Electrical Properties of Nerve Cells 24.1.13 +29.1.13 The resting membrane potential Action Potential of Excitable Cells Graded and Action Potential Cells which can respond to a stimulus are said to be excitable (nerve and muscle cells) There is a threshold value for the intensity of a stimulus which can generate an action potential Stimulus less than threshold value will generate a “graded potential” which cannot be transferred over long distances When stimulus is strong enough it will generate an “action potential” that can be transferred over large distance Dendrites: receive information and undergo graded potentials. Neuron Axons: undergo action potentials to deliver information, Excitable Nerve cell A single neuron typically consists of three basic parts 1. Dendrites 2. The cell body 3. The axon • Dendrites and cell body are specialized to receive information and produce graded potentials: In most neurons, the plasma membrane of the dendrites and cell body contain protein receptors that bind chemical messengers/neurotransmitters from other neurons Graded potentials are produced in response to stimulus (here neurotransmitter released from other neurons) • Axons are specialized to deliver information by generation of action potentials Axon hillock is the portion of the axon where action potentials are triggered or initiated by the graded potential if it is of sufficient magnitude The action potentials are then conducted along the axon from axon hillock to axon terminals Electrical Signal Generation • Action potentials can be initiated only in portions of the membrane with abundant voltage gated Na+ channels • Sites of a nerve cell specialized for graded potentials such as dendrites and cell body do not undergo action potentials because they have less voltage gated Na+ channels • Graded potentials generated in response to a stimulus can spread to adjacent areas of the membrane before dying out Electrical Signal Generation • The graded responses produced throughout the dendrites or cell body is summed spatially and temporally, and if the summed response is large enough to pass the threshold by the time it reaches axon hillock, an action potential will be generated at axon hillock. • The axon hillock has the lowest threshold in the neuron because this region has a much higher density of voltage gated Na+ channels than anywhere else in the neurons • Action potential originates at axon hillock Graded Potential • They are local changes in membrane potential that occur in varying grades or degrees of magnitude or strength in response to a stimulus • For example membrane potential could change from -70mV to -60mV (a 10mV graded potential) or from -70mV to -50mV (a 20mV graded potential) • Graded potentials are usually produced by a specific triggering event that causes gated ion channels to open in a specialized region of the excitable cell membrane • The resultant ion movement produces the graded potential which most commonly is a depolarization resulting from Na+ entry The size of a graded potential (here, graded depolarizations) is proportionate to the intensity of the stimulus. The duration of a graded potential (here, graded depolarizations) is proportionate to the duration of the triggering event Graded potential spread by passive current flow Graded Potential die out over short distance • As the current flows along the membrane, some of the current leaks through open leak channels (mainly K+) in the neighboring areas. As a result the membrane potential progressively decreases with increasing distance from the source point Length constant of a membrane • This spatial pattern is exponential and the distance where the voltage changes to 37% of its original value is the “ length constant” • Larger the length constant, farther the graded potential can travel along the membrane Spatial or temporal summation of graded potentials • Graded responses can interact with each other and can be spatially or temporally summed • If two graded potentials occur at the same time in close enough /same places, their effects add up. This is called “spatial summation” • If two graded potentials occur at the same place in succession, their effects add up. This is called temporal summation • As an analogy, spatial summation is like using many shovels to fill up a hole all at once. Temporal summation is like using a single shovel to fill up a hole over time. Both methods work to fill up the hole Panel 1: Panel 2: Panel 3: Panel 4: Panel 5: Two distinct, non-overlapping, graded depolarizations. Two overlapping graded depolarizations demonstrate temporal summation. Distinct actions of stimulating neurons A and B demonstrate spatial summation. A and B are stimulated enough to cause a suprathreshold graded depolarization, so an action potential results. Neuron C causes a graded hyperpolarization; A and C effects add, cancel each other out. Graded Potential • Graded potential causes potential change in limited areas • The graded potential spreads along the membrane by changing the charge on the membrane capacitance and by flowing through opened channels Remember: 1. Membrane potential changes due to change in stored charge on membrane capacitor 2. Membrane conductance changes due to flow of ions through gated channels during graded and action potentials Action Potential Depolarization: When a stimulus opens the voltage gated channels, both Na+ and K+ channels get opened, however Na+ channels open faster and are responsible for the rising phase of action potential. The opening of Na+ channels will depolarize the membrane potential which will cause more Na+ channels to open The membrane then becomes overwhelmingly permeable to Na+ ions. Sodium ions diffuse into the cell down a concentration gradient. The entry of Na+ disturbs the resting potential and causes the inside of the cell to become more positive relative to the outside. Membrane potential shoots up to +30 mV Once open, however the Na+ channels spontaneously close by inactivation gate and they cannot open again until the membrane potential returns to resting membrane potential Action Potential Repolarization: Closing of Na+ channels causes the membrane potential to return to its resting level. In addition K+ channels start to open slowly and this facilitates the falling phase. Membrane becomes more permeable to K+ now than to Na+ Potassium rushes outside, membrane potential drops back down to -70mV. Lots of sodium inside, lots of potassium outside (opposite of the resting state at start) Hyperpolarization: Potassium channels doesn't close fast enough, so the membrane potential actually drops below the resting potential for a bit. The outside becomes comparatively more positive relative to the interior than that at the resting stage Action potential mechanism The rapid opening of voltage-gated Na+ channels allows rapid entry of Na+, moving membrane potential closer to the sodium equilibrium potential (+40 mv) A cell is “polarized” because its interior is more negative than its exterior. Repolarization is movement back toward the resting potential. The slower opening of voltage-gated K+ channels allows K+ exit, moving membrane potential closer to the potassium equilibrium potential (-90 mv) The rapid opening of voltage-gated Na+ channels explains the rapid-depolarization phase at the beginning of the action potential. The slower opening of voltage-gated K+ channels explains the repolarization and hyperpolarization phases that complete the action potential. Important Potentials • • • • Resting membrane potential is -70mV Depolarization peak is at +40mV Hyperpolarization peak is at -90mV Threshold potential is about -55mV • +40mV is close to the Na+ equilibrium potential • -90mV is the K+ equilibrium potential