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CLASS #1: Tues, 9 Jan 2007 PSB 2000 -01, -02, -03, -04 Spring 2007 INTRODUCTION: THE BRAIN AND BEHAVIOR I. OVERVIEW ● Attendance ● General information about mechanics of the class; syllabus ● What is “Neuroscience?” ● An example: how your instructor (Dr. Berkley) became a neuroscientist, and what she studies. II. THE NERVOUS SYSTEM A. THE MIND-BODY DILEMMA DUALISM MONISM mind body mind body B. IMPORTANT PHILOSOPHERS (besides ourselves) ● Hippocrates: 460-379 BC---brain is seat of intelligence. ● Aristotle: 384-322 BC---heart = mind/soul; brain serves as radiator to cool overheated blood. Monism ● Galen: 130-200AD---brain secretes fluids conveyed by nerves to the body—behaves like a gland. ● Versalius: 1514-1564 ---mind/soul is the nervous system. Dualism ● Descartes: 1596-1650: MIND pineal gland brain body. C. FUNCTION: The main functions of the nervous system are to organize and control an individual’s appropriate interactions with her/his environment. Examples: Neural input capabilities are appropriate to lifestyle of species: e.g., rodents can detect ultrasounds (‘bat detector’) Impact of abnormal neural input can have social consequences: e.g., your instructor’s hearing aids The nervous system is plastic. It learns and processes information so that actions are appropriate to circumstances: e.g., pain—a motivating perception: horse breaking leg in race; deer running from a lion; humans in the ER D. EVOLUTION, THE GENETIC REVOLUTION AND ITS SIGNIFICANCE FOR NEUROSCIENCE (if time)—Module 1.2 in your book. PSB 2000 Class #2: Thurs 11 Jan 2007 cCELLS OF THE NERVOUS SYSTEMm I. COMPONENTS OF A CELL ● Nucleus: chromosomes (23 pairs-humans)-DNA/genes; nucleolus (makes rRNA-ribosomal RNA); nuclear membrane ● Cytoplasm: mitochondria (energy), ribosomes (synthesis of protein); endoplasmic reticulum (rough; smooth); golgi complex, lysosomes. ● Plasma membrane II. PROTEIN SYNTHESIS ● composition of DNA: double stranded molecule, with sequences of any of 4 bases attached in various orders to a skeleton made of a sugar (deoxyribose) and a phosphate: Guanine (G), Cytosine (C), Adenine (A), **Thymine (T) ● compsition of RNA: single stranded molecule, with 4 bases on a skeleton made of a sugar (ribose) and a phophate: G, C, A, and **Uracil (U) ● DNA mRNA* G C Genetic info in nucleus is transcribed to mRNA. The mRNA* moves into the C G cytoplasm and onto a ribosome, where A U the mRNA is translated into a protein**. T A *messenger’ RNA; **each sequence of 3 RNA bases codes for one AMINO ACID (example: G-C-G = arginine). Proteins are composed of combinations of amino acids. III. NEURONS ● dendrites, soma, axon hillock, axon, myelin sheath (node, internode), terminal branches, terminal boutons (presynaptic swellings) ● variations in the arrangements of these elements ● synapses: specialized junctions between two neurons ● synapse composition: presynaptic swelling (vesicles, mitochondria, presynaptic membrane); synaptic cleft; postsynaptic membrance and thickening ● synapse types axodendritic; axosomatic; dendrodentritic; axoaxonic IV. CONNECTIONS BETWEEN NEURONS (“WIRING”): divergence, convergence and plasticity IV. GLIAL CELLS ● astrocytes: “nerve glue”, inactivate neurotransmitters, nutrition, cleanup, potassium (K+) buffer ●oligodendrocytes myelin (CNS); Schwann cells myelin (PNS) ●microglia (associated with injury) V. BLOOD-BRAIN BARRIER (if time) PSB 2000 Class #3: Tues, 16 Jan 2007 ELECTRICAL ACTIVITY: RESTING AND ACTION POTENTIALS Communication between neurons depends on active (energy-requiring) properties of the neuron’s plasma membrane. I. MEMBRANE CHANNELS (FOR IONS**): 3 TYPES TYPE LOCATION leakage voltage-gated Mostly axon, also everywhere soma, dendrites ligand-gated* Dendrites and soma (not axon), terminal boutons Type of potential for Resting Action potential, which the chanel is sometimes synaptic Synaptic potentials potential important potentials *Ligand-gated types: ● ionotropic (just ions) ● metabotropic (involves G-proteins) + + + II. RESTING MEMBRANE POTENTIAL: steady state voltage across + + membrane; inside of cell negative to outside; produced by: ● an unequal distribution of ions OUTSIDE OUTSIDE INSIDE between inside and outside of membrane: Na+ K+ ● semipermeability of membrane Cl● forces acting on movement of ions in solution K+ A- Na+ Cl-- Na+ Cl- -concentration gradient; electrostatic (like forces repel; opposite attract) ● sodium-potassium pump (needs energy): pumps K+ in and Na+ out III. ACTION POTENTIAL: A rapid, all-or-none, reversible change in membrane potential of ~1msec duration (duration can vary) ● sequential opening , then closing of voltage-gated ion channels: 1st: Na+ chan. open (Na+ rushes in—causing depolarization; 2nd: K+ chan. open (K+ moves out), causing repolarization and hyperpolarization. ● usually initiated at axon hillock (threshold is lowest there) ● propagates from there down to boutons (via passive, cable properties of axon and saltatory conduction-if axon myelinated) ● refractory periods (absolute and relative) prevent it going in both directions constantly; absol ref per + duration of APsets top frequency. ● conduction velocity = speed of conduction of action potential down the axon; depends on diameter of axon and how it is myelinated; varies from ~0.3 meters/sec to ~120 meters/sec! ** IONS are electrically-charged molecules: cations are positively charged—such as potassium (K+), sodium (Na+), calcium (Ca++); anions are negatively charged—such as chloride (Cl-), some large proteins (A-) K+ PSB 2000 Class #4: Thurs, 18 Jan 2007 SYNAPTIC POTENTIALS, TRANSMITTERS, DRUG ACTIONS I. WHAT HAPPENS AT A SYNAPSE? A. Transmitter release: AP activates voltage-gated Ca++ channels in the presynaptic membrane, causing vesicles to fuse with presynaptic membrane and release their transmitter into synaptic cleft. B. Transmitter recognition: Receptors in the post-synaptic membrane recognize and bind the transmitter. C. Transmitter inactivation: 4 mechanisms of inactivation: ● reuptake; ● uptake by post-synaptic neuron; ● glial uptake; ● enzyme deactivation. D. Actions at post-synaptic membrane: If receptors in post-synaptic membrane recognize the transmitter (=ligand-gated channels), the membrane’s conformation changes, causing a change in ion movement across the post-synaptic membrane and therefore a change in voltage across the membrane at that point. Depolarization = excitatory post-synptic potential (EPSP); hyperpolarization = inhibitory post-synaptic potential (IPSP). II. Summation of EPSPs and IPSPs across the membrane of the post-synaptic neuron: EPSPs and IPSPs summate in time and in space. If the summation produces enough depolarization to reach the neuron’s threshold anywhere on the membrane, then an action potential will occur there and spread over the whole membrane. The threshold is usually lowest at the axon hillock. A. Types of effects of neurotransmitters on the post-synap. membrane: ● ionotropic: direct action on ion channels ● metabotropic: indirect, either via a G-protein, or a G-protein and a 2nd messenger B. Types of neurotransmitters: ● acetylcholine ● monoamines: --indolamine: serotonin --catecholamines: dopamine, norepinephrine, epinephrine ● amino acids: glutamate, GABA, glycine, many others ● peptides (contain amine groups-NH2): endorphins, substance P, oxytocin, neuropeptide Y, vasopressin, many others ● purines: adeniosine, ATP, others ● gases: nitric oxide, carbon dioxide (NO, CO) ● others: cannabinoids, others… III. DRUG ACTIONS: ● Concepts: antagonist, agonist, affinity, efficacy, dependency, addiction ● Drugs can influence synaptic activity in a huge number of ways, by acting on any of the processes associated with neurotransmitter action, such as: ●synthesis of transmitters, their receptors and their inactivators; ●release of transmitters; ●inactivation of neurotransmitters,● the recognition of neurotransmitters, etc… PSB 2000 Class #5: Tues, 23 Jan 2007 GROSS ANATOMY-part 1Y I. DEVELOPMENT OF THE NERVOUS SYSTEM A. Origin: The early embryo differentiates into groups of cells that are the origin of ectodermal tissues, mesodermal tissues and endodermal tissues. The nervous system originates from ectodermal cells. B. Process: A thin sheet of cells forms on the dorsal* surface. This sheet curls up into a tube-like structure along the length of the developing body. The most dorsal part of this tube (where the two ends of the sheet have joined) is composed of neural crest cells. The rest of the tube is composed of neural tube cells. The neural crest cells proliferate and migrate out into the rest of the body, thereby forming the PERIPHERAL NERVOUS SYSTEM. The neural tube cells proliferate to enlarge the tube, thereby forming the CENTRAL NERVOUS SYSTEM, whose rostral* part is the BRAIN, and whose caudal* part is the SPINAL CORD. II. PERIPHERAL NERVOUS SYSTEM Derived from neural crest cells. Consists mainly of the nerves (groups of axons) and ganglia (groups of nerve cell bodies) that lie outside the spinal vertebrae and skull. A. Cranial nerves: 12 pairs MEMORIZE THESE! I = olfactory II = optic “On Old Olympics Towering see p. 88, III = oculomotor Tops A Finn And German Table 4.4 in IV = trochlear Viewed Some (or A) Hat(s)” your text V = trigeminal VI = abducens VII = facial VIII = auditory (or statoacoustic) IX = Glossopharyngeal X = vagus XI = spinal accessory (or accessory) XII = hypoglossal B. Spinal nerves: 31 pairs (humans): ● 8 Cervical = C1-C8 ● 5 Sacral = S1-S5) ● 12 Thoracic = T1-T12 ● 1 Coccygeal = Co1 ● 5 Lumbar = L1-L5) C. Divisions: ● Somatic: those nerves that innervate skin and muscles ● Autonomic: those nerves that innervates internal organs. It has 2 subdivisions --Sympathetic --Parasympathetic III. SPINAL CORD A. Segmentation: There are 31 segments that are associated with each of the 31 pairs of spinal nerves (named the same way; e.g., T12 segment receives input from/ sends output through T12 spinal nerves. B. Organization: “Grey matter” surrounded by “white matter.” Through the middle runs a “central canal” that contains cerebrospinal fluid (CSF). Grey matter is composed of neuronal soma and synapses. White matter is composed of axon tracts heading rostrally to the brain or descending from the brain to the spinal cord. C. Association with spinal nerves: Each spinal nerve is divided in 2 parts inside the spinal vertebrae. One division enters the dorsal* aspect of the spinal cord. It is called the “dorsal root,” and brings information into the spinal cord (afferent*, sensory input). The other division exits from the ventral* aspect of the spinal cord (“ventral root”), bringing information out to the body (efferent*, motor output). This arrangement, discovered by Bell and Magendie (thus the “Bell-Magendie Law”) creates a situation in which the dorsal part of the spinal cord has mainly sensory functions, whereas the ventral part has mainly motor functions. The cell soma for the the nerve fibers of the dorsal (sensory) division of each spinal nerve are located outside the spinal cord in ganglia (“dorsal root ganglion”). These ganglia have the same name as the spinal nerve (e.g., the T12 dorsal root ganglion). The cell soma of the ventral root axons are located in the ventral part of the spinal cord--motoneurons and “preganglionic autonomic neurons.” (Preganglionic neurons will discussed later.) *Learn the following orientation terms: anterior, posterior, rostral, caudal, dorsal, ventral, medial, lateral, proximal, distal, ipsilateral, contralateral. (p. 83, Table 4.1 in your text) *Learn the following planes of section through the spinal cord and brain: coronal (frontal) section, saggital section, horizontal section, transverse section (cross section). – (p. 83, Figure 4.2 in your text) PSB 2000 Class #6: Thurs, 25 Jan 2007 (Class #7 on Tues 30 Jam 2007 is EXAM # 1) GROSS ANATOMY-part 2Y I. SUBDIVISIONS OF THE BRAIN MIDBRAIN FOREBRAIN telencephalon diencephalon mesencephal basal ganglia internal capsule paleocortex neocortex epithalamus thalamus subthalamus hypothalamus = midbrain 1st 2nd tectum tegmentum 3rd, 4th HINDBRAIN cerebellum pons medulla --associated with the cerebellum via the “cerebellar peduncles” Dorsal part is sensory Ventral part is motor 4th, 5th, 6th, 7th, 8th 9th, 10th, 11th 12th grey areas = “brainstem” II. ASSOCIATION WITH CRANIAL NERVES Each of the subdivisions of the brainstem receives input from or sends output through one or more of the cranial nerves, a situation which has functional implications for that subdivision. Memorizing the functions of the cranial nerves will help you learn this (Table 4.4, p. 88). Here is some room for notes from class lectures: MEDULLA PONS MIDBRAIN DIENCEPHALON III. TELENCEPHALON A. Basal ganglia: embedded in the internal capsule-motor functions B. Internal capsule: axons conveying information to and from the cerebral cortex C. Cerebral cortex: folded up to increase surface area (sulcus or fissure=base; gyrus=top of the fold). There are two main types of cerebral cortex: ● paleocortex: 3-4 layers: “limbic cortex”: hippocampus, amygdala, cingulate gyrusclosely associated with the hypothalamus in the diencephalon ● neocortex: 6 layers: 4 lobes: FRONTAL Primary motor ctx; eye movement; “Executive” functions; Speech:Broca’s area-left side. Olfaction: associated with paleocortex PARIETAL OCCIPITAL TEMPORAL Primary somatosensory cortex Vision Audition; Memory (buried in hippocampus); Speech (Wernike’s area-left side) PSB 2000 Class #6: Thurs, 25 Jan 2007 (Class #7 on Tues 30 Jam 2007 is EXAM # 1) Here are three pages of diagrams that will be useful during lectures in Classes #6 & 7, GROSS ANATOMY, Part 2: PSB 2000 Class #6: Thurs, 25 Jan 2007 (Class #7 on Tues 30 Jam 2007 is EXAM # 1) PSB 2000 Class #6: Thurs, 25 Jan 2007 (Class #7 on Tues 30 Jam 2007 is EXAM # 1) PSB 2000 Class #8: Thurs, 1 Feb 2007 (Class #7 on Tues, 30 Jan was Exam # 1) EETHICS AND METHODS OF NEUROSCIENCE RESEARCHH I. FACTORS ASSOCIATED WITH THE USE OF ANIMALS IN RESEARCH A. Respect for the welfare of animals is: ● very important ● strictly regulated—by federal law, Nat Institutes of Health (NIH) Also: granting organizations, professional research societies, scientific journals) ● strictly monitored--by local Animal Care and Use Committee (ACUC) and federal government (US Dept. of Agriculture), ● strictly enforced—ACUC, NIH, USDA ACUC: minimal membership B. Before research can be done: ● training or evidence of training required ● literature review must be done ● ACUC must give approval ● for funding: government (NIH, etc) review #1 ● funding: government review #2 1. community ‘lay’ person (non- scientist, non-academic) 2. veterinarian 3. scientist who works with animals 4. another academic who does not work with animals C. During research: ● continued training ● site visits (often unannounced): local ACUC committee; USDA; sometimes NIH D. Publishing results: All journals have ethical standards that must be met—usually 3 reviews! II. METHODS USED IN NEUROSCIENCE RESEARCH (both humans and nonhuman animals)---All methods have limitations; best approach is combinations. A. Anatomical: (What is its structure? What does it look like?)—subcellular to whole brain): 1. Terms: ●cytoarchitecture (how cells are grouped) special stains, radioactivity, ●histochemistry (makes molecules visible) fluorescent dyes 2. Techniques: electron microscopy; light microscopy; confocal microscopy; CAT scans; MRI 3. Important aspects: ● orientation (3 planes, brain atlas, 3-D maps); ● stereotaxic device 4. “Neuronavigation” and Tracing neural connections B. Functional: 1. Genetisc, Molecular Biology; Neurochemistry 2. Electrophysiological: (how electrical activity relates to function) a. direct devices: ●oscilloscope (measures voltage changes with time); ●electrodes: microelectrode—single neuron, ‘patch’ of membrane; macroelectrode---regional activity; EEG MEG (magnetoencephalography; whole brain) b. indirect: ●PET--positron emission tomography—decay of injected or inhaled radioactive elements (133Xenon-inhaled; 11C, 18F, 15O-injected); ●fMRI--functional MRI-- magnetic properties of hemoglobin after O2 release 3. Behavioral: a. Lesion/Ablation: destroy an area of brain, then measure behavioral change b. Stimulation: artifically activate an area, then measure behavior elicited c. experimentally, both a and b have serious interpretative problems for both positive and negative results; controls are therefore very important. PSB 2000 Class #9: Tues, 6 Feb 2007 SENSORY MECHANISMS I. GENERAL ASPECTS OF SENSATION AND PERCEPTION A. Transduction: the process by which sensory receptors change stimulus energy action potentials B. Receptors: types (mechanical, chemical thermal); sources (telereceptors-distance; exteroceptors-on body; proprioceptors-muscles, joints, vestibular apparatus; interoceptors-internal organs) C. Adequate stimulus: That stimulus to which a particular sensory receptor is normally AND most efficiently responsive. C. Information necessary for perception: Modality, Intensity, Position, Timing “Law of specific nerve energies” (Müller in 1826): Different perceptual qualities of the stimulus world are produced by activity in different neurons. In other words, matter how the neurons are activated the perception produced is the same D. Relationship between stimulus and perception: Sensations and perceptions are NOT copies of physical stimuli. The CNS constructs our perceptions, using its ability to integrate, modulate, process, and control stimulus information. II. “CHEMOSENSES” (smell = olfaction, taste = gustation) SMELL FUNCTION RECEPTORS ADEQUATE STIMULUS SENSITIVITY SUBMODALITIES SUBMODALITY CODING AFFERENT NERVES MAIN CNS ROUTES TASTE ●long distance; ●test environment (foraging, communication; ● strong affect ● contact sense; ●test food; ●control intake; ● begin digestive process ●telereceptors; ● in olfactory mucosa of nose; ●like a dendrite with cilia; ●primary sense cell; ●~60-day life cycle; ●>1,000 types ●exteroceptors; ●on tongue; ●made up of papillae (10, front – 300, back) “taste buds”; ●each bud has 20-50 receptor cells: ●secondary sense cell; 10-day life cycle; ● 4 or possibly more types molecules in air (mostly organic; Molecules in fluid (organic and dissolve in mucous at receptor inorganic) surface VERY high (107 mol/ml (102 – 103 Less sensitive: 1016 molecules/ml in some species!) ●Thousands, ● sweet: front tongue (fungiform papillae)- G prot, Ca++ open (foliate poorly defined ● sour: side (front) tongue K+ chan. close papillae) ●Many theories ● salty: side (back) Na+ chan. open ● bitter, back of tongue, K+close, 2nd mess (circumvallate papil.) ● unami (MSG)? carbohydrates?, glutamate? Spatiotemporal pattern in CNS Which peripheral receptor? ● Ist cranial n. (olfactory n.) ● some in Vth n. (trigeminal n.) VIIth (facial n., chorda tympani branch — front of tongue) IXth (glossopharyngeal n., lingual br. —back of tongue) Xth (vagus n., only a few, ---palate, epiglottis) Ist n. olfactory bulb (mitral cells, glomeruli) paleocortex (●amygdala hypothalamus; ●enterorhinal ctx hippocampus; ●pyriform cortex hypothalamus & orbitofrontal cortex) VIIth, IXth, Xth medulla (solitary n.) pons (parabrachial n.) thalamus (VPMp) frontal/parietal lobes of neocortex orbitofrontal cortex, amygdala, hypothalamus, basal forbrain PSB 2000 Class #10: Thurs, 8 Feb 2007 SOMATIC AND VISCERAL SENSATION FUNCTION Provides information about: --our immediate environments: external-skin; internal-viscera (=internal organs) --our body position (for ‘kinesthesia’ and posture): muscles & joints Important for: protection, warning, social communication Involves varying amounts of affect RECEPTORS Located all over and inside the body (skin, muscles, tendons, joints, ligaments, viscera Many specializations: i.e., many different types Cell bodies located in dorsal root ganglia or trigeminal ganglion: pseudounipolar cells Axons: LARGE MYELINATED SMALL-- little myelin SMALL--unmyelinated ‘Aβ’ fibers ‘Aδ’ fibers; ‘C’ fibers fast conduction slow condution very slow conduction Receptive fields can be very large → tiny The timing of responses can be: (a) rapidly adapting; (b) slowly adapting-stimulus bound; (c) slowly adapting with afterdischarge. A B C | | | | |||||||| | | |||||||||||||||||||| response→| | ||| stimulus→ ADEQUATE STIMULI mechanical: pressure, tension chemical: CO2, O2, many other molecules (e.g., inflammatory agents) thermal: high temperature, moderately high temp, low temp, very low temp very high to almost insensitive (requires very little stimulation) (requires very intense stim.) “low threshold” “nociceptors” can change! That is, they can become desensitized or hypersensitized hard to categorize---many?—touch, temperature, pain, itch, tickle, vibration SUBMODALITIES roughness, wetness, etc??? all have varying degrees of affect varying precision (e.g., stomachache vs Braille reading) some not conscious (e.g., blood pressure, posture) SUBMODALITY It is almost impossible to relate the different perceptual submodalities to CODING specific receptors, but there have been attempts: for example,--SENSITIVITY Nociceptors pain Low threshold mechanoreceptors touch AFFERENT NERVES (topographically organized) We shall see the problems with this idea later on ! from skin and muscle—well ordered somatotopy --via spinal nerves (cervical-8; thoracic-12; lumbar-5; sacral-5, coccyxgeal-1) --concept of DERMATOME: The area of skin innervated by sensory fibers in a single spinal nerve. (Thus, we speak of the C8 dermatome, the T12 dermatome, etc…; see next page) from viscera—somatotopy not as well ordered --via nerves of autonomic n.s. (Fig 4.6—p.86 in your text) --we will discuss the parasympathetic and sympathetic systems in class (reread pgs 85-87 in your text) PSB 2000 DERMATOMES Class #10: Thurs, 8 Feb 2007 PSB 2000 Class #11: Tues, 13 Feb 2007 TOUCH and PAIN: CNS mechanisms - From conceptualization to therapy There are TWO ways of conceptualizing how the nervous system creates the perceptions of touch and pain Traditional “pathway” conceptualization This view is misleading. This diagram is similar to the one that is provided in most modern textbooks to illustrate so-called “pain pathways” and “touch pathways.” It is important that we begin with this diagram so that we can then discuss better alternatives. •• • • •••• • • •• • A new view: DYNAMIC DISTRIBUTED SYSTEM SYSTEMS This view fits the data better because: …it takes into consideration: (1) ..convergence and divergence of information flow (2) ..neural plasticity (learning over the lifespan). (3) ...that we cannot equate a perception with a sensory receptor or a pathway example: phantom limbs. brain imaging data PSB 2000 Class #11: Tues, 13 Feb 2007 PSB 2000 Class #11 (cont’d): Tues, 13 Feb 2007 TREATING PAIN A growing list of therapies for pain1 DRUGS Primary analgesics NSAIDS acetaminophen opioids Other analgesics -2 agonists adrenergic antagonists antidepressants anticonvulsants antiarrhythmics calcium channel blockers cannabinoids corticosteroids COX-2 inhibitors GABAB agonists serotonin agonists nitric oxide Adjuvants antihistamines laxatives neuroleptics phenothiazines Routes topical, transdermal, oral buccal, sublingual, intranasal vaginal, rectal inhalation intramuscular, intraperitoneal intravenous epidural, intrathecal intraventricular SOMATIC INTERVENTIONS SITUATIONAL APPROACHES Simple Clinician education attitude clinical setting and arrangement heat/cold exercise massage vibration relaxation Minimally invasive physical therapy traction manipulation ultrasound TENS acupuncture local anesthetics Invasive radiation therapy dorsal column stimulation nerve blocks neurectomy local ganglion blocks sympathectomy rhizotomy DREZ lesions punctate midline myelotomy limited myelotomy commissural myelotomy cordotomy brain stimulation brain lesions 1 January 2007 Self education meditation Diet, herbal supplements art, music, poetry, performing arts sports, gardening, hobbies humor, virtual reality aroma therapy religion pets Interactive hypnosis biofeedback support groups advocacy groups networking self-help groups Structured settings group therapy family counseling job counseling cognitive therapy behavioral therapy psychotherapy multidisciplinary clinic hospice PSB 2000 Class #12&13: Thus & Tues, 15 & 20 Feb 2007 (class #14 on Thurs 23 Feb is Exam #2) AUDITION FUNCTION STIMULI RECEPTOR APPARATUS To provide us with information about the environment that is distant from us. For communication between individuals. Sound pressure waves: alternating “compression” and “rarefaction” of air. The waves have an AMPLITUDE and FREQUENCY. Amplitude is measured in ‘bels’ (or decibels-dB—10ths of bels). A bel is actually a psychophysical measure that is related to a reference “sound” intensity. 0 dB is about level of softest sound an average person can hear at 1000 cycles/sec = 0.0002 dynes/cm2 (a dyne is a physical measure of force; 1dyne = 1gm/cm/sec). dB = 20 log observed force/reference force. [examples: 130dB is max level before tissue damage occurs (stand behind a jet airplane). 20dB is soft whisper at ~10ft.] Outer ear through middle ear: Sound pressure waves are funneled via the pinna (external ear) through the external auditory canal to the eardrum (tympanic membrane), then through the bones of the middle ear (malleus-hammer; incus-anvil; stapes-stirrup) to the OVAL window of the cochlea. The tymp. Membrane is 16X size of the oval window! Thus, the middle ear transforms low energy sound waves to smaller amplitude, but much more forceful displacements of the stapes to move fluid in the cochlea. (The middle ear lies between the tymp. membrane and the oval window.) Cochlea: Coiled structure filled with fluid & divided into three compartments (see text). The receptor organ is in the central compartment, & is called the “organ of Corti” (composed of basilar membrane, hair cells, tectorial membrane). Cochlear fluid is displaced by movements of stapes against oval window, with pressure being released at the ROUND window. Thus, sound pressure waves are transmitted as vibrations of the fluid surrounding the organ of Corti. The base end of the cochlea (stapes end) is stiffer and can transmit higher frequencies than the apical end, which is floppier and transmits lower frequencies. We lose some stiffness with age— therefore we lose ability to hear high frequency components of sound waves. Hair cells: cilia-like structures anchored in the basilar membrane, with tips held in place in the tectorial membrane. Flexing of the basilar/tectorial membranes moves hair cells and changes the flow of K+ and Ca++ ions to produce action potentials in cochlear nerve fibers. AFFERENT NERVES Fibers in the auditory (8th cranial) nerve. The cell bodies of these axons are in the “spiral receptopically organized ganglion” which runs along the length of and adjacent to the cochlea. RECEPTORS (thus, “tonotopically” organized?) AIN ROUTES THROUGH CNS SUBMODALITIES AND THEIR CODING Auditory nerve ipsilateral cochlear nucleus (in medulla/pons) contralateral superior olive (pons) inferior colliculus (midbrain) medial geniculate nucleus (in thalamus) “auditory” cortex in temporal lobe. In other words, the cochlear n. receives input from the ipsilateral ear, but most of the information is then relayed to the contralateral superior olive. Loudness: related to the amplitude of the sound wave, BUT depends on frequency too. Loudness is coded mainly by the # neural fibers activated (not their frequency) (We will discuss audiograms and hearing loss in class.) Pitch: related to both the frequency of firing of receptor fibers and place (i.e., which neurons). From 15 100hZ: frequency code. From 100 4,000Hz: volley coding and place coding (i.e., receptor location on cochlea). (We will discuss cochlear implants in class.) NOTE: Your book states that place coding occurs only at frequencies >4,00Hz. This conclusion is now known to be wrong, mainly because of cochlear implant success. Direction: The brain compares input from two ears, which arrives at different parts of the sound wave (phase) and with different amplitudes. Distance: Depends on sound wave composition. [Higher frequencies are lost faster as they travel towards the ear. Therefore the closer the sound, the more the sound contains higher frequency components. PSB 2000 Class #11: Tues, 13 Feb 2007