic membrane. The pre-synaptic action potential depolarises the axon terminals and Ca2+-channels open. Ca2+ enters the pre-synaptic cell by which the transmitter vesicles are forced to open and release the transmitter. The pre-synaptic action potential depolarises the axon terminals and Ca2+-channels open.
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Neurotransmitters and their (main) Actions:
Transmitter Channel-typ Ion-current Action Acetylecholin nicotin. Receptor Na+ and K+ excitatory Glutamate AMPA / Kainate Na+ and K+ excitatory GABA GABAA-Receptor Cl- inhibitory Glycine Cl- inhibitory Acetylecholin muscarin. Rec. - metabotropic, Ca2+ Release Transmitter Channel-typ Ion-current Action Acetylecholin nicotin. Receptor Na+ and K+ excitatory Glutamate AMPA / Kainate Na+ and K+ excitatory GABA GABAA-Receptor Cl- inhibitory Glycine Cl- inhibitory Acetylecholin muscarin. Rec. - metabotropic, Ca2+ Release Glutamate NMDA Na+, K+, Ca2+ voltage dependent blocked at resting potential Transmitter Channel-typ Ion-current Action Acetylecholin nicotin. Receptor Na+ and K+ excitatory Glutamate AMPA / Kainate Na+ and K+ excitatory GABA GABAA-Receptor Cl- inhibitory Glycine Cl- inhibitory Transmitter Channel-typ Ion-current Action Acetylecholin nicotin. Receptor Na+ and K+ excitatory Glutamate AMPA / Kainate Na+ and K+ excitatory Transmitter Channel-typ Ion-current Action Transmitter Channel-typ Ion-current Action Acetylecholin nicotin. Receptor Na+ and K+ excitatory Transmitter Channel-typ Ion-current Action Acetylecholin nicotin. Receptor Na+ and K+ excitatory Glutamate AMPA / Kainate Na+ and K+ excitatory GABA GABAA-Receptor Cl- inhibitory
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Synaptic Plasticity
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