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Sensory receptors Action potential 26.09.2012. Sensory receptors Special type of cells which can collect and transfer different informations from the environment. Function: „translation”, signal processing. Receptors (sensory organs) Nerve fibre Central nervous system stimulus: (physico-chemical) effect from the environment. → metabolic changes inside the cells induced by a stimulus. sensation: will involve the work of the CNS. Classification of sensory receptors Type of stimulus • light-photoreceptors • temperature-thermoreceptors • pressure-mechanoreceptors… Localisation of the receptors • head, ear… The origin of the information • Exteroceptor (informations form the environment) • Interoceptor (informations from the body) • Proprioceptor (position of the different parts of the body) Work of the receptors I. ! Receptors (sensory organs) stimulus Nerve fibre Central nervous system Local change of the receptor-potential thresholdpotential Action-potential frequency ~ strength of the stimulus „all or nothing” Work of the receptors II. Φ (Strength of the) stimulus Φthreshold Below the threshold level there is no sensation (no action potential). t Urec receptor Uthreshold t Uaction Action-potential 0 mV -70 mV t Nerve fibre The origin of the resting membrane potential Bernstein potassium hypothesis Nernst-equlibrium potential (electro-chemical potential) Donnan equlibrium: the membrane is impermeable for some components (e.g. intracellular proteins). Goldman equation: The membrane potential is the result of a „compromise” between the various equlibrium potentials, each weighted by the membrane permeability and absolute concentration of the ions. Equlibrium potential Nernst equation: What membrane potential (E) can compensate (balance) the concentration gradient (X1/X2). RT X 1 E= ln zF X 2 The inward and outward flows of the ions are balanced (net current = zero → equilibrium = stable, balanced, or unchanging system). Ionic concentrations inside and outside of a muscle cell battery Na+ : 120 mM Na+ : 20 mM K+ : 2.5 mM K+ : 139 mM Cl- : 120 mM Cl- : 3.8 mM [K+] ⇒ EmV = -58/1 log (139/2.5) = - 101.2 mV [Na+] ⇒ EmV = -58/1 log (20/120) = + 45.1 mV [Cl-] ⇒ EmV = -58/1 log (3.8/120) = + 86.9 mV = 30.8 mV EmV=-92mV Passive or leakage channels → a membrane-potential is not equal with any of the equlibrium potentials for the different ions EmV_K+ = -101.2 mV EmV_Na+ = +45.1 mV EmV_Cl- = +86.9 mV EmV= - 92mV → the ions are trying to move through the membrane ⇒ „leakage” „Leakage channels” K+ K+ Na+ + Na • Slow movement of the ions. • Has to be compensated somehow. Na-K ATPase The passive flux of Na+ and K+ (leakage) is balanced by the active work of Na-K pump → contribution to the membrane potential. 3 Na+ move out vs. 2 K+ move in (exchanger) ATP (energy source) is needed Na-K ATPase Sodium/potassium pump active transport → ATP hydrolysis - transport the ions against their concentration gradient (antiport transport ↔ symport). Antiport transport mechanism: a coupled movement of two compounds in opposite direction. Symport transport mechanism: a coupled movement of two compounds in the same direction. http://www.brookscole.com/chemistry_d/templates/student_resources/shared_resources/animations/ion_pump/ionpump.html Potassium channels ● Ca2+ sensitive potassium channels (KCa) ● Inwardly rectifying potassium channels (KIR) ● “Tandem pore domain potassium channel” – “leak channel” (K2p) ● Voltage-gated potassium channels (KV) Sensitive (dependent) to voltage changes in the membrane potential Islas LD, Sigworth FJ. Voltage sensitivity and gating charge in Shaker and Shab family potassium channels. J Gen Physiol. 1999 Nov;114(5):723-42. Sodium channels ● Ligand gated sodium channels ● Voltage gated (sensitive, dependent) sodium channels contains a voltage sensor Sensitive (dependent) to voltage changes in the membrane potential Planells-Cases R, Caprini M, Zhang J, Rockenstein EM, Rivera RR, Murre C, Masliah E, Montal M. Neuronal death and perinatal lethality in voltage-gated sodium channel alpha(II)-deficient Bernstein,J.(1902).Untersuchungen zur Thermodynamik der bioelektrischen Strome. Pflugers Arch.ges. Physiol. 92, 521–562. mice. Biophys J. 2000 Jun;78(6):2878-91. Sodium channels ● Ligand gated sodium channels ● Voltage gated (sensitive, dependent) sodium channels contains a voltage sensor Sensitive (dependent) to voltage changes in the membrane potential activation gate (extracellular side) inactivation gate (intracellular side) Planells-Cases R, Caprini M, Zhang J, Rockenstein EM, Rivera RR, Murre C, Masliah E, Montal M. Neuronal death and perinatal lethality in voltage-gated sodium channel alpha(II)-deficient Bernstein,J.(1902).Untersuchungen zur Thermodynamik der bioelektrischen Strome. Pflugers Arch.ges. Physiol. 92, 521–562. mice. Biophys J. 2000 Jun;78(6):2878-91. Action potential Action potential: a momentary reversal of membrane potential (- 65mV to + 40 mV) that will be followed by the restoration of the original membrane potential after a certain time period (1-400ms). Action potentials happens in different phases (depolarisation and repolarisation). Action potentials are triggered by the depolarization of the membrane if it can reach a critical value (voltage threshold). Action potentials are all or none phenomena any stimulation above the voltage threshold results in the same action potential response. any stimulation below the voltage threshold will not result action potential response. Membrane potential (mV) Action potential (nerve cell) overshoot ~+40 0 Falling phase Rising phase Voltage threshold ~-50 Resting potential ~-70 undershoot Stimulus 0 1 2 time (ms) 3 4 Equlibrium situation NaV channels: activation gate closed inactivation gate open Membrane potential (mV) Resting phase overshoot ~+40 0 Rising phase Falling phase Voltage threshold ~-50 Resting potential ~-70 Stimulus 0 1 undershoot 2 time (ms) 3 The voltage-gated sodium channels will open-up if the voltage threshold reached by a stimulus NaV channels: activation gate open inactivation gate open → Na+ will move into the cell → the inner surface of the cell will be positively charged Membrane potential (mV) Rising phase (depolarization) overshoot ~+40 0 Rising phase Falling phase Voltage threshold ~-50 Resting potential ~-70 Stimulus 0 1 undershoot 2 time (ms) 3 The movement of the Na+ will slow down EmV_Na+ = +45.1 mV (Nernst –equlibrium potential) Na+ channels will start to form an inactive conformation K+ channels are starting to open-up Membrane potential (mV) Overshoot overshoot ~+40 0 Rising phase Falling phase Voltage threshold ~-50 Resting potential ~-70 Stimulus 0 1 undershoot 2 time (ms) 3 All the voltage gated K+ channels are open K+ move out from the cell NaV channels: activation gate open inactivation gate closed → refractory period Membrane potential (mV) Falling phase (repolarization) overshoot ~+40 0 Rising phase Falling phase Voltage threshold ~-50 Resting potential ~-70 Stimulus 0 1 undershoot 2 time (ms) 3 The movement of the K+ ions will slow down EmV_K+ = -101.2 mV (Nernst-equlibrium potential) The K+ channels will get into a closed conformation The numerous and slowly inactivating K+ channels will cause some hyperpolarisation Membrane potential (mV) Undershoot (hyperpolarization) overshoot ~+40 0 Rising phase Falling phase Voltage threshold ~-50 Resting potential ~-70 Stimulus 0 1 undershoot 2 time (ms) 3 Membrane potential (mV) Resting phase overshoot ~+40 0 Rising phase Falling phase Voltage threshold ~-50 Resting potential ~-70 Stimulus 0 1 undershoot 2 time (ms) 3 Recovery after the AP Intracellular ion concentrations change only a small amount with each AP (0.0001% - 1%). Na+/K+ ATPases will slowly restore the original ion concentrations. If the Na/K ATPases of a squid giant axon is poisoned, it can still generate 100,000 impulses while the internal sodium concentration is increased only by 10%. Propagation of the action potential Action potentials can propagate: the AP at one site on the membrane can itself initiate an AP at a neighboring region. Unidirectional movement because of the refractory period – inactivated Na+ channels. The electrical signal can propagate down the axon without a decrease in amplitude. Refractory period The cell is resistant to a stimulus. It is hard to get a response (action potential). Membrane potential (mV) Absolute refractory period: The formation of a new AP is totaly blocked Relative refractory period: larger depolarisation is needed than the threshold to initialize an AP overshoot ~+40 0 Rising phase Falling phase Voltage threshold ~-50 Resting potential ~-70 0 Stimulus 1 time (ms) undershoot 2 3 4 Action potential in the cardiac muscle cells K+ and Cl- ion movement Membrane potential plateau phase Equlibrium between the Ca2+ release and the outward movement of the K+ ions ~+35mV depolarisation •Na+ moving into the cell K+ ion moving out from the cell ~-90mV 0 ms 350 ms Resting membrane potential Plateau: can help to direct the flow of the APs. time (ms) Resting membrane potential The end!!!