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3: Neurons and Synapses Brain and Behavior David Eagleman Jonathan Downar Chapter Outline The Cells of the Brain Synaptic Transmission: Chemical Signaling in the Brain Spikes: Electrical Signaling in the Brain What Do Spikes Mean? The Neural Code Individuals and Populations 2 The Cells of the Brain Neurons: A Close-Up View Many Different Types of Neurons Glial Cells 3 Neurons: A Close-Up View Ramon y Cajal established the Neuron Doctrine, which states that the brain is made of many small, discrete cells. There are almost 100 billion neurons in the human brain. These neurons are like any other cell in the body, with a membrane, a nucleus, and specialized organelles. 4 Neurons: A Close-Up View 5 Neurons: A Close-Up View Neurons have four important regions. Dendrites: Branching projections that collect information 6 Neurons: A Close-Up View Neurons have four important regions. Soma (Cell Body): Contains the nucleus and integrates information 7 Neurons: A Close-Up View Neurons have four important regions. Axon: Conducts the neural signal across a long distance 8 Neurons: A Close-Up View Neurons have four important regions. Axon terminals: Small swellings that release signals to affect other neurons Chemical signals, known as neurotransmitters, cross small gaps, known as synapses. It is estimated that there are about 500 trillion synapses in the adult brain. 9 Neurons: A Close-Up View 10 Many Different Types of Neurons Neurons can be classified by their function: Sensory neurons carry information to the brain. Motor neurons carry information from the brain to the muscles. Interneurons convey the signals around the nervous system. 11 Many Different Types of Neurons 12 Many Different Types of Neurons Neurons can be classified by their shape: Multipolar neurons have many dendrites. Bipolar neurons have one dendrite and one axon. Monopolar neurons have only one projection from the soma, which branches to form the axon and the dendrite. 13 Many Different Types of Neurons 14 Glial Cells Glia play many roles within the nervous system: Speeding up the neuronal signaling Regulating extracellular chemicals Enabling neurons to modify their connections 15 Glial Cells Oligodendrocytes, in the central nervous system, and Schwann cells, in the peripheral nervous system, wrap myelin around axons to speed up signals. Nodes of Ranvier are small gaps in the myelin sheath. 16 Glial Cells 17 Glial Cells Astrocytes regulate extracellular chemicals and regulate local blood flow. Microglia provide immune system functions for the central nervous system. 18 Synaptic Transmission: Chemical Signaling in the Brain Release of Neurotransmitter at the Synapse Types of Neurotransmitters Receptors Postsynaptic Potentials 19 Release of Neurotransmitter at the Synapse Neurotransmitters are chemicals released by the presynaptic cell to affect the postsynaptic cell. The synaptic cleft is the 20- to 30-nm space between the cells. The small size of the synaptic cleft allows the concentration of the neurotransmitter to change rapidly. 20 Release of Neurotransmitter at the Synapse 21 Types of Neurotransmitters There are small-molecular-weight neurotransmitters, such as monoamines and amino acids, soluble gases, such as NO and CO, and large-molecular-weight neurotransmitters, which are peptides. Most neurons release one or two small transmitters as well as a peptide. 22 Types of Neurotransmitters 23 Receptors Specialized proteins in the cell membrane Neurotransmitters interact with receptors to affect the postsynaptic cell. Ionotropic receptors allow ions to flow across the membrane, changing the charge of the cell membrane. Metabotropic receptors relay information into the cell using a series of proteins. 24 Receptors 25 Receptors Neurotransmitters only bind to receptors for a short time and need a way to be removed. Degradation: The neurotransmitter is broken apart. Diffusion: The neurotransmitter moves down the concentration gradient and out of the synapse. Reuptake: Neurotransmitter is transported back into the original cell. 26 Receptors 27 Postsynaptic Potentials When at rest, there is a voltage difference between the inside and the outside of the cell. The inside of the cell is more negative than the outside, about -70 mV. 28 Postsynaptic Potentials Excitatory postsynaptic potentials alter the membrane voltage, moving the voltage closer to 0. Inhibitory postsynaptic potentials move the voltage further from 0. Postsynaptic potentials are small (about 1 mV) and fast (a few milliseconds). 29 Postsynaptic Potentials 30 Spikes: Electrical Signaling in the Brain Adding up the Signals How an Action Potential Travels Myelinating Axons to Make the Action Potential Travel Faster Action Potentials Reach the Terminals and Cause Neurotransmitter Release 31 Adding up the Signals Action potentials are all or none. EPSPs and IPSPs combine to affect the membrane voltage. In temporal summation, PSPs arriving at the soma at close to the same time are combined. In spatial summation, PSPs arriving at different locations on the soma are combined. 32 Adding up the Signals 33 Adding up the Signals The soma receives 100s or 1000s of PSPs at a time. EPSPs sum together to depolarize the cell (move the voltage closer to 0). If the membrane voltage reaches threshold (approximately -60 mV), an action potential is generated at the axon hillock. 34 How an Action Potential Travels In neurons at rest, there are more Na+ ions outside the cell and more K+ ions inside the cell. At threshold, voltage-gated Na+ channels open, allowing Na+ ions to flow into the cell, down the chemical concentration and electrical gradients. Voltage-gated K+ channels open, allowing K+ ions to flow out of the cell. 35 How an Action Potential Travels 36 How an Action Potential Travels The current formed by the Na+ ions flows down the neuron, depolarizing the next part of the neuron. There is a refractory period after the action potential, when the voltage-gated Na+ ion channels are less likely to open. Calcium and chloride ions also contribute to the action potential. 37 Myelinating Axons to Make the Action Potential Travel Faster Myelin is interrupted by gaps, known as nodes of Ranvier, where the action potential is regenerated. The action potential jumps from node to node, greatly speeding up transmission. Myelination decreases the amount of energy used by the neuron. 38 Myelinating Axons to Make the Action Potential Travel Faster 39 Action Potentials Cause Neurotransmitter Release Action potentials cause voltage changes in the axon terminals, causing voltage-gated calcium channels to open. Calcium ions cause vesicles with neurotransmitters to bind to the presynaptic membrane. Neurotransmitters are released and cross the synapse. 40 Action Potentials Cause Neurotransmitter Release 41 What Do Spikes Mean? The Neural Code Encoding Stimuli in Spikes Decoding Spikes 42 Encoding Stimuli in Spikes In the brain, there are approximately 100 billion neurons, each sending up to a few hundred action potentials per second. The number of spikes per second is used to describe the neuron’s response to a stimulus. 43 Encoding Stimuli in Spikes 44 Encoding Stimuli in Spikes Neurons have a baseline level of activity, so the neuron can either increase or decrease the firing rate. Research suggests that there may be other coding methods. 45 Encoding Stimuli in Spikes 46 Decoding Spikes A typical neuron receives 10,000 incoming synapses. Neurons may be responding not to individual input but to the average input. 47 Decoding Spikes 48 Individuals and Populations Populations of Neurons Forming a Coalition: What Constitutes a Group? Open Questions for Future Investigation 49 Populations of Neurons Local coding is the idea that stimuli in the outside world are encoded by different neurons. Population coding is the idea that each stimulus is represented by a collection of neurons. Each individual neuron many participate in multiple collections of neurons. 50 Forming a Coalition: What Constitutes a Group? Neurons can be mutually excitatory or a coalition of neurons can support the high firing rate of the population. Neurons may form a coalition by firing in synchrony. 51 Forming a Coalition: What Constitutes a Group? 52 Open Questions for Future Investigation At present, the neural code is not understood. Why do neurons have random changes in membrane voltage? What is the role of the non-spiking neurons in the brain? What is the role of glia in information processing? 53