Lecture 07, 13 Sept 2005 Chapters 12 and 13. Vertebrate Physiology ECOL 437 (aka MCB 437, VetSci 437) University of Arizona Fall 2005
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1 Lecture 07, 13 Sept 2005 Chapters 12 and 13 Vertebrate Physiology ECOL 437 (aka MCB 437, VetSci 437) University of Arizona Fall 2005 instr: Kevin Bonine t.a.: Kristen Potter Vertebrate Physiology 437 Chapter Synapses, Neurotransmitters Chapter Sensory Processes/Systems 1 2 Hill et al. 2004, Fig Hill et al. 2004, Fig different domains 4 identical subunits Voltage-gated channel superfamily 3 4 Frequency and number! Proposed Evolution Voltage-gated Channels 5 Hill et al. 2004, pg. 301 Silverthorn nd ed. Human Physiology. Prentice 6Hall 1
2 SYNAPSES -communication between neurons or between neuron and effector organ Electrical Synapse (rapid) - direct ionic coupling via gap junctions -examples in retina, CNS, smooth muscle, cardiac muscle, etc. 1-electrical (rapid) 2-chemical( fast or slow) 6-9 Randall et al In postsynaptic neuron: 1. De- or hyper-polarize 2. Change # ion channels in membrane 6-22 Randall et al Randall et al Alter rate of ion channel activity 4. Modify sensitivity to activation signals 7 gap junctions 8 Chemical Electrical Agonist (mimics) (e.g., heroin mimics natural opiates) vs. Antagonist (blocks) (e.g., curare blocks ACh reception) Hill et al. 2004, Fig Chemical (neurotransmitter) 20-30nm apart Chemical synapses Electrical (gap junction, connexons) 3nm apart Role of Ca ++ 1 amplify 2 excitatory or inhibitory 3 ~one-way 4 modifiable ionotropic metabotropic Hill et al. 2004, Fig 12.2,3 11 Hill et al. 2004, Fig
3 Silverthorn nd ed. Human Physiology. Prentice Hall Fast Slow 13 Hill et al Postsynaptic Neurotransmitter Effects NT role depends primarily on receptor characteristics on postsynaptic neuron 1. Fast and direct 2. Slow and indirect e.g., ACh receptors 1. Nicotinic (muscles, autonomic/sympathetic NS) 2. Muscarinic (parasympathetic, indirect) Norepi G-protein camp 2 nd messenger Phosphorylation (kinases) (amplification) 15 Hill et al. 2004, Fig Postsynaptic Neurotransmitter Effects e.g., indirect, metabotropic muscarinic ACh receptors acting to reduce heart cell excitability Hill et al. 2004, Fig Randall et al
4 Postsynaptic Neurotransmitter Effects 1. Fast and direct e.g., fast nicotinic ACh receptors Nicotinic ACh receptor 6-32 Randall et al ACh binds alpha subunits 6-34 Randall 19 et al Hill et al. 2004, Fig Neuromuscular Junction Quantal packets (~5,000 Ach/vesicle) Hill et al. 2004, Fig Neurotransmitters: 1. small-molecule neurotransmitters (often made in axon terminals) 2. neuroactive peptides (often made in soma and shipped down axon) (10%) (1%) (abundant and widespread) Nematodes use a lot of the same neurotransmitters. (IPSP) (IPSP)
5 Synaptic Plasticity Longterm Potentation Change synaptic efficacy Alter rate of NT production and release Learning and Memory Facilitation vs. antifacilitation/depression Calcium-dependent -Research on-going Hippocampus Learning and Memory Neurons that fire together wire together NMDA glutamate receptors Hill et al. 2004, Fig Vertebrate Physiology 437 Chapter Sensory Processes/Systems Doogie Mice? Sensing the Environment Sensory Reception -Environment -Within body Integrated and Processed by NS Properties of Receptor Cells Sensory Modality Modalities include: vision, hearing, touch, taste, smell, chemical, thermal, proprioceptors Sensory Receptors send signals to brain so perceive sensations Sensory Receptor cells often organized into Qualities within each modality e.g., Red or yellow; High or low-pitched 7-1 Randall et al organs 7-1 Randall et 29 al
6 Properties of Receptor Cells 7-2 Randall et al Receptor Cells - Specialized - Selective for energy type and modality -either is a neuron or -Synapses immediately on a neuron (1 afferent neuron to CNS) Stimulus modifies conformation of receptor Properties of Receptor Cells Mechanisms and Molecules Transduction= Stimulus energy converted to nerve impulse Sensory Adaptation - orders of magnitude different stimulus strength Example Mechanoreceptors (touch) 1- Proteins respond to membrane distortion 2- Signal often amplified 3- Ion channels opened directly or indirectly 4- Current flows across membrane (often Na + ) 5-Vm changes (aka receptor potential changes) 6- AP sent or NT released causing AP 33 - often controlled via Ca++ availability -local controlor feedback from CNS Type of stimulus received depends on where in CNS (~brain) AP arrives (LABELED LINES). Rub eyes and see light! Intensity signalled by frequency of APs, but 34 Stimulus Intensity and Dynamic Range From lowest threshold, to upper limit imposed by refractory period: Note log axis Dynamic Range Shifting range of appropriate AP frequency Detectable light intensity varies over 9 orders magnitude Detectable sound intensity varies over 12 orders magnitude Range Fractionation - Function of sensory adaptation - Also recruit receptors with different tunage or sensitivity (e.g., rods and cones in eye) 7-7 Randall et al
7 Sensory Adaptation Possibilities: 1. Receptor cell mechanical properties may filter 2. Receptor cells may be depleted (e.g., visual pigments; need to be regenerated) 3. Enzyme cascade (during amplification) may be inhibited by (intermediate) product 4. Electrical properties change b/c [Ca ++ ] 5. Accommodation of spike initiating zone 6. Sensory adaptation in downstream neurons (CNS) 37 Hill et al. 2004, Fig Sensory Adaptation; Pacinian Corpuscle - Touch Example 7-5 Randall et al Movement of Oil between layers is what triggers APs Signal changes in pressure, not steady pressure Randall et al Mechanisms and Molecules Lots of Evolutionarily Conserved Elements e.g., 7 transmembrane helices and G-protein intermediate Mechanisms and Molecules Enzymatic Cascade to amplify Threshold of Detection e.g., 1 photon or hair cell movement of H diam. Sour (ph; H+) and salt (Na+) move directly no amplification e.g., Vision, olfaction, sweet and bitter taste (also muscarinic ACh receptors and many hormone receptors) To measure quality need many receptors grouped into organ; different tunage (e.g, wavelength of light or frequency of sound) 7-3 Randall et al
8 Enhancing Sensitivity - Spontaneous basal activity 7-12 Randall et al Tonic vs. Phasic receptors fast-adapting - Constant rate of APs - Directionality if or AP frequency Slow-adapting 5-19 Randall et al Accommodation Enhancing Sensitivity - Efferent Control e.g., stretch receptors in muscle control length so can perceive stretch - Feedback Inhibition Auto (helps keep in dynamic range) vs. Lateral Randall et al Enhancing Receptor Sensitivity - Lateral Inhibition 7-14 Randall et al Vision FOCUS - light is focused by cornea to create an image on the retina 7-34 Randall et al refraction by cornea (85%) and by lens (15%) e.g., improve touch sensitivity and visual acuity (edges especially) -alter focal length by altering shape and curvature of lens (zonular fibers and ciliary muscle sphincter ) - binocular convergence (both eyes on same part of retina) 47 LIGHT INTENSITY - pupil for variable aperture via iris and radial muscle 48 8
9 10-27 Silverthorn Randall et al distant Out of focus close Vision ~ANATOMY - sclera white tough outer layer - choroid lots of blood vessels - pigment layer with photoreceptors - fovea where highest acuity and highest # cones -(visual streak?) TRANSDUCTION - photoreceptors (rods and cones) -Transduce photons (light) into electrical signal Silverthorn rhodopsins (visual pigments) opsin (7-transmembrane lipoprotein) plus retinal (absorbs photon) 52 Vision Receptor Cells Silverthorn 2001 Rods and Cones -Dim light, low resolution -Bright light, high resolution Rods and Cones 7-38 Randall et al
10 7-39 Randall et al Rhodopsins (visual pigments) -located in stacked lamellae Membranes hyperpolarize in response to light Na + dark current When light hits, the Na + current into the cell is stopped and membrane hyperpolarizes stopping release of 55 NT Bleaching of retinal photoreceptors Expectation after 15 seconds? Photoreceptors called cones respond to particular wavelengths of light. Their response involves bleaching of their responsive pigment, so that for some seconds they are unable to respond again. 56 Bleaching of retinal photoreceptors Bleaching of retinal photoreceptors Expectation after 15 seconds? Expectation after 15 seconds? Photoreceptors called cones respond to particular wavelengths of light. Their response involves bleaching of their responsive pigment, so that for some seconds they are unable to respond again. 57 Photoreceptors called cones respond to particular wavelengths of light. Their response involves bleaching of their responsive pigment, so that for some seconds they are unable to respond again. 58 Rhodopsin mechanism: cis-trans isomerization of retinal molecule 7-44 Randall et al Activated retinal changes conformation of opsin molecule (opsin and retinal separate) and initiates transduction 7-43 Randall et al activated Changes conformation of opsin molecule and therefore initiates 59 transduction 7-45 Randall et al G-protein amplification Need to reconstitute the rhodopsin 60 (night blindness) 10
11 Rod and Cone details Action spectrum (where absorb light) 3 (e.g., humans, fish)-5 (e.g,. birds) different photopigments Different opsins, same retinal Porphyropsins (different retinal) seem better than rhodopsins in freshwater Sensitivity vs. Acuity Silverthorn Physiology Players Theatre -2 competing casts -Judge(s) -accuracy -enthusiasm 7-49 Randall et al Actors: 1. Photon 4. Transducin 7. Ion channel 2. Retinal 5. PDE 8. Cation (Na + ) 3. Opsin 6. cgmp Act I Photon enters stage right. Other players assembled within or near membrane. photo transduction... Dark current reduced as curtain closes. Red vs. Green opsin 62 11
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