LECTURE 2 Basic Electrical Properties of Neurons I. Basic neural system II. Membrane properties III. Synapses IV. Action potential generation Two classes of cells • Nerve cells (neurons,神经细胞或神经元) − For electrical signaling − The human brain has 1011 (1000 亿) neurons • Glial cells (神经胶质细胞) − Not for electrical signaling − 10 to 50 times more glial cells than neurons − Play an essential role in brain metabolism − Support neurons − Cover neurons with myelin − Clean up debris Neuron structure • cell body – contains nucleus (核)& organelles (细胞器) • dendrite – conducts signals to cell body • axon – long fibers – specialized for electrical conduction Structure of a typical neuron Dendrite Axon Terminal Node of Ranvier Cell Body Axon Nucleus Schwann Cell Myelin Sheath Differences between neurons and other cells 1. Neurons have specialized extensions called dendrites and axons. Dendrites bring information to the cell body and axons take information away from the cell body 2. Neurons communicate with each other through an electrochemical process 3. Neurons contain some specialized structures (for example, synapses) and chemicals (for example, neurotransmitters) Classification of neurons by function Sensory (or afferent) neurons (感觉神经元) They send information from sensory receptors (e.g., in skin, eyes, nose, tongue, ears) TOWARD the central nervous system Interneurons(中间神经元) They send information between sensory neurons and motor neurons. Most interneurons are located in the central nervous system Motor (or efferent) neurons(运动神经元) They send information AWAY from the central nervous system to muscles or glands By the number of extensions extending from the neuron's cell body Pseudounipolar cells-伪单极神经元 One process extends centrally toward the spinal cord, the other extends toward the skin or muscle ( dorsal root ganglion cells) Bipolar neurons - 双极神经元 Two processes extending from the cell body. (retinal cells, olfactory epithelium cells) Multipolar neurons - 多极神经元 Many processes but has only one axon. (spinal motor neurons, pyramidal neurons, Purkinje cells) By neuron‘s shape 105 synaptic inputs thousands of synaptic inputs (103) A cortical pyramidal cell A Purkinje cell of the cerebellum A stellate cell of the cerebral cortex (magnified about 150 fold) (Drawings from Cajal 1911) I. Basic neural system II. Membrane properties III. Synapses IV. Action potential generation The soma of a neuron: 4 - 100 μm in diameter Nucleus • 1 μm3: 1010 water molecules, 108 ions, 107 small molecules such as amino acids and nucleotides, and 105 proteins • In water, the molecules Na+, K+, Cl- and Ca2+ are in ionic form A schematic diagram of a section of the lipid bilayer 3 to 4 nm ~10 nm long (Dayan and Abbott 2001) Basic points: physiological specializations • A wide variety of membrane-spanning ion channels (Na+, K+, Ca2+, and Cl−) • Ion channels control the flow of ions across the cell membrane (voltage-gated, ligand-gated, and others) • This type of membrane is called semipermeable I. Basic neural system II. Membrane properties III. Synapses IV. Action potential generation Diagram of a synapse (Kandel et al. 1991) Signal transmission at synapse Electrical synapses At gap junctions, cells approach within about 3.5 nm of each other, rather than the 20 to 40 nm distance that separates cells at chemical synapses Postsynaptic potential in electrical synapses is not caused by the opening of ion channels by chemical transmitters, but by direct electrical coupling between both neurons Electrical synapses are therefore faster and more reliable than chemical synapses. (Most of time, biderectional) 电突触主要存在于蚯蚓、虾、软体动物等无脊椎动物 I. Basic neural system II. Membrane properties III. Synapses IV. Action potential generation A rough estimation of membrane potential qVT kT kT VT q kT: The thermal energy of an ion K: Boltzmann constant q: the charge of a single proton VT: 24 ~27 mV Membrane potentials: about -3 to +2 times VT Intracellular Resistance the intracellular resistivity Membrane potentials measured at different places within a neuron can take different values For neurons with electrotonic compactness, we have … Membrane Capacitance and Resistance membrane resistance 1. membrane capacitance membrane conductance: g = 1/ rm 2. membrane time constant: τm = RmCm = rmcm (10 and 100 ms) (the basic time scale for changes in the membrane potential) The restriction in measuring membrane resistance Why is the restriction to small currents and small voltage needed here? Equilibrium potentials -+ [inside] + + + + + + + + [outside] + + + + + + + + + + + + + Concentration gradient Voltage gradient zqE zE [outside] [inside] exp( ) [inside] exp( ) kT VT + + Nernst equation VT [outside] E ln( ) z [inside] Examples: Sodium ions: o = 60 mM, I = 440mM Potassium ions: o = 400 mM, i=20 mM E = 27·ln(440/60) = 54mV E = 27·ln(20/400) = -80mV Chloride ions: E = -65mV (near the resting potential of many neurons) Calcium ions: E = 130mV Equilibrium and reversal potentials Equilibrium potentials (The Nernst equation applies when the channels allow only one type of ion to pass through them) Some channels are not so selective, and in this case the potential E is estimated by the Goldman equation Reversal potentials takes a value intermediate between the equilibrium potentials of the individual ion types that it conducts The Membrane Current • For one ion type with reversal potential E: i g (V E ) • For several ions through different channels: i gi (V Ei ) i Leakage current the resting potential i g L (V EL ) Leakage conductance: a passive conductance All of the time-independent contributions to the membrane current can be lumped together into a single leakage term. For example, the currents carried by ion pumps that maintain the concentration gradients that make equilibrium potentials nonzero Action potential An action potential: a roughly 100 mV fluctuation in the electrical potential across the cell membrane that lasts for about 1ms Depolarization and hyperpolarization Absolute refractory period: a few milliseconds just after an action potential Relative refractory period: lasting up to tens of milliseconds after a spike Subthreshold potential fluctuations Subthreshold potential fluctuations are severely attenuated over distances of 1 mm or less Three simulated recordings from a neuron (Dayan and Abbott 2001) 作业及思考题 1. 胶质细胞主要类型和功能. 2. 如何估计膜电位数量级? 3. 说出钠、钾和氯离子通道平衡电位的数量级。 4. 如果用电极分别在锥体细胞的胞体内、胞外和距 胞体一定距离的轴突内记录动作电位串,结果有 何异同?