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Neural Basis of Speech 2/29/00 Neuron • Neuron = Nervous system cell – Neuron cell body (contains nucleus) – Nucleus (contains genetic material) – Dendrites (projections; communication from 1 neuron to another) – Axon (single long process which conducts nerve impulses to muscles, glands or other neurons) – Rarely can be replaced – Cannot regenerate Neuron Structure Nucleus Dendrites Nucleolus Axon Neuron • Three basic types: – Sensory Neurons • Conduct nerve impulses from sensory receptor (eye or ear) to the brain & spinal cord • Travel from periphery to central site • Direction of travel is afferent – Motor Neurons • Carry neural instructions from the brain to muscles or glands • Travel from central nervous system to the periphery • Direction of travel is efferent – Interneurons • Most numerous of all types • Constitute neural tissue of brain & spinal cord Neuron • Three primary structure types: – Monopolar • Cell body located in a collateral section that connects to transmitting zone of dendrite & axon • Cell of somatic sense (sense of touch & pressure) – Bipolar • Cell body along the main structure of the neuron with the dendrite extending in one direction from body and the axon in the other direction • Found in special senses (vision, audition, olfaction) – Multipolar • Multiple dendrites project from cell body • Neuron of the CNS & motor neuron innervating muscle Types of Neurons CB= Cell Body Bipolar Unipolar Axons Multipolar Neural Connections • Communication between neurons is achieved by the release of neurotransmitters – Synapse= Tiny gap between 2 neurons – Presynaptic Neuron= Transmits impulse – Postsynaptic Neuron= Receiving impulse • Excitation (promoting neural activity) • Inhibition (reducing neural activity) – Neurotransmitter=Chemicals involved in neural communication • Released from terminal boutons of one neuron into cleft of synaptic junction • Contained in synaptic vesicles Synapse PRESYNAPTIC NEURON Vesicles Synaptic Cleft POSTSYNAPTIC NEURON Lock & Key Neurotransmitters • 100 different kinds • Major: – – – – Glutamate Aspartate Gamma-aminobutyric acid (GABA) Glycine • Relatively simple & fast action • Central to basic life processes • Slower Neurotransmitters: – Seratonin – Norepinephrine – Dopamine Synaptic Connections Neuron B Neuron C Synapses Myelin Neuron A Myelin & Glia • Larger axon insulated with fatty coating- Myelin – Increases speed of neural transmission – Reduces interference with the neural message – Multiple sclerosis- dymyelinating • Neurons outnumbered by glial cells – Hold neurons in place & provide nutrients – Oligodendroglia (form myelin in CNS) – Schwann (form myelin in the PNS) Neural Impulse • Neurons generate electrical impulse traveling the length of the nerve fibers • Neural activity= electrical & chemical activity • Neuron is like a battery – Stores electrical potential by accumulating positive charge in one terminal & excessive negative at the other terminal – An electrical potential across the membrane is created • Extracellular positive compared to intracellular • Ions carry charges – positive: sodium (Na+ ) & potassium (K+) – Negative: chlorine (CL=) Neural Impulse • Positive ions- concentrated outside the cell (sodium) • Negative ions- concentrated inside the cell • Resting membrane potential (-70 millivolts) created due to excessive positive outside cell – Maintained through sodium-potassium pump • act to exchange sodium ions found inside the cell with potassium ions found outside the ell • Neuron at rest= polarized • Neural activity= depolarization Sodium (Na+) Extracellular fluid Sodium-Potassium Pump Sodium Channel Potassium (K+) Intracellular fluid Potassium Channel Neural Impulse • Action potential occurs= Wave of depolarization – Depolarization occurs when an action in another neuron momentarily lowers the voltage of a region of a membrane • Causes voltage-controlled gates to open that regulate sodium channels • Sodium floods into the cell • Polarity reverses from -70 to +30 mV • Cell returns to rest (sodium-potassium pump) Neural Impulse • Depolarization effects tiny portion of membrane at a time. • Causes a wave down the entire membrane by causing voltage gated channels to open • Wave continues until the axon terminal – Synapse with other neuron – Transmitted to next neuron Na+ A C Na+ CL- A= resting state; ionic imbalance B K+ B= depolarization; Sodium channels open; potential positive Na+ K+ D K+ K+ C= Opening of potassium channels; potential returns negative D= return to rest as sodium-potassium pump works A B C D Neuroanatomy of the Vocal Mechanism Neuroanatomy of the vocal Mechanism • Volitional control of muscles of the larynx resides in the brain. • Connecting points in brain that have a role in control of phonation: cortex, subcortical areas, midbrain & medulla. • Next slides will briefly review phonation neuroanatomy & neurophysiology. Cortical Mechanisms of Phonatory Control • The cerebral cortex is responsible for: – conceptualization, planning, and execution of speech , including phonation. • Three major areas of the cortex responsible for vocalization: – a) Precentral & postcentral gyrus, – b) Anterior (Broca’s) area, – c) Supplementary motor area. Cortical Areas Involved in Speech Movement Control Premotor & Supplementary Cortex Primary Motor Cortex Somatosensory Cortex Broca’s Area -Stimulation of these areas can initiate, stop or distort vocalization. -These behaviors occur in dominant & nondominant hemispheres. Speech and Phonation are complex motor acts • Involves simultaneous activation and control of many muscles. • Control of these motor acts occurs primarily in the cortex. • Control of individual muscles occurs lower in the brain. – No evidence that cortical stimulation produces a response in a single solitary muscle. • Higher brain function = idealization of the event, integration of sensory information, feedback control, and coordination of various muscles. Subcortical Mechanisms • Motor cortex has connections to the Thalamus ( egg shaped in the middle of the cerebral cortex), – A major portion of the diencephalon or interbrain. – Contain nuclei for language & speech – Relay station from cortical to subcortical brain – Thalamus has major pathways to the motor cortex & Broca’s area. • Parts of the diencephalon: a) hypothalamus, b) metathalumus, c) epithalumus, d) subthalumus, & e) third ventricle. Projections to Cerebral Cortex •Acts as a relay for impulses in lower areas of the brain. Diencepahalon Thalamus: What Does it Do? •Integrates emotion into a complex motor act. Pons Thalamus Midbrain Projections to Cerebellar Cortex •Plays a major role in: • coordinating outgoing information from cortex, • integrating incoming sensory information • adding emotionality to speech Nuclei in thalamus that project to parts of the cerebral cortex • Motor area receives its projections from the ventrolateral nucleus. to & from Prenucleus to & from Sup. Parietal Lobule to & from Parietal Lobe Massa Intermedia • 1971- ventrolateral nucleus shown to be responsible for initiation of speech movements & control of loudness, pitch, rate & articulation. • Broca’s area- receives connections from dorsomedian nuclei. Dorsal Median Lateral Dorsal Ventral Lateral Ventral posterior Lateral Midbrain Structures • Midbrain (mesencephalon) lies beneath the thalamus. • Cerebral peduncles lie on anterior surface of the midbrain and connect the cerebrum with the brainstem and spinal cord. • Posterior side has four colliculi: Superior (visual function), inferior (audition). • Within midbrain lies the cerebral aqueduct of Sylvius, surrounded by periaqueductal gray. Periaqueductal Gray: What does it do? • Stimulation of dorsal and ventrolateral areas of periaqueductal gray = activity in some laryngeal muscles. • 1985- Larson reported some cells in ventrolateral area stimulate muscle activity, whereas some suppress activity. • Periaqueductal gray is an intermediate area between recognition of a stimulus and the production of a motor act. Brainstem • Bilateral structures in brainstem implicated in the neural control of phonation: • Nucleus ambiguus • Nucleus tractus solitarii • Nucleus parabrachialis • How do we know these structures are involved in phonation? Yoshida, Mitsumasu, Hirano Study • Traced connections among brainstem structures. • Injected tracer chemical into one nucleus ambiguus. • Found evidence of tracer throughout the contralateral nuclei, nuclei tractus solitarri bilaterally, in nucleus parabrachialis and bilaterally in the lateral and ventrolateral parts of the periaqueductal gray area, with a predominance ipsilaterally. • Conclusion: Many interconnections bilaterally among the nucleus ambiguous, nucleus tractus solitarri, and motor roots of vagus. Cerebellum • Structure lying posterior to the midbrain area. • Implicated in the control of movement. • Three main portions: a) vermis, b) pars intermedia, c) hemispheres • Consists of many traverse folia- increases surface area. • Fissura prima- fissure separating anterior & posterior lobes. References: • Colton, R.H. & Casper, J.K.,(1990), Understanding Voice Problems: A physiological perspective for diagnosis and treatment,, Williams & Wilkins. • Bhatnager, S.C. & Andy, O.J., (1995), Neuroscience for the study of communicative disorders, Williams & Wilkins. • Kuehn, D.P., Lemme, M.L. & Baumgartner, J.M., (1989), Neural basis of speech, hearing, and language, CollegeHill Press. • Lieberman, M., (1991), Neuroanatomy made easy and understandable, Aspen Publishers. • Netsell, R., (1985), Speech and language evaluation in neurology-adult disorders, Grune & Stratton. • Poritsky, R., (1992), Neuroanatomy: a functional atlas of parts & pathways, Mosby-Year Book.