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Chapter 14 *Lecture PowerPoint The Brain and Cranial Nerves *See separate FlexArt PowerPoint slides for all figures and tables preinserted into PowerPoint without notes. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Introduction • The human brain is complex • Brain function is associated with life • This chapter is a study of brain and cranial nerves directly connected to it • Will provide insight into brain circuitry and function 14-2 Introduction • Aristotle thought brain was “radiator” to cool blood • Hippocrates was more accurate: “from the brain only, arises our pleasures, joys, laughter, and jests, as well as our sorrows, pains, griefs, and tears” • Cessation of brain activity—clinical criterion of death • Evolution of the central nervous system shows spinal cord has changed very little while brain has changed a great deal – Greatest growth in areas of vision, memory, and motor control of the prehensile hand 14-3 Overview of the Brain • Expected Learning Outcomes – Describe the major subdivisions and anatomical landmarks of the brain. – Describe the locations of its gray and white matter. – Describe the embryonic development of the CNS and relate this to adult brain anatomy. 14-4 Major Landmarks Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Rostral—toward the forehead Rostral Caudal Central sulcus • Caudal—toward the spinal cord • Brain weighs about 1,600 g (3.5 lb) in men, and 1,450 g in women Gyri Cerebrum Lateral sulcus Cerebellum Temporal lobe Brainstem Spinal cord (b) Lateral view Figure 14.1b 14-5 Major Landmarks Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Three major portions of the brain – Cerebrum is 83% of brain volume; cerebral hemispheres, gyri and sulci, longitudinal fissure, corpus callosum Rostral Caudal Central sulcus Gyri Cerebrum Lateral sulcus – Cerebellum contains 50% of the Temporal lobe neurons; second largest brain region, located in posterior cranial fossa Brainstem – Brainstem is the portion of the brain that remains if the cerebrum and cerebellum are removed; diencephalon, midbrain, pons, and medulla oblongata Cerebellum Spinal cord (b) Lateral view Figure 14.1b 14-6 Major Landmarks • Longitudinal fissure—deep groove that separates cerebral hemispheres Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Gyri—thick folds Frontal lobe • Sulci—shallow grooves Cerebral hemispheres Central sulcus Parietal lobe • Corpus callosum—thick nerve bundle at bottom of longitudinal fissure that connects hemispheres Occipital lobe Longitudinal fissure (a) Superior view Figure 14.1a 14-7 Major Landmarks Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Occupies posterior cranial fossa Rostral Caudal Central sulcus • Marked by gyri, sulci, and fissures Gyri Cerebrum Lateral sulcus Cerebellum Temporal lobe • About 10% of brain volume Brainstem Spinal cord (b) Lateral view • Contains over 50% of brain neurons Figure 14.1b 14-8 Major Landmarks • Brainstem—what remains of the brain if the cerebrum and cerebellum are removed Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Rostral Caudal Central sulcus Gyri Cerebrum Lateral sulcus Cerebellum Temporal lobe • Major components Brainstem – Diencephalon – Midbrain – Pons Spinal cord (b) Lateral view Figure 14.1b – Medulla oblongata 14-9 Major Landmarks Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Central sulcus Parietal lobe Cingulate gyrus Corpus callosum Parieto–occipital sulcus Frontal lobe Occipital lobe Thalamus Anterior commissure Hypothalamus Habenula Pineal gland Optic chiasm Posterior commissure Mammillary body Epithalamus Cerebral aqueduct Pituitary gland Fourth ventricle Temporal lobe Cerebellum Midbrain Pons Medulla oblongata (a) Figure 14.2a 14-10 Major Landmarks Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Cingulate gyrus Lateral ventricle Corpus callosum Parieto–occipital sulcus Choroid plexus Pineal gland Thalamus Hypothalamus Occipital lobe Midbrain Posterior commissure Pons Fourth ventricle Cerebellum Medulla oblongata (b) b: © The McGraw-Hill Companies, Inc./Dennis Strete, photographer Figure 14.2b 14-11 Gray and White Matter • Gray matter—the seat of neuron cell bodies, dendrites, and synapses – Dull white color when fresh, due to little myelin – Forms surface layer (cortex) over cerebrum and cerebellum – Forms nuclei deep within brain • White matter—bundles of axons – Lies deep to cortical gray matter, opposite relationship in the spinal cord – Pearly white color from myelin around nerve fibers – Composed of tracts, or bundles of axons, that connect one part of the brain to another, and to the spinal cord 14-12 Embryonic Development • Nervous system develops from ectoderm – Outermost tissue layer of the embryo • Early in third week of development – Neuroectoderm: dorsal streak appears along the length of embryo – Thickens to form neural plate • Destined to give rise to most neurons and all glial cells except microglia, which come from mesoderm • As thickening progresses – Neural plate sinks and its edges thicken • Forming a neural groove with a raised neural fold on each side • Neural folds fuse along the midline – Beginning in the cervical region and progressing rostrally and caudally 14-13 Embryonic Development • By fourth week, creates a hollow channel—neural tube – Neural tube separates from overlying ectoderm – Sinks deeper – Grows lateral processes that later will form motor nerve fibers – Lumen of neural tube becomes fluid-filled space • Central canal in spinal cord • Ventricles of the brain 14-14 Embryonic Development Cont. • Neural crest—formed from ectodermal cells that lay along the margins of the groove and separate from the rest forming a longitudinal column on each side – Gives rise to the two inner meninges, most of the peripheral nervous system, and other structures of the skeletal, integumentary, and endocrine systems • By fourth week, the neural tube exhibits three anterior dilations (primary vesicles) – Forebrain (prosencephalon) – Midbrain (mesencephalon) – Hindbrain (rhombencephalon) 14-15 Embryonic Development • By fifth week, it subdivides into five secondary vesicles – Forebrain divides into two of them • Telencephalon—becomes cerebral hemispheres • Diencephalon—has optic vesicles that become retina of the eye – Midbrain remains undivided • Mesencephalon – Hindbrain divides into two vesicles • Metencephalon • Myelencephalon 14-16 Embryonic Development Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Neural plate Neural crest Neural crest Neural fold Ectoderm Neural groove Notochord (a) 19 days (b) 20 days Neural crest Neural tube Somites (c) 22 days (d) 26 days Figure 14.3 14-17 Embryonic Development Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Fourth week Telencephalon Prosencephalon Diencephalon – Forebrain – Midbrain – Hindbrain • Fifth week – – – – – Optic vesicle Mesencephalon Metencephalon Rhombencephalon Myelencephalon Spinal cord (a) 4 weeks (b) 5 weeks Telencephalon Diencephalon Mesencephalon Metencephalon Myelencephalon Telencephalon Forebrain Diencephalon Mesencephalon Midbrain Pons Metencephalon Cerebellum Hindbrain Myelencephalon (medulla oblongata) Spinal cord (c) Fully developed Figure 14.4 14-18 Meninges, Ventricles, Cerebrospinal Fluid, and Blood Supply • Expected Learning Outcomes – Describe the meninges of the brain. – Describe the fluid-filled chambers within the brain. – Discuss the production, circulation, and function of the cerebrospinal fluid that fills these chambers. – Explain the significance of the brain barrier system. 14-19 Meninges • Meninges—three connective tissue membranes that envelop the brain – Lies between the nervous tissue and bone – As in spinal cord, they are the dura mater, arachnoid mater, and the pia mater – Protect the brain and provide structural framework for its arteries and veins 14-20 Meninges • Dura mater – In cranial cavity; has two layers • Outer periosteal—equivalent to periosteum of cranial bones • Inner meningeal—continues into vertebral canal and forms dural sac around spinal cord – Cranial dura mater is pressed closely against cranial bones • No epidural space • Not attached to bone except: around foramen magnum, sella turcica, the crista galli, and sutures of the skull • Layers separated by dural sinuses—collect blood circulating through brain 14-21 Meninges Cont. – Folds inward to extend between parts of the brain • Falx cerebri separates the two cerebral hemispheres • Tentorium cerebelli separates cerebrum from cerebellum • Falx cerebelli separates the right and left halves of cerebellum 14-22 Meninges • Arachnoid mater and pia mater are similar to those in the spinal cord • Arachnoid mater – Transparent membrane over brain surface – Subarachnoid space separates it from pia mater below – Subdural space separates it from dura mater above in some places • Pia mater – Very thin membrane that follows contours of brain, even dipping into sulci – Not usually visible without a microscope 14-23 Meninges Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Skull Dura mater: Periosteal layer Meningeal layer Subdural space Subarachnoid space Arachnoid granulation Arachnoid mater Superior sagittal sinus Blood vessel Falx cerebri (in longitudinal fissure only) Pia mater Brain: Gray matter White matter Figure 14.5 14-24 Meningitis • Meningitis—inflammation of the meninges – Serious disease of infancy and childhood – Especially between 3 months and 2 years of age • Caused by bacterial and virus invasion of the CNS by way of the nose and throat • Pia mater and arachnoid are most often affected • Bacterial meningitis can cause swelling of the brain, enlargment of the ventricles, and hemorrhage • Signs include high fever, stiff neck, drowsiness, and intense headache; may progress to coma then death within hours of onset • Diagnosed by examining the CSF for bacteria – Lumbar puncture (spinal tap) draws fluid from subarachnoid space between two lumbar vertebrae 14-25 Ventricles and Cerebrospinal Fluid Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Caudal Rostral Lateral ventricles Cerebrum Interventricular foramen Lateral ventricle Interventricular foramen Third ventricle Third ventricle Cerebral aqueduct Fourth ventricle Cerebral aqueduct Lateral aperture Fourth ventricle Median aperture Lateral aperture Median aperture Central canal (b) Anterior view (a) Lateral view Figure 14.6a,b 14-26 Ventricles and Cerebrospinal Fluid Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Rostral (anterior) Longitudinal fissure Frontal lobe Gray matter (cortex) White matter Corpus callosum (anterior part) Lateral ventricle Caudate nucleus Septum pellucidum emporal lobe T Third ventricle Sulcus Gyrus Lateral sulcus Insula Thalamus Lateral ventricle Choroid plexus Corpus callosum (posterior part) Occipital lobe Longitudinal fissure Figure 14.6c (c) Caudal (posterior) c: © The McGraw-Hill Companies, Inc./Rebecca Gray,photographer/Don Kincaid, dissections 14-27 Ventricles and Cerebrospinal Fluid • Ventricles—four internal chambers within the brain – Two lateral ventricles: one in each cerebral hemisphere • Interventricular foramen—a tiny pore that connects to third ventricle – Third ventricle: single narrow medial space beneath corpus callosum • Cerebral aqueduct runs through midbrain and connects third to fourth ventricle – Fourth ventricle: small triangular chamber between pons and cerebellum • Connects to central canal, runs down through spinal cord 14-28 Ventricles and Cerebrospinal Fluid Cont. • Choroid plexus—spongy mass of blood capillaries on the floor of each ventricle • Ependyma—neuroglia that lines the ventricles and covers choroid plexus – Produces cerebrospinal fluid 14-29 Ventricles and Cerebrospinal Fluid • Cerebrospinal fluid (CSF)—clear, colorless liquid that fills the ventricles and canals of CNS – Bathes its external surface • Brain produces and absorbs 500 mL/day – – – – 100 to 160 mL normally present at one time 40% formed in subarachnoid space external to brain 30% by the general ependymal lining of the brain ventricles 30% by the choroid plexuses • Production begins with the filtration of blood plasma through the capillaries of the brain – Ependymal cells modify the filtrate, so CSF has more sodium and chloride than plasma, but less potassium, calcium, glucose, and very little protein 14-30 Ventricles and Cerebrospinal Fluid • CSF continually flows through and around the CNS – Driven by its own pressure, beating of ependymal cilia, and pulsations of the brain produced by each heartbeat • CSF secreted in lateral ventricles flows through intervertebral foramina into third ventricle • Then down the cerebral aqueduct into the fourth ventricle • Third and fourth ventricles add more CSF along the way 14-31 Ventricles and Cerebrospinal Fluid • Small amount of CSF fills the central canal of the spinal cord – All escapes through three pores • Median aperture and two lateral apertures • Leads into subarachnoid space of brain and spinal cord surface • CSF is reabsorbed by arachnoid villi – Cauliflower-shaped extension of the arachnoid meninx – Protrudes through dura mater – Into superior sagittal sinus – CSF penetrates the walls of the villi and mixes with the blood in the sinus 14-32 Ventricles and Cerebrospinal Fluid • Functions of CSF – Buoyancy • Allows brain to attain considerable size without being impaired by its own weight • If it rested heavily on floor of cranium, the pressure would kill the nervous tissue – Protection • Protects the brain from striking the cranium when the head is jolted • Shaken child syndrome and concussions do occur from severe jolting – Chemical stability • Flow of CSF rinses away metabolic wastes from nervous tissue and homeostatically regulates its chemical environment 14-33 Ventricles and Cerebrospinal Fluid Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Arachnoid villus 8 Superior sagittal sinus Arachnoid mater 1 CSF is secreted by choroid plexus in each lateral ventricle. Subarachnoid space Dura mater 1 2 CSF flows through interventricular foramina into third ventricle. 2 Choroid plexus Third ventricle 3 3 Choroid plexus in third ventricle adds more CSF. 7 4 Cerebral aqueduct 4 CSF flows down cerebral aqueduct to fourth ventricle. Lateral aperture 5 Choroid plexus in fourth ventricle adds more CSF. Fourth ventricle 6 5 6 CSF flows out two lateral apertures and one median aperture. Median aperture 7 CSF fills subarachnoid space and bathes external surfaces of brain and spinal cord. 7 8 At arachnoid villi, CSF is reabsorbed into venous blood of dural venous sinuses. Central canal of spinal cord Subarachnoid space of spinal cord Figure 14.7 14-34 Blood Supply and the Brain Barrier System • Brain is only 2% of the adult body weight, and receives 15% of the blood – 750 mL/min. • Neurons have a high demand for ATP, and therefore, oxygen and glucose, so a constant supply of blood is critical to the nervous system – A 10-second interruption of blood flow may cause loss of consciousness – A 1- to 2-minute interruption can cause significant impairment of neural function – Going 4 minutes without blood causes irreversible brain damage 14-35 Blood Supply and the Brain Barrier System • Blood is also a source of antibodies, macrophages, bacterial toxins, and other harmful agents • Brain barrier system—strictly regulates what substances can get from the bloodstream into the tissue fluid of the brain • Two points of entry must be guarded – Blood capillaries throughout the brain tissue – Capillaries of the choroid plexus 14-36 Blood Supply and the Brain Barrier System • Blood–brain barrier—protects blood capillaries throughout brain tissue – Consists of tight junctions between endothelial cells that form the capillary walls – Astrocytes reach out and contact capillaries with their perivascular feet – Induce the endothelial cells to form tight junctions that completely seal off gaps between them – Anything leaving the blood must pass through the cells, and not between them – Endothelial cells can exclude harmful substances from passing to the brain tissue while allowing necessary ones to pass 14-37 Blood Supply and the Brain Barrier System • Blood–CSF barrier—protects the brain at the choroid plexus – Forms tight junctions between the ependymal cells – Tight junctions are absent from ependymal cells elsewhere • Important to allow exchange between brain tissue and CSF • Blood barrier system is highly permeable to water, glucose, and lipid-soluble substances such as oxygen, carbon dioxide, alcohol, caffeine, nicotine, and anesthetics • Slightly permeable to sodium, potassium, chloride, and the waste products urea and creatinine 14-38 Blood Supply and the Brain Barrier System • Obstacle for delivering medications such as antibiotics and cancer drugs • Trauma and inflammation can damage BBS and allow pathogens to enter brain tissue – Circumventricular organs (CVOs)—places in the third and fourth ventricles where the barrier is absent • Blood has direct access to the brain • Enables the brain to monitor and respond to fluctuations in blood glucose, pH, osmolarity, and other variables • CVOs afford a route for invasion by the human immunodeficiency virus (HIV) 14-39 The Hindbrain and Midbrain • Expected Learning Outcomes – List the components of the hindbrain and midbrain and their functions. – Describe the location and functions of the reticular formation. 14-40 The Medulla Oblongata • Embryonic myelencephalon becomes medulla oblongata Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Central sulcus Parietal lobe Cingulate gyrus Corpus callosum • Begins at foramen magnum of the skull Parieto–occipital sulcus Frontal lobe Occipital lobe Thalamus Anterior commissure Hypothalamus Habenula Pineal gland Optic chiasm Posterior commissure Mammillary body Epithalamus Cerebral aqueduct Pituitary gland • Extends for about 3 cm rostrally and ends at a groove between the medulla and pons • Slightly wider than spinal cord Fourth ventricle Temporal lobe Midbrain Cerebellum Pons Medulla oblongata (a) Figure 14.2a • Pyramids—pair of external ridges on anterior surface – Resembles side-by-side baseball bats 14-41 The Medulla Oblongata • Olive—a prominent bulge lateral to each pyramid Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Central sulcus Parietal lobe Cingulate gyrus Corpus callosum Parieto–occipital sulcus Frontal lobe • Posteriorly, gracile and cuneate fasciculi of the spinal cord continue as two pair of ridges on the medulla Occipital lobe Thalamus Anterior commissure Hypothalamus Optic chiasm Mammillary body Pituitary gland Temporal lobe Habenula Pineal gland Epithalamus Posterior commissure Cerebral aqueduct Fourth ventricle Midbrain Cerebellum Pons Medulla oblongata • All nerve fibers connecting the brain to the spinal cord pass through the medulla (a) Figure 14.2a • Four pairs of cranial nerves begin or end in medulla—IX, X, XI, XII 14-42 The Medulla Oblongata • Cardiac center – Adjusts rate and force of heart • Vasomotor center – Adjusts blood vessel diameter • Respiratory centers – Control rate and depth of breathing • Reflex centers – For coughing, sneezing, gagging, swallowing, vomiting, salivation, sweating, movements of tongue and head 14-43 The Medulla Oblongata Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. (a) Midbrain (b) Pons (c) Medulla Nucleus of hypoglossal nerve Gracile nucleus Fourth ventricle Cuneate nucleus Nucleus of vagus nerve Dorsal spinocerebellar tract Trigeminal nerve: Nucleus Tract Reticular formation Tectospinal tract Medial lemniscus Inferior olivary nucleus Olive Hypoglossal nerve Pyramids of medulla Corticospinal tract (c) Medulla oblongata Figure 14.9c • Pyramids contain descending fibers called corticospinal tracts – Carry motor signals to skeletal muscles • Inferior olivary nucleus—relay center for signals to cerebellum • Reticular formation—loose network of nuclei extending throughout the medulla, pons, and midbrain – Contains cardiac, vasomotor, and respiratory centers 14-44 The Medulla Oblongata Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Diencephalon: Thalamus Infundibulum Optic tract Mammillary body Cranial nerves: Midbrain: Optic nerve (II) Cerebral peduncle Oculomotor nerve (III) Trochlear nerve (IV) Trigeminal nerve (V) Abducens nerve (VI) Pons Facial nerve (VII) Vestibulocochlear nerve (VIII) Glossopharyngeal nerve (IX) Vagus nerve (X) Accessory nerve (XI) Medulla oblongata: Pyramid Hypoglossal nerve (XII) Anterior median fissure Regions of the brainstem Diencephalon Midbrain Pyramidal decussation Spinal nerves Spinal cord Pons Medulla oblongata (a) Ventral view Figure 14.8a 14-45 The Pons Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Regions of the brainstem Diencephalon Midbrain Pons Medulla oblongata Diencephalon: Thalamus Lateral geniculate body Pineal gland Medial geniculate body Optic tract Midbrain: Superior colliculus Inferior colliculus Cerebral peduncle Pons Superior cerebellar peduncle Middle cerebellar peduncle Fourth ventricle Inferior cerebellar peduncle Olive Medulla oblongata Cuneate fasciculus Gracile fasciculus Spinal cord (b) Dorsolateral view Figure 14.8b 14-46 The Pons Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Central sulcus Parietal lobe Cingulate gyrus Corpus callosum Parieto–occipital sulcus Frontal lobe Occipital lobe Thalamus Anterior commissure Hypothalamus Habenula Pineal gland Epithalamus Posterior commissure Optic chiasm Mammillary body Cerebral aqueduct Pituitary gland Fourth ventricle Temporal lobe Midbrain Cerebellum Pons Medulla oblongata (a) Figure 14.2a • Metencephalon—develops into the pons and cerebellum • Pons—anterior bulge in brainstem, rostral to medulla • Cerebral peduncles—connect cerebellum to pons and midbrain 14-47 The Pons • Ascending sensory tracts • Descending motor tracts • Pathways in and out of cerebellum • Cranial nerves V, VI, VII, and VIII – Sensory roles: hearing, equilibrium, taste, facial sensations – Motor roles: eye movement, facial expressions, chewing, swallowing, urination, and secretion of saliva and tears • Reticular formation in pons contains additional nuclei concerned with sleep, respiration, posture 14-48 The Pons Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. (a) Midbrain (b) Pons (c) Medulla Vermis of cerebellum Fourth ventricle Superior cerebellar peduncle Middle cerebellar peduncle Ventral spinocerebellar tract Tectospinal tract Trigeminal nerve nuclei Anterolateral system Sensory root of trigeminal nerve Trigeminal nerve Reticular formation Transverse fascicles Medial lemniscus Longitudinal fascicles (b) Pons Figure 14.9b 14-49 The Midbrain • Mesencephalon becomes one mature brain structure, the midbrain – Short segment of brainstem that connects the hindbrain to the forebrain – Contains cerebral aqueduct – Contains continuations of the medial lemniscus and reticular formation – Contains the motor nuclei of two cranial nerves that control eye movements: CN III (oculomotor) and CN IV (trochlear) 14-50 The Midbrain • Mesencephalon (cont.) – Tectum: rooflike part of the midbrain posterior to cerebral aqueduct • Exhibits four bulges, the corpora quadrigemina • Upper pair, the superior colliculi, function in visual attention, tracking moving objects, and some reflexes • Lower pair, the inferior colliculi, receives signals from the inner ear – Relays them to other parts of the brain, especially the thalamus 14-51 The Midbrain Cont. – Cerebral peduncles: two stalks that anchor the cerebrum to the brainstem anterior to the cerebral aqueduct • Cerebral peduncles—three main components – Tegmentum • Dominated by the red nucleus • Pink color due to high density of blood vessels • Connections go to and from cerebellum • Collaborates with cerebellum for fine motor control 14-52 The Midbrain – Substantia nigra • Dark gray to black nucleus pigmented with melanin • Motor center that relays inhibitory signals to thalamus and basal nuclei preventing unwanted body movement • Degeneration of neurons leads to tremors of Parkinson disease – Cerebral crus • Bundle of nerve fibers that connect the cerebrum to the pons • Carries corticospinal tracts 14-53 The Midbrain Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Posterior Superior colliculus Tectum Cerebral aqueduct Medial geniculate nucleus Reticular formation Central gray matter Cerebral peduncle: Oculomotor nucleus Medial lemniscus Tegmentum Red nucleus Substantia nigra (a) Midbrain Cerebral crus Oculomotor nerve (III) (b) Pons Anterior (c) Medulla (a) Midbrain Figure 14.9a 14-54 The Reticular Formation Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Radiations to cerebral cortex Thalamus Auditory input Visual input Reticular formation Ascending general sensory fibers Descending motor fibers to spinal cord Figure 14.10 • Loosely organized web of gray matter that runs vertically through all levels of the brainstem • Clusters of gray matter scattered throughout pons, midbrain, and medulla • Occupies space between white fiber tracts and brainstem nuclei • Has connections with many areas of cerebrum – More than 100 small neural networks without distinct boundaries 14-55 The Reticular Formation • Networks – Somatic motor control • Adjust muscle tension to maintain tone, balance, and posture • Especially during body movements • Relays signals from eyes and ears to the cerebellum • Integrates visual, auditory, and balance and motion stimuli into motor coordination • Gaze center—allows eyes to track and fixate on objects • Central pattern generators—neural pools that produce rhythmic signals to the muscles of breathing and swallowing 14-56 The Reticular Formation Cont. – Cardiovascular control • Includes cardiac and vasomotor centers of medulla oblongata – Pain modulation • One route by which pain signals from the lower body reach the cerebral cortex • Origin of descending analgesic pathways—fibers act in the spinal cord to block transmission of pain signals to the brain 14-57 The Reticular Formation – Sleep and consciousness • Plays central role in states of consciousness, such as alertness and sleep • Injury to reticular formation can result in irreversible coma – Habituation • Process in which the brain learns to ignore repetitive, inconsequential stimuli while remaining sensitive to others 14-58 The Cerebellum Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Anterior Vermis Anterior lobe Posterior lobe Cerebellar hemisphere Folia Posterior Figure 14.11b (b) Superior view • The largest part of the hindbrain and the second largest part of the brain as a whole • Consists of right and left cerebellar hemispheres connected by vermis • Cortex of gray matter with folds (folia) and four deep nuclei in each hemisphere • Contains more than half of all brain neurons—about 100 billion – Granule cells and Purkinje cells synapse on deep nuclei • White matter branching pattern is called arbor vitae 14-59 The Cerebellum Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Superior colliculus Inferior colliculus Pineal gland Posterior commissure Cerebral aqueduct Mammillary body Midbrain White matter (arbor vitae) Gray matter Oculomotor nerve Fourth ventricle Pons Medulla oblongata Figure 14.11a (a) Median section • Cerebellar peduncles—three pairs of stalks that connect the cerebellum to the brainstem – Inferior peduncles: connected to medulla oblongata • Most spinal input enters the cerebellum through inferior peduncle – Middle peduncles: connected to the pons • Most input from the rest of the brain enters by way of middle peduncle – Superior peduncles: connected to the midbrain • Carries cerebellar output • Consist of thick bundles of nerve fibers that carry signals to and from the cerebellum 14-60 The Cerebellum • Monitors muscle contractions and aids in motor coordination • Evaluation of sensory input – Comparing textures without looking at them – Spatial perception and comprehension of different views of threedimensional objects belonging to the same object • Timekeeping center – Predicting movement of objects – Helps predict how much the eyes must move in order to compensate for head movements and remain fixed on an object 14-61 The Cerebellum Cont. • Hearing – Distinguish pitch and similar-sounding words • Planning and scheduling tasks • Lesions may result in emotional overreactions and trouble with impulse control 14-62 The Forebrain • Expected Learning Outcomes – Name the three major components of the diencephalon and describe their locations and functions. – Identify the five lobes of the cerebrum and their functions. – Identify the three types of tracts in the cerebral white matter. – Describe the distinctive cell types and histological arrangement of the cerebral cortex. – Describe the location and functions of the basal nuclei and limbic system. 14-63 The Forebrain • Forebrain consists of two parts – Diencephalon Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Encloses the third ventricle Telencephalon Forebrain • Most rostral part of the brainstem Diencephalon Mesencephalon Pons Metencephalon Cerebellum • Has three major embryonic derivatives – Thalamus – Hypothalamus – Epithalamus – Telencephalon Midbrain Hindbrain Myelencephalon (medulla oblongata) Spinal cord (c) Fully developed Figure 14.4c • Develops chiefly into the cerebrum 14-64 The Diencephalon: Thalamus Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Thalamic Nuclei Anterior group Part of limbic system; memory and emotion Medial group Emotional output to prefrontal cortex; awareness of emotions Ventral group Somesthetic output to postcentral gyrus; signals from cerebellum and basal nuclei to motor areas of cortex Lateral group Somesthetic output to association areas of cortex; contributes to emotional function of limbic system Posterior group Relay of visual signals to occipital lobe (via lateral geniculate nucleus) and auditory signals to temporal lobe (via medial geniculate nucleus) Lateral geniculate nucleus Medial geniculate nucleus (a) Thalamus Figure 14.12a • Thalamus—ovoid mass on each side of the brain perched at the superior end of the brainstem beneath the cerebral hemispheres – Constitutes about four-fifths of the diencephalon – Two thalami are joined medially by a narrow intermediate mass – Composed of at least 23 nuclei; to consider five major functional groups 14-65 The Diencephalon: Thalamus Cont. – “Gateway to the cerebral cortex”: nearly all input to the cerebrum passes by way of synapses in the thalamic nuclei, filters information on its way to cerebral cortex – Plays key role in motor control by relaying signals from cerebellum to cerebrum and providing feedback loops between the cerebral cortex and the basal nuclei – Involved in the memory and emotional functions of the limbic system: a complex of structures that include some cerebral cortex of the temporal and frontal lobes and some of the anterior thalamic nuclei 14-66 The Diencephalon: Hypothalamus Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Hypothalamus—forms part of the walls and floor of the third ventricle • Extends anteriorly to optic chiasm and posteriorly to the paired mammillary bodies Central sulcus Parietal lobe Cingulate gyrus Corpus callosum Parieto–occipital sulcus Frontal lobe Occipital lobe Thalamus Anterior commissure Hypothalamus Optic chiasm Mammillary body Pituitary gland Temporal lobe Habenula Pineal gland Epithalamus Posterior commissure Cerebral aqueduct Fourth ventricle Cerebellum Midbrain Pons Medulla oblongata (a) Figure 14.2a • Each mammillary body contains three or four mammillary nuclei – Relay signals from the limbic system to the thalamus 14-67 The Diencephalon: Hypothalamus • Infundibulum—a stalk that attaches the pituitary gland to the hypothalamus • Major control center of autonomic nervous system and endocrine system – Plays essential role in homeostatic regulation of all body systems • Functions of hypothalamic nuclei – Hormone secretion • Controls anterior pituitary • Regulates growth, metabolism, reproduction, and stress responses 14-68 The Diencephalon: Hypothalamus Cont. – Hormone secretion • Controls anterior pituitary • Regulates growth, metabolism, reproduction, and stress responses – Autonomic effects • Major integrating center for autonomic nervous system • Influences heart rate, blood pressure, gastrointestinal secretions, motility, etc. – Thermoregulation • Hypothalamic thermostat monitors body temperature • Activates heat-loss center when temp is too high • Activates heat-promoting center when temp is too low 14-69 The Diencephalon: Hypothalamus Cont. – Food and water intake • Hunger and satiety centers monitor blood glucose and amino acid levels – Produce sensations of hunger and satiety • Thirst center monitors osmolarity of the blood – Rhythm of sleep and waking • Controls 24-hour (circadian) rhythm of activity – Memory • Mammillary nuclei receive signals from hippocampus – Emotional behavior • Anger, aggression, fear, pleasure, and contentment 14-70 The Diencephalon: Epithalamus Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Central sulcus Parietal lobe Cingulate gyrus Corpus callosum Parieto–occipital sulcus Frontal lobe Occipital lobe Thalamus Anterior commissure Hypothalamus Habenula Pineal gland Epithalamus Posterior commissure Optic chiasm Mammillary body Cerebral aqueduct Pituitary gland Fourth ventricle Temporal lobe Midbrain Cerebellum Pons Medulla oblongata (a) Figure 14.2a • Epithalamus—very small mass of tissue composed of: – Pineal gland: endocrine gland – Habenula: relay from the limbic system to the midbrain – Thin roof over the third ventricle 14-71 The Telencephalon: Cerebrum Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Central sulcus Parietal lobe Cingulate gyrus Corpus callosum Parieto–occipital sulcus Frontal lobe Occipital lobe Thalamus Anterior commissure Hypothalamus Habenula Pineal gland Epithalamus Posterior commissure Optic chiasm Mammillary body Cerebral aqueduct Pituitary gland Fourth ventricle Temporal lobe Midbrain Cerebellum Pons Medulla oblongata Figure 14.2a (a) • Cerebrum—largest and most conspicuous part of the human brain – Seat of sensory perception, memory, thought, judgment, and voluntary motor actions 14-72 The Cerebrum Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Cerebral hemispheres Rostral Caudal Central sulcus Frontal lobe Gyri Cerebrum Lateral sulcus Central sulcus Cerebellum Temporal lobe Parietal lobe Brainstem Occipital lobe Longitudinal fissure (a) Superior view Figure 14.1a,b Spinal cord (b) Lateral view • Two cerebral hemispheres divided by longitudinal fissure – – – – Connected by white fibrous tract, the corpus callosum Gyri and sulci: increase amount of cortex in the cranial cavity Gyri increases surface area for information-processing capability Some sulci divide each hemisphere into five lobes named for the cranial bones overlying them 14-73 The Cerebrum • Frontal lobe – Voluntary motor functions – Motivation, foresight, planning, memory, mood, emotion, social judgment, and aggression • Parietal lobe – Receives and integrates general sensory information, taste, and some visual Precentral gyrus processing • Occipital lobe – Primary visual center of Central sulcus brain Insula • Temporal lobe – Areas for hearing, smell, learning, memory, and some aspects of vision and emotion • Insula (hidden by other regions) – Understanding spoken language, taste and sensory information from visceral receptors Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Rostral Caudal Frontal lobe Parietal lobe Postcentral gyrus Occipital lobe Lateral sulcus Temporal lobe 14-74 Tracts of Cerebral White Matter Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Association tracts Projection tracts Frontal lobe Corpus callosum Parietal lobe Temporal lobe Occipital lobe (a) Sagittal section Longitudinal fissure Corpus callosum Commissural tracts Lateral ventricle Thalamus Basal nuclei Third ventricle Mammillary body Cerebral peduncle Pons Projection tracts Pyramid Decussation in pyramids Medulla oblongata (b) Frontal section Figure 14.14 14-75 The Cerebral White Matter • Most of the volume of cerebrum is white matter – Glia and myelinated nerve fibers transmitting signals from one region of the cerebrum to another and between cerebrum and lower brain centers 14-76 The Cerebral White Matter • Three types of tracts – Projection tracts • Extends vertically between higher and lower brain and spinal cord centers • Carries information between cerebrum and rest of the body – Commissural tracts • Cross from one cerebral hemisphere through bridges called commissures – Most pass through corpus callosum – Anterior and posterior commissures – Enables the two sides of the cerebrum to communicate with each other 14-77 The Cerebral White Matter Cont. – Association tracts • Connect different regions within the same cerebral hemisphere • Long association fibers—connect different lobes of a hemisphere to each other • Short association fibers—connect different gyri within a single lobe 14-78 The Cerebral Cortex Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Neural integration is carried out in the gray matter of the cerebrum Figure 14.15 Cortical surface • Cerebral gray matter found in three places – Cerebral cortex – Basal nuclei – Limbic system I Small pyramidal cells II III Stellate cells IV Large pyramidal cells V VI White matter 14-79 The Cerebral Cortex Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Cerebral cortex—layer covering the surface of the hemispheres – Only 2 to 3 mm thick – Cortex constitutes about 40% of brain mass – Contains 14 to 16 billion neurons Figure 14.15 Cortical surface I Small pyramidal cells II III Stellate cells IV Large pyramidal cells V VI White matter 14-80 The Cerebral Cortex • Contains two principal types of neurons – Stellate cells • Have spheroid somas with dendrites projecting in all directions • Receive sensory input and process information on a local level – Pyramidal cells • Tall, and conical, with apex toward the brain surface • A thick dendrite with many branches with small, knobby dendritic spines • Include the output neurons of the cerebrum • Only neurons that leave the cortex and connect with other parts of the CNS. 14-81 The Cerebral Cortex • Neocortex—six-layered tissue that constitutes about 90% of the human cerebral cortex – Relatively recent in evolutionary origin 14-82 The Basal Nuclei Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Cerebrum Corpus callosum Lateral ventricle Thalamus Internal capsule Caudate nucleus Insula Putamen Lentiform nucleus Third ventricle Globus pallidus Hypothalamus Subthalamic nucleus Optic tract Pituitary gland Corpus striatum Figure 14.16 • Basal nuclei—masses of cerebral gray matter buried deep in the white matter, lateral to the thalamus – Receives input from the substantia nigra of the midbrain and the motor areas of the cortex – Send signals back to both these locations – Involved in motor control 14-83 The Basal Nuclei • At least three brain centers form basal nuclei – Caudate nucleus – Putamen – Globus pallidus • Lentiform nucleus—putamen and globus pallidus collectively • Corpus striatum—putamen and caudate nucleus collectively 14-84 The Limbic System • Limbic system—important center of emotion and learning • Most anatomically prominent components are: – Cingulate gyrus: arches over the top of the corpus callosum in the frontal and parietal lobes – Hippocampus: in the medial temporal lobe; memory – Amygdala: immediately rostral to the hippocampus; emotion Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Medial prefrontal cortex Corpus callosum Cingulate gyrus Fornix Orbitofrontal cortex Mammillary body Hippocampus Thalamic nuclei Basal nuclei Amygdala Temporal lobe Figure 14.17 14-85 The Limbic System • Limbic system components are connected through a complex loop of fiber tracts allowing for somewhat circular patterns of feedback • Limbic system structures have centers for both gratification and aversion – Gratification: sensations of pleasure or reward – Aversion: sensations of fear or sorrow Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Medial prefrontal cortex Corpus callosum Cingulate gyrus Fornix Orbitofrontal cortex Mammillary body Hippocampus Thalamic nuclei Basal nuclei Amygdala Temporal lobe Figure 14.17 14-86 Integrative Functions of the Brain • Expected Learning Outcomes – List the types of brain waves and discuss their relationship to mental states. – Describe the stages of sleep, their relationship to the brain waves, and the neural mechanisms of sleep. – Identify the brain regions concerned with consciousness and thought, memory, emotion, sensation, motor control, and language. – Discuss the functional differences between the right and left cerebral hemispheres. 14-87 Integrative Functions of the Brain • Higher brain functions—sleep, memory, cognition, emotion, sensation, motor control, and language • Involve interactions between cerebral cortex and basal nuclei, brainstem, and cerebellum • Functions of the brain do not have easily defined anatomical boundaries • Integrative functions of the brain focus mainly on the cerebrum, but involve combined action of multiple brain levels 14-88 The Electroencephalogram • Alpha waves 8 to 13 Hz – Awake and resting with eyes closed and mind wandering – Suppressed when eyes open or performing a mental task • Beta waves 14 to 30 Hz – Eyes open and performing mental tasks – Accentuated during mental activity and sensory stimulation • Theta waves 4 to 7 Hz – Drowsy or sleeping adults – If awake and under emotional stress • Delta waves (high amplitude) <3.5 Hz – Deep sleep in adults 14-89 The Electroencephalogram Copyright © The McGraw-Hill Companies, Inc. 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Alpha () Beta () Theta () Delta (δ) Figure 14.18a (b) 1 second Figure 14.18b • Electroencephalogram (EEG)—monitors surface electrical activity of the brain waves – Useful for studying normal brain functions as sleep and consciousness – In diagnosis of degenerative brain diseases, metabolic abnormalities, brain tumors, etc. 14-90 The Electroencephalogram Cont. • Brain waves—rhythmic voltage changes resulting from synchronized postsynaptic potentials at the superficial layer of the cerebral cortex – Four types distinguished by amplitude (mV) and frequency (Hz) • Persistent absence of brain waves is common clinical and legal criterion of brain death 14-91 Sleep • Sleep occurs in cycles called circadian rhythms – Events that reoccur at intervals of about 24 hours • Sleep—temporary state of unconsciousness from which one can awaken when stimulated – Characterized by stereotyped posture • Lying down with eyes closed – Sleep paralysis: inhibition of muscular activity – Resembles unconsciousness but can be aroused by sensory stimulation – Coma or hibernation: states of prolonged unconsciousness where individuals cannot be aroused by sensory stimulation 14-92 Sleep Cont. • Restorative effect – Brain glycogen and ATP levels increase in non-REM sleep – Memories strengthened in REM sleep • Synaptic connections reinforced • Four stages of sleep – Stage 1 • Feel drowsy, close eyes, begin to relax • Often feel drifting sensation, easily awakened if stimulated • Alpha waves dominate EEG 14-93 Sleep Cont. – Stage 2 • Pass into light sleep • EEG declines in frequency but increases in amplitude • Exhibits sleep spindles—high spikes resulting from interactions between neurons of the thalamus and cerebral cortex – Stage 3 • • • • Moderate to deep sleep About 20 minutes after stage 1 Theta and delta waves appear Muscles relax and vital signs fall (body temperature, blood pressure, heart and respiratory rates) 14-94 Sleep Cont. – Stage 4 • Called slow-wave sleep (SWS)—EEG dominated by lowfrequency, high-amplitude delta waves • Muscles now very relaxed, vital signs at their lowest, and more difficult to awaken • About five times a night, a sleeper backtracks from stage 3 or 4 to stage 2 – Exhibits bouts of rapid eye movement (REM) sleep, eyes oscillate back and forth – Also called paradoxical sleep, because EEG resembles the waking state, but sleeper is harder to arouse than any other stage 14-95 Sleep Cont. – Vital signs increase, brain uses more oxygen than when awake – Sleep paralysis stronger to prevent sleeper from acting out dreams • Dreams occur in both REM and non-REM sleep – REM tends to be longer and more vivid • Parasympathetic nervous system active during REM sleep – Causing constriction of the pupils – Erection of the penis and clitoris 14-96 Sleep • Rhythm of sleep is controlled by a complex interaction between the cerebral cortex, thalamus, hypothalamus, and reticular formation – Arousal induced in the upper reticular formation, near junction of pons and midbrain – Sleep induced by nuclei below pons, and in ventrolateral preoptic nucleus in the hypothalamus 14-97 Sleep • Suprachiasmatic nucleus (SCN)—another important control center for sleep – Above optic chiasma in anterior hypothalamus – Input from eye allows SCN to synchronize multiple body rhythms with external rhythms of night and day • Sleep, body temperature, urine production, secretion, and other functions 14-98 Sleep • Sleep has a restorative effect, and sleep deprivation can be fatal to experimental animals – Bed rest alone does not have the restorative effect of sleep— why must we lose consciousness? – Sleep may be the time to replenish such energy sources as glycogen and ATP – REM sleep may consolidate and strengthen memories by reinforcing some synapses, and eliminating others 14-99 Sleep Stages and Brain Activity Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Awake Sleep spindles REM sleep Stage Stage 1 Drowsy Stage 2 Light sleep Stage 3 Moderate to deep sleep Stage 4 Deepest sleep 0 10 20 30 40 50 60 70 Time (min.) (a) One sleep cycle Awake EEG stages REM REM REM REM REM Stage 1 Stage 2 Stage 3 Stage 4 0 1 (b) Typical 8-hour sleep period 2 3 4 5 6 7 8 Time (hours) Figure 14.19 14-100 Cognition • Cognition—the range of mental processes by which we acquire and use knowledge – Such as sensory perception, thought, reasoning, judgment, memory, imagination, and intuition • Association areas of cerebral cortex have above functions – Constitute about 75% of all brain tissue 14-101 Cognition • Studies of patients with brain lesions, cancer, stroke, and trauma yield information on cognition – Parietal lobe association area: perceiving stimuli • Contralateral neglect syndrome—unaware of objects on opposite side of the body – Temporal lobe association area: identifying stimuli • Agnosia—inability to recognize, identify, and name familiar objects • Prosopagnosia—person cannot remember familiar faces – Frontal lobe association area: planning our responses and personality; inability to execute appropriate behavior 14-102 Memory • Information management entails: – Learning: acquiring new information – Memory: information storage and retrieval – Forgetting: eliminating trivial information; as important as remembering • Amnesia—defects in declarative memory: inability to describe past events • Procedural memory—ability to tie one’s shoes – Anterograde amnesia: unable to store new information – Retrograde amnesia: person cannot recall things known before the injury 14-103 Memory • Hippocampus—important memory-forming center – Does not store memories – Organizes sensory and cognitive information into a unified long-term memory – Memory consolidation: the process of “teaching the cerebral cortex” until a long-term memory is established – Long-term memories are stored in various areas of the cerebral cortex 14-104 Memory Cont. – Vocabulary and memory of familiar faces stored in superior temporal lobe – Memories of one’s plans and social roles stored in the prefrontal cortex • Cerebellum—helps learn motor skills • Amygdala—emotional memory 14-105 Accidental Lobotomy • Phineas Gage—railroad construction worker; severe injury with metal rod • Injury to the ventromedial region of both frontal lobes • Extreme personality change – Fitful, irreverent, grossly profane – Opposite of previous personality • Prefrontal cortex functions – Planning, moral judgment, and emotional control Figure 14.20 14-106 Emotion • Emotional feelings and memories are interactions between prefrontal cortex and diencephalon • Prefrontal cortex—seat of judgment, intent, and control over expression of emotions • Feelings come from hypothalamus and amygdala – Nuclei generate feelings of fear or love 14-107 Emotion • Amygdala receives input from sensory systems – Role in food intake, sexual behavior, and drawing attention to novel stimuli – One output goes to hypothalamus influencing somatic and visceral motor systems • Heart races, raises blood pressure, makes hair stand on end, induces vomiting – Other output to prefrontal cortex important in controlling expression of emotions • Ability to express love, control anger, or overcome fear • Behavior shaped by learned associations between stimuli, our responses to them, and the reward or punishment that results 14-108 Sensation • Primary sensory cortex—sites where sensory input is first received and one becomes conscious of the stimulus • Association areas nearby to sensory areas that process and interpret that sensory information – Primary visual cortex is bordered by visual association area: interprets and makes cognitive sense of visual stimuli Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Anterior Precentral gyrus Frontal lobe Central sulcus Parietal lobe Postcentral gyrus – Multimodal association areas: receive input from multiple senses and integrate this into an overall perception of our surroundings Occipital lobe Posterior (a) Figure 14.22a 14-109 The Special Senses • Special senses—limited to the head and employ relatively complex sense organs • Primary cortices and association areas listed below • Vision – Visual primary cortex in far posterior region of occipital lobe – Visual association area: anterior, and occupies all the remaining occipital lobe • Much of inferior temporal lobe deals with facial recognition and other familiar objects • Hearing – Primary auditory cortex in the superior region of the temporal lobe and insula – Auditory association area: temporal lobe deep and inferior to primary auditory cortex • Recognizes spoken words, a familiar piece of music, or a voice on the phone 14-110 The Special Senses • Equilibrium – Signals for balance and sense of motion project mainly to the cerebellum and several brainstem nuclei concerned with head and eye movements and visceral functions – Association cortex in the roof of the lateral sulcus near the lower end of the central sulcus • Seat of consciousness of our body movements and orientation in space • Taste and smell – Gustatory (taste) signals received by primary gustatory cortex in inferior end of the postcentral gyrus of the parietal lobe and anterior region of insula – Olfactory (smell) signals received by the primary olfactory cortex in the medial surface of the temporal lobe and inferior surface of the frontal lobe 14-111 The General Senses • General (somesthetic, somatosensory, or somatic) senses—distributed over the entire body and employ relatively simple receptors – Senses of touch, pressure, stretch, movement, heat, cold, and pain • Several cranial nerves carry general sensations from head • Ascending tracts bring general sensory information from the rest of the body – Thalamus processes the input – Selectively relays signals to the postcentral gyrus • Fold of the cerebrum that lies immediately caudal to the central sulcus and forms the rostral border of the parietal lobe – Primary somesthetic cortex is the cortex of the postcentral gyrus – Somesthetic association area: caudal to the gyrus and in the roof of the lateral sulcus 14-112 The General Senses • Awareness of stimulation occurs in primary somesthetic cortex • Making cognitive sense of the stimulation occurs in the somesthetic association area • Because of decussation, the right postcentral gyrus receives input from the left side of the body and vice versa • Sensory homunculus—upside-down sensory map of the contralateral side of the body • Somatotopy—point-for-point correspondence between an area of the body and an area of the CNS 14-113 The General Senses • Sensory homunculus— diagram of the primary somesthetic cortex which resembles an upside-down sensory map of the contralateral side of the body Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Figure 14.22b II III I IV V II V IV III Toes Genitalia • Shows receptors in the lower limbs projecting to the superior and medial parts of the gyrus Teeth, gums Tongue Abdominal viscera • Shows receptors from the face projecting to the inferior and lateral parts Insula Lateral • Somatotopy—point-to-point correspondence between an area of the body and an area of the CNS Viscerosensory area Lateral sulcus Medial (b) 14-114 Functional Regions of the Cerebral Cortex Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Primary somatosensory cortex Primary motor cortex Somatosensory association area Motor association area Primary gustatory cortex Wernicke area Broca area Visual association area Prefrontal cortex Primary visual cortex Olfactory association area Primary auditory cortex Auditory association area Figure 14.21 14-115 Motor Control • The intention to contract a muscle begins in motor association (premotor) area of frontal lobes – Where we plan our behavior – Where neurons compile a program for degree and sequence of muscle contraction required for an action – Program transmitted to neurons of the precentral gyrus (primary motor area) • Most posterior gyrus of the frontal lobe 14-116 Motor Control • Premotor area (cont.) – Neurons send signals to the brainstem and spinal cord – Ultimately resulting in muscle contraction – Precentral gyrus also exhibits somatotopy • Neurons for toe movement are deep in the longitudinal fissure of the medial side of the gyrus • The summit of the gyrus controls the trunk, shoulder, and arm • The inferolateral region controls the facial muscles – Motor homunculus has a distorted look because the amount of cortex devoted to a given body region is proportional to the number of muscles and motor units in that body region 14-117 Motor Control • Pyramidal cells of the precentral gyrus are called upper motor neurons – – – – – Their fibers project caudally About 19 million fibers ending in nuclei of the brainstem About 1 million forming the corticospinal tracts Most fibers decussate in lower medulla oblongata Form lateral corticospinal tracts on each side of the spinal cord • In the brainstem or spinal cord, the fibers from upper motor neurons synapse with lower motor neurons whose axons innervate the skeletal muscles • Basal nuclei and cerebellum are also important in muscle control 14-118 • Basal nuclei Motor Control – Determines the onset and cessation of intentional movements • Repetitive hip and shoulder movements in walking – Highly practiced, learned behaviors that one carries out with little thought • Writing, typing, driving a car – Lies in a feedback circuit from the cerebrum, to the basal nuclei, to the thalamus, and back to the cerebrum – Dyskinesias: movement disorders caused by lesions in the basal nuclei • Cerebellum – – – – – – – Highly important in motor coordination Aids in learning motor skills Maintains muscle tone and posture Smoothes muscle contraction Coordinates eye and body movements Coordinates the motions of different joints with each other Ataxia: clumsy, awkward gait 14-119 Motor Homunculus Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. V IV III II Toes I Vocalization Salivation Mastication Swallowing Lateral Medial Figure 14.23b (b) 14-120 Motor Pathways Involving the Cerebellum Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. (a) Input to cerebellum (b) Output from cerebellum Motor cortex Cerebrum Cerebrum Cerebellum Reticular formation Brainstem Cerebellum Brainstem Eye Inner ear Spinocerebellar tracts of spinal cord Muscle and joint proprioceptors Reticulospinal and vestibulospinal tracts of spinal cord Figure 14.24 Limb and postural muscles 14-121 Language • Language include several abilities: reading, writing, speaking, and understanding words assigned to different regions of the cerebral cortex • Wernicke area – Permits recognition of spoken and written language and creates plan of speech – When we intend to speak, Wernicke area formulates phrases according to learned rules of grammar – Transmits plan of speech to Broca area • Broca area – Generates motor program for the muscles of the larynx, tongue, cheeks, and lips – Transmits program to primary motor cortex for commands to the lower motor neurons that supply relevant muscles • Affective language area lesions produce aprosody—flat emotionless speech 14-122 Language Centers of the Left Hemisphere Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Anterior Posterior Precentral gyrus Postcentral gyrus Speech center of primary motor cortex Angular gyrus Primary auditory cortex (in lateral sulcus) Primary visual cortex Broca area Wernicke area Figure 14.25 14-123 Aphasia • Aphasia—any language deficit from lesions in same hemisphere (usually left) containing the Wernicke and Broca areas • Nonfluent (Broca) aphasia – Lesion in Broca area – Slow speech, difficulty in choosing words, using words that only approximate the correct word • Fluent (Wernicke) aphasia – Lesion in Wernicke area – Speech normal and excessive, but uses senseless jargon – Cannot comprehend written and spoken words • Anomic aphasia – Can speak normally and understand speech, but cannot identify written words or pictures 14-124 Cerebral Lateralization Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Left hemisphere Right hemisphere Olfaction, right nasal cavity Olfaction, left nasal cavity Anterior Verbal memory Memory for shapes Speech (Limited language comprehension, mute) Left hand motor control Right hand motor control Feeling shapes with left hand Feeling shapes with right hand Hearing nonvocal sounds (left ear advantage) Hearing vocal sounds (right ear advantage) Musical ability Rational, symbolic thought Intuitive, nonverbal thought Superior language comprehension Superior recognition of faces and spatial relationships Vision, right field Posterior Figure 14.26 Vision, left field 14-125 Cerebral Lateralization • Cerebral lateralization—the difference in the structure and function of the cerebral hemispheres • Left hemisphere—categorical hemisphere – Specialized for spoken and written language – Sequential and analytical reasoning (math and science) – Breaks information into fragments and analyzes it in a linear way • Right hemisphere—representational hemisphere – – – – – Perceives information in a more integrated holistic way Seat of imagination and insight Musical and artistic skill Perception of patterns and spatial relationships Comparison of sights, sounds, smells, and taste 14-126 Cerebral Lateralization • Highly correlated with handedness – Left hemisphere is the categorical one in 96% of right-handed people • Right hemisphere in 4% – Left-handed people: right hemisphere is categorical in 15% and left in 70% • Lateralization develops with age – Males exhibit more lateralization than females and suffer more functional loss when one hemisphere is damaged 14-127 The Cranial Nerves • Expected Learning Outcomes – List the 12 cranial nerves by name and number. – Identify where each cranial nerve originates and terminates. – State the functions of each cranial nerve. 14-128 The Cranial Nerves • Brain must communicate with rest of body – Most of the input and output travels by way of the spinal cord – 12 pairs of cranial nerves arise from the base of the brain – Exit the cranium through foramina – Lead to muscles and sense organs located mainly in the head and neck 14-129 Cranial Nerve Pathways • Most motor fibers of the cranial nerves begin in nuclei of brainstem and lead to glands and muscles • Sensory fibers begin in receptors located mainly in head and neck and lead mainly to the brainstem – Sensory fibers for proprioception may travel to brain in a different nerve from motor nerve 14-130 Cranial Nerve Pathways • Most cranial nerves carry fibers between brainstem and ipsilateral receptors and effectors – Lesion in left brainstem causes sensory or motor deficit on same side • Exceptions: optic nerve where half the fibers decussate, and trochlear nerve where all efferent fibers lead to a muscle of the contralateral eye 14-131 Cranial Nerve Classification • Some cranial nerves are classified as motor, some sensory, others mixed – Sensory (I, II, and VIII) – Motor (III, IV, VI, XI, and XII) • Stimulate muscle but also contain fibers of proprioception – Mixed (V, VII, IX, X) • Sensory functions may be quite unrelated to their motor function – Facial nerve (VII) has sensory role in taste and motor role in facial expression 14-132 The Cranial Nerves Figure 14.27a,b Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Frontal lobe Frontal lobe Longitudinal fissure Cranial nerves: Olfactory bulb (from olfactory nerve, I) Olfactory tract Olfactory tract Optic chiasm Optic nerve (II) Temporal lobe Temporal lobe Oculomotor nerve (III) Trochlear nerve (IV) Infundibulum Trigeminal nerve (V) Optic chiasm Pons Abducens nerve (VI) Facial nerve (VII) Pons Medulla Vestibulocochlear nerve (VIII) Glossopharyngeal nerve (IX) Cerebellum Vagus nerve (X) Medulla oblongata Hypoglossal nerve (XII) Accessory nerve (XI) Cerebellum Spinal cord Spinal cord (a) (b) b: © The McGraw-Hill Companies, Inc./Rebecca Gray, photographer/DonKincaid, dissections 14-133 The Olfactory Nerve (I) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Olfactory bulb Olfactory tract Cribriform plate of ethmoid bone Fascicles of olfactory nerve (I) Nasal mucosa Figure 14.28 • Sense of smell • Damage causes impaired sense of smell 14-134 The Optic Nerve (II) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Eyeball Optic nerve (II) Optic chiasm Optic tract Pituitary gland Figure 14.29 • Provides vision • Damage causes blindness in part or all of visual field 14-135 The Oculomotor Nerve (III) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Oculomotor nerve (III): Superior branch Inferior branch Ciliary ganglion Superior orbital fissure Figure 14.30 • Controls muscles that turn the eyeball up, down, and medially, as well as controlling the iris, lens, and upper eyelid • Damage causes drooping eyelid, dilated pupil, double vision, difficulty focusing, and inability to move eye in certain directions 14-136 The Trochlear Nerve (IV) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Superior oblique muscle Superior orbital fissure Trochlear nerve (IV) Figure 14.31 • Eye movement (superior oblique muscle) • Damage causes double vision and inability to rotate eye inferolaterally 14-137 The Trigeminal Nerve (V) • Largest of the cranial nerves • Most important sensory nerve of the face Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Superior orbital fissure Ophthalmic division (V1) Trigeminal ganglion Trigeminal nerve (V) Maxillary division (V2) Foramen ovale Mandibular division (V3) Foramen rotundum Infraorbital nerve Superior alveolar nerves Lingual nerve • Forks into three divisions Anterior trunk of V3 to chewing muscles Inferior alveolar nerve – Ophthalmic division (V1): sensory – Maxillary division (V2): sensory – Mandibular division (V3): mixed Temporalis muscle Lateral pterygoid muscle Medial pterygoid muscle V1 Masseter muscle V2 Anterior belly of digastric muscle V3 Motor branches of the mandibular division (V3) Distribution of sensory fibers of each division Figure 14.32 14-138 The Abducens Nerve (VI) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Lateral rectus muscle Superior orbital fissure Abducens nerve (VI) Figure 14.33 • Provides eye movement (lateral rectus m.) • Damage results in inability to rotate eye laterally and at rest eye rotates medially 14-139 The Facial Nerve (VII) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Facial nerve (VII) Internal acoustic meatus Geniculate ganglion Pterygopalatine ganglion Lacrimal (tear) gland Chorda tympani branch (taste and salivation) Submandibular ganglion Sublingual gland Parasympathetic fibers Submandibular gland Stylomastoid foramen Motor branch to muscles of facial expression Figure 14.34a (a) • Motor—major motor nerve of facial muscles: facial expressions; salivary glands and tear, nasal, and palatine glands • Sensory—taste on anterior two-thirds of tongue • Damage produces sagging facial muscles and disturbed sense of taste (no sweet and salty) 14-140 Five Branches of Facial Nerve Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Temporal Zygomatic Buccal Buccal Mandibular Cervical (b) (c) c: © The McGraw-Hill Companies, Inc./Joe DeGrandis, photographer Figure 14.34b,c Clinical test: test anterior two-thirds of tongue with substances such as sugar, salt, vinegar, and quinine; test response of tear glands to ammonia fumes; test motor functions by asking subject to close eyes, smile, whistle, frown, raise eyebrows, etc. 14-141 The Vestibulocochlear Nerve (VIII) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Semicircular ducts Vestibular ganglia Vestibular nerve Cochlear nerve Vestibulocochlear nerve (VIII) Internal acoustic meatus Cochlea Vestibule Figure 14.35 • Nerve of hearing and equilibrium • Damage produces deafness, dizziness, nausea, loss of balance, and nystagmus (involuntary rhythmic oscillation of the eyes from side to side) 14-142 The Glossopharyngeal Nerve (IX) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Swallowing, salivation, gagging, control of BP and respiration Glossopharyngeal nerve (IX) Jugular foramen Superior ganglion Inferior ganglion Otic ganglion Parotid salivary gland • Sensations from posterior one-third of tongue Carotid sinus Pharyngeal muscles • Damage results in loss of bitter and sour taste and impaired swallowing Figure 14.36 14-143 The Vagus Nerve (X) • Most extensive distribution of any cranial nerve Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • Major role in the control of cardiac, pulmonary, digestive, and urinary function Jugular foramen Pharyngeal nerve Laryngeal nerve Carotid sinus Vagus nerve (X) • Swallowing, speech, regulation of viscera Lung • Damage causes hoarseness or loss of voice, impaired swallowing, and fatal if both are cut Heart Spleen Liver Kidney Stomach Colon (proximal portion) Small intestine Figure 14.37 14-144 The Accessory Nerve (XI) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Jugular foramen Vagus nerve Accessory nerve (XI) Foramen magnum Sternocleidomastoid muscle Spinal nerves C3 and C4 Trapezius muscle Posterior view Figure 14.38 • Swallowing; head, neck, and shoulder movement – Damage causes impaired head, neck, shoulder movement; head turns toward injured side 14-145 The Hypoglossal Nerve (XII) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Hypoglossal canal Intrinsic muscles of the tongue Extrinsic muscles of the tongue Hypoglossal nerve (XII) Figure 14.39 • Tongue movements for speech, food manipulation, and swallowing – If both are damaged: cannot protrude tongue – If one side is damaged: tongue deviates toward injured side; see ipsilateral atrophy 14-146 Cranial Nerve Disorders • Trigeminal neuralgia (tic douloureux) – Recurring episodes of intense stabbing pain in trigeminal nerve area (near mouth or nose) – Pain triggered by touch, drinking, washing face – Treatment may require cutting nerve • Bell palsy – Degenerative disorder of facial nerve causes paralysis of facial muscles on one side – May appear abruptly with full recovery within 3 to 5 weeks 14-147 Images of the Mind • Positron emission tomography (PET) and MRI visualize increases in blood flow when brain areas are active – Injection of radioactively labeled glucose • Busy areas of brain “light up” • Functional magnetic resonance imaging (fMRI) looks at increase in blood flow to an area (additional glucose is needed in active area)—magnetic properties of hemoglobin depend on how much oxygen is bound to it (additional oxygen is there due to additional blood flow) – Quick, safe, and accurate method to see brain function 14-148 Images of the Mind Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Rostral Caudal Primary auditory cortex Visual cortex 1 The word car is seen in the visual cortex. Wernicke area 2 Wernicke area conceives of the verb drive to go with it. Premotor area Primary motor cortex Broca area 3 Broca area compiles a motor program to speak the word drive. 4 The primary motor cortex executes the program and the word is spoken. Figure 14.40 14-149