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CAMPBELL BIOLOGY TENTH EDITION Reece • Urry • Cain • Wasserman • Minorsky • Jackson 49 Nervous Systems Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick © 2014 Pearson Education, Inc. Command and Control Center The human brain contains about 100 billion neurons, organized into circuits more complex than the most powerful supercomputers Powerful imaging techniques allow researchers to monitor multiple areas of the brain while the subject performs various tasks A recent advance uses expression of combinations of colored proteins in brain cells, a technique called “brainbow” © 2014 Pearson Education, Inc. Figure 49.1 © 2014 Pearson Education, Inc. Concept 49.1: Nervous systems consist of circuits of neurons and supporting cells By the time of the Cambrian explosion more than 500 million years ago, specialized systems of neurons had appeared that enables animals to sense their environments and respond rapidly © 2014 Pearson Education, Inc. The simplest animals with nervous systems, the cnidarians, have neurons arranged in nerve nets A nerve net is a series of interconnected nerve cells More complex animals have nerves, in which the axons of multiple neurons are bundled together Nerves channel and organize information flow through the nervous system © 2014 Pearson Education, Inc. Figure 49.2 Eyespot Nerve net (a) Hydra (cnidarian) Brain Brain Nerve cords Radial nerve Nerve ring Ventral nerve cord Transverse nerve (b) Sea star (echinoderm) Segmental ganglia (c) Planarian (flatworm) (d) Leech (annelid) Brain Ganglia Brain Ventral nerve cord Segmental ganglia (e) Insect (arthropod) © 2014 Pearson Education, Inc. Anterior nerve ring Brain Ganglia Spinal cord (dorsal nerve cord) Sensory ganglia Longitudinal nerve cords (f) Chiton (mollusc) (g) Squid (mollusc) (h) Salamander (vertebrate) Figure 49.2a Nerve net (a) Hydra (cnidarian) Radial nerve Nerve ring (b) Sea star (echinoderm) Eyespot Brain Nerve cords Transverse nerve (c) Planarian (flatworm) © 2014 Pearson Education, Inc. Brain Ventral nerve cord Segmental ganglia (d) Leech (annelid) Figure 49.2a, b Nerve net (a) Hydra (cnidarian) © 2014 Pearson Education, Inc. Radial nerve Nerve ring (b) Sea star (echinoderm) Bilaterally symmetrical animals exhibit cephalization, the clustering of sensory organs at the front end of the body The simplest cephalized animals, flatworms, have a central nervous system (CNS) The CNS consists of a brain and longitudinal nerve cords The peripheral nervous system (PNS) consists of neurons carrying information into and out of the CNS © 2014 Pearson Education, Inc. Figure 49.2c Eyespot Brain Nerve cords Transverse nerve (c) Planarian (flatworm) © 2014 Pearson Education, Inc. Annelids and arthropods have segmentally arranged clusters of neurons called ganglia © 2014 Pearson Education, Inc. Figure 49.2d, e Brain Ventral nerve cord Segmental ganglia (d) Leech (annelid) © 2014 Pearson Education, Inc. Brain Ventral nerve cord Segmental ganglia (e) Insect (arthropod) Figure 49.2b Ganglia Brain Ventral nerve cord Segmental ganglia Anterior nerve ring Longitudinal nerve cords (e) Insect (arthropod) (f) Chiton (mollusc) Brain Brain Ganglia (g) Squid (mollusc) © 2014 Pearson Education, Inc. Spinal cord (dorsal nerve cord) Sensory ganglia (h) Salamander (vertebrate) Nervous system organization usually correlates with lifestyle Sessile molluscs (for example, clams and chitons) have simple systems, whereas more complex molluscs (for example, octopuses and squids) have more sophisticated systems © 2014 Pearson Education, Inc. Figure 49.2f, g Ganglia Anterior nerve ring Brain Ganglia Longitudinal nerve cords (f) Chiton (mollusc) © 2014 Pearson Education, Inc. (g) Squid (mollusc) In vertebrates The CNS is composed of the brain and spinal cord The peripheral nervous system (PNS) is composed of nerves and ganglia Region specialization is a hallmark of both systems © 2014 Pearson Education, Inc. Figure 49.2h Brain Spinal cord (dorsal nerve cord) Sensory ganglia (h) Salamander (vertebrate) © 2014 Pearson Education, Inc. Glia Glial cells, or glia have numerous functions to nourish, support, and regulate neurons Embryonic radial glia form tracks along which newly formed neurons migrate Astrocytes induce cells lining capillaries in the CNS to form tight junctions, resulting in a bloodbrain barrier and restricting the entry of most substances into the brain © 2014 Pearson Education, Inc. Figure 49.3 Capillary CNS Neuron PNS VENTRICLE Cilia Ependymal cells © 2014 Pearson Education, Inc. Astrocytes Oligodendrocytes Microglia Schwann cells Radial glial cells and astrocytes can both act as stem cells Researchers are trying to find a way to use neural stem cells to replace brain tissue that has ceased to function normally © 2014 Pearson Education, Inc. Figure 49.4 © 2014 Pearson Education, Inc. Organization of the Vertebrate Nervous System The CNS develops from the hollow nerve cord The cavity of the nerve cord gives rise to the narrow central canal of the spinal cord and the ventricles of the brain The canal and ventricles fill with cerebrospinal fluid, which supplies the CNS with nutrients and hormones and carries away wastes © 2014 Pearson Education, Inc. Figure 49.5 Gray matter White matter Ventricles © 2014 Pearson Education, Inc. The brain and spinal cord contain Gray matter, which consists of neuron cell bodies, dendrites, and unmyelinated axons White matter, which consists of bundles of myelinated axons © 2014 Pearson Education, Inc. Figure 49.6 Central nervous system (CNS) Brain Spinal cord Cranial nerves Ganglia outside CNS Spinal nerves © 2014 Pearson Education, Inc. Peripheral nervous system (PNS) The spinal cord conveys information to and from the brain and generates basic patterns of locomotion The spinal cord also produces reflexes independently of the brain A reflex is the body’s automatic response to a stimulus For example, a doctor uses a mallet to trigger a knee-jerk reflex © 2014 Pearson Education, Inc. Figure 49.7 Cell body of sensory neuron in dorsal root ganglion Gray matter Quadriceps muscle Spinal cord (cross section) Hamstring muscle Key Sensory neuron Motor neuron Interneuron © 2014 Pearson Education, Inc. White matter The Peripheral Nervous System The PNS transmits information to and from the CNS and regulates movement and the internal environment In the PNS, afferent neurons transmit information to the CNS and efferent neurons transmit information away from the CNS © 2014 Pearson Education, Inc. Figure 49.8 CENTRAL NERVOUS SYSTEM (information processing) PERIPHERAL NERVOUS SYSTEM Efferent neurons Afferent neurons Sensory receptors Autonomic nervous system Motor system Control of skeletal muscle Internal and external stimuli Sympathetic Parasympathetic division division Enteric division Control of smooth muscles, cardiac muscles, glands © 2014 Pearson Education, Inc. The PNS has two efferent components: the motor system and the autonomic nervous system The motor system carries signals to skeletal muscles and is voluntary The autonomic nervous system regulates smooth and cardiac muscles and is generally involuntary © 2014 Pearson Education, Inc. The autonomic nervous system has sympathetic and parasympathetic divisions The sympathetic division regulates arousal and energy generation (“fight-or-flight” response) The parasympathetic division has antagonistic effects on target organs and promotes calming and a return to “rest and digest” functions © 2014 Pearson Education, Inc. Figure 49.9 Parasympathetic division Sympathetic division Constricts pupil of eye Dilates pupil of eye Stimulates salivary gland secretion Inhibits salivary gland secretion Constricts bronchi in lungs Cervical Sympathetic ganglia Relaxes bronchi in lungs Slows heart Accelerates heart Stimulates activity of stomach and intestines Inhibits activity of stomach and intestines Thoracic Stimulates activity of pancreas Inhibits activity of pancreas Stimulates gallbladder Stimulates glucose release from liver; inhibits gallbladder Lumbar Stimulates adrenal medulla Promotes emptying of bladder Promotes erection of genitalia © 2014 Pearson Education, Inc. Inhibits emptying of bladder Sacral Synapse Promotes ejaculation and vaginal contractions Figure 49.9a Parasympathetic division Sympathetic division Constricts pupil of eye Dilates pupil of eye Stimulates salivary gland secretion Inhibits salivary gland secretion Constricts bronchi in lungs Slows heart Stimulates activity of stomach and intestines Stimulates activity of pancreas Stimulates gallbladder © 2014 Pearson Education, Inc. Cervical Sympathetic ganglia Figure 49.9b Parasympathetic division Sympathetic division Relaxes bronchi in lungs Sympathetic ganglia Accelerates heart Inhibits activity of stomach and intestines Thoracic Inhibits activity of pancreas Stimulates glucose release from liver; inhibits gallbladder Lumbar Stimulates adrenal medulla Promotes emptying of bladder Promotes erection of genitalia © 2014 Pearson Education, Inc. Inhibits emptying of bladder Sacral Synapse Promotes ejaculation and vaginal contractions Concept 49.2: The vertebrate brain is regionally specialized Specific brain structures are particularly specialized for diverse functions The vertebrate brain has three major regions: the forebrain, midbrain, and hindbrain © 2014 Pearson Education, Inc. Figure 49.UN01 Forebrain Midbrain Hindbrain Cerebellum Olfactory bulb Cerebrum © 2014 Pearson Education, Inc. The forebrain has activities including processing of olfactory input, regulation of sleep, learning, and any complex processing The midbrain coordinates routing of sensory input The hindbrain controls involuntary activities and coordinates motor activities © 2014 Pearson Education, Inc. Comparison of vertebrates shows that relative sizes of particular brain regions vary These size differences reflect the relative importance of the particular brain function Evolution has resulted in a close match between structure and function © 2014 Pearson Education, Inc. Figure 49.10 Lamprey ANCESTRAL VERTEBRATE Shark Ray-finned fish Amphibian Crocodilian Key Forebrain Midbrain Hindbrain © 2014 Pearson Education, Inc. Bird Mammal During embryonic development the anterior neural tube gives rise to the forebrain, midbrain, and hindbrain The midbrain and part of the hindbrain form the brainstem, which joins with the spinal cord at the base of the brain The rest of the hindbrain gives rise to the cerebellum The forebrain divides into the diencephelon, which forms endocrine tissues in the brain, and the telencephalon, which becomes the cerebrum © 2014 Pearson Education, Inc. Figure 49.11a © 2014 Pearson Education, Inc. Figure 49.11b Embryonic brain regions Brain structures in child and adult Telencephalon Cerebrum (includes cerebral cortex, basal nuclei) Diencephalon Diencephalon (thalamus, hypothalamus, epithalamus) Forebrain Midbrain Mesencephalon Midbrain (part of brainstem) Metencephalon Pons (part of brainstem), cerebellum Myelencephalon Medulla oblongata (part of brainstem) Hindbrain Cerebrum Mesencephalon Metencephalon Midbrain Hindbrain Diencephalon Diencephalon Myelencephalon Forebrain Embryo at 1 month © 2014 Pearson Education, Inc. Telencephalon Cerebellum Spinal cord Embryo at 5 weeks Spinal cord Child Brainstem Midbrain Pons Medulla oblongata Figure 49.11ba Embryonic brain regions Telencephalon Forebrain Diencephalon Midbrain Mesencephalon Metencephalon Hindbrain Myelencephalon Brain structures in child and adult Cerebrum (includes cerebral cortex, basal nuclei) Diencephalon (thalamus, hypothalamus, epithalamus) Midbrain (part of brainstem) Pons (part of brainstem), cerebellum Medulla oblongata (part of brainstem) © 2014 Pearson Education, Inc. Figure 49.11bb Mesencephalon Metencephalon Midbrain Hindbrain Forebrain Embryo at 1 month © 2014 Pearson Education, Inc. Diencephalon Myelencephalon Telencephalon Spinal cord Embryo at 5 weeks Figure 49.11bc Cerebrum Diencephalon Cerebellum Spinal cord Child © 2014 Pearson Education, Inc. Brainstem Midbrain Pons Medulla oblongata Figure 49.11c Left cerebral hemisphere Right cerebral hemisphere Cerebral cortex Corpus callosum Cerebrum Basal nuclei Cerebellum Adult brain viewed from the rear © 2014 Pearson Education, Inc. Figure 49.11d Diencephalon Thalamus Pineal gland Hypothalamus Pituitary gland Midbrain Pons Medulla oblongata Spinal cord © 2014 Pearson Education, Inc. Arousal and Sleep The brainstem and cerebrum control arousal and sleep The core of the brainstem has a diffuse network of neurons called the reticular formation These neurons control the timing of sleep periods characterized by rapid eye movements (REMs) and by vivid dreams Sleep is also regulated by the biological clock and regions of the forebrain that regulate intensity and duration © 2014 Pearson Education, Inc. Figure 49.12 Eye Reticular formation Input from touch, pain, and temperature receptors © 2014 Pearson Education, Inc. Input from nerves of ears Sleep is essential and may play a role in the consolidation of learning and memory Some animals have evolutionary adaptations allowing for substantial activity during sleep Dolphins sleep with one brain hemisphere at a time and are therefore able to swim while “asleep” © 2014 Pearson Education, Inc. Figure 49.13 Key Low-frequency waves characteristic of sleep High-frequency waves characteristic of wakefulness Location Left hemisphere Right hemisphere © 2014 Pearson Education, Inc. Time: 0 hours Time: 1 hour Biological Clock Regulation Cycles of sleep and wakefulness are examples of circadian rhythms, daily cycles of biological activity Such rhythms rely on a biological clock, a molecular mechanism that directs periodic gene expression and cellular activity Biological clocks are typically synchronized to light and dark cycles © 2014 Pearson Education, Inc. In mammals, circadian rhythms are coordinated by a group of neurons in the hypothalamus called the suprachiasmatic nucleus (SCN) The SCN acts as a pacemaker, synchronizing the biological clock © 2014 Pearson Education, Inc. Emotions Generation and experience of emotions involve many brain structures, including the amygdala, hippocampus, and parts of the thalamus These structures are grouped as the limbic system © 2014 Pearson Education, Inc. Figure 49.14 Thalamus Hypothalamus Olfactory bulb Amygdala © 2014 Pearson Education, Inc. Hippocampus Generating and experiencing emotion often require interactions between different parts of the brain The structure most important to the storage of emotion in the memory is the amygdala, a mass of nuclei near the base of the cerebrum © 2014 Pearson Education, Inc. Functional Imaging of the Brain Brain structures are probed and analyzed with functional imaging methods Positron-emission tomography (PET) enables a display of metabolic activity through injection of radioactive glucose Today, many studies rely on functional magnetic resonance imaging (fMRI), in which brain activity is detected through changes in local oxygen concentration © 2014 Pearson Education, Inc. Figure 49.15 Nucleus accumbens Happy music © 2014 Pearson Education, Inc. Amygdala Sad music Figure 49.15a Nucleus accumbens Happy music © 2014 Pearson Education, Inc. Figure 49.15b Amygdala Sad music © 2014 Pearson Education, Inc. The range of applications for fMRI include monitoring recovery from stroke, mapping abnormalities in migraine headaches, and increasing the effectiveness of brain surgery © 2014 Pearson Education, Inc. Concept 49.3: The cerebral cortex controls voluntary movement and cognitive functions The cerebrum, the largest structure in the human brain, is essential for language, cognition, memory, consciousness, and awareness of our surroundings Four regions, or lobes (frontal, temporal, occipital, and parietal), are landmarks for particular functions © 2014 Pearson Education, Inc. Figure 49.16 Motor cortex (control of skeletal muscles) Somatosensory cortex (sense of touch) Sensory association cortex (integration of sensory information) Frontal lobe Parietal lobe Prefrontal cortex (decision making, planning) Visual association cortex (combining images and object recognition) Broca’s area (forming speech) Temporal lobe Occipital lobe Auditory cortex (hearing) Cerebellum Wernicke’s area (comprehending language) © 2014 Pearson Education, Inc. Visual cortex (processing visual stimuli and pattern recognition) Information Processing The cerebral cortex receives input from sensory organs and somatosensory receptors Somatosensory receptors provide information about touch, pain, pressure, temperature, and the position of muscles and limbs The thalamus directs different types of input to distinct locations © 2014 Pearson Education, Inc. Information received at the primary sensory areas is passed to nearby association areas that process particular features of the input Integrated sensory information passes to the prefrontal cortex, which helps plan actions and movements In the somatosensory cortex and motor cortex, neurons are arranged according to the part of the body that generates input or receives commands © 2014 Pearson Education, Inc. Figure 49.17 Frontal lobe Parietal lobe Toes Genitalia Lips Jaw Tongue © 2014 Pearson Education, Inc. Primary motor cortex Abdominal organs Primary somatosensory cortex Language and Speech Studies of brain activity have mapped areas responsible for language and speech Patients with damage in Broca’s area in the frontal lobe can understand language but cannot speak Damage to Wernicke’s area causes patients to be unable to understand language, though they can still speak © 2014 Pearson Education, Inc. Figure 49.18 Max Hearing words Seeing words Min Speaking words © 2014 Pearson Education, Inc. Generating words Lateralization of Cortical Function The two hemispheres make distinct contributions to brain function The left hemisphere is more adept at language, math, logic, and processing of serial sequences The right hemisphere is stronger at facial and pattern recognition, spatial relations, and nonverbal thinking © 2014 Pearson Education, Inc. The differences in hemisphere function are called lateralization The two hemispheres work together by communicating through the fibers of the corpus callosum © 2014 Pearson Education, Inc. Frontal Lobe Function Frontal lobe damage may impair decision making and emotional responses but leave intellect and memory intact The frontal lobes have a substantial effect on “executive functions” © 2014 Pearson Education, Inc. Figure 49.UN03 © 2014 Pearson Education, Inc. Evolution of Cognition in Vertebrates Previous ideas that a highly convoluted neocortex is required for advanced cognition may be incorrect The anatomical basis for sophisticated information processing in birds (without a highly convoluted neocortex) appears to be the clustering of nuclei in the top or outer portion of the brain (pallium) © 2014 Pearson Education, Inc. Figure 49.19 Cerebrum (including pallium) Cerebellum Thalamus Midbrain (a) Songbird brain Cerebrum (including cerebral cortex) Thalamus Midbrain (b) Human brain © 2014 Pearson Education, Inc. Cerebellum Concept 49.4: Changes in synaptic connections underlie memory and learning Two processes dominate embryonic development of the nervous system Neurons compete for growth-supporting factors in order to survive Only half the synapses that form during embryo development survive into adulthood © 2014 Pearson Education, Inc. Neuronal Plasticity Neuronal plasticity describes the ability of the nervous system to be modified after birth Changes can strengthen or weaken signaling at a synapse Autism, a developmental disorder, involves a disruption in activity-dependent remodeling at synapses Children affected with autism display impaired communication and social interaction, as well as stereotyped, repetitive behaviors © 2014 Pearson Education, Inc. Figure 49.20 N1 N1 N2 N2 (a) Connections between neurons are strengthened or weakened in response to activity. (b) If two synapses are often active at the same time, the strength of the postsynaptic response may increase at both synapses. © 2014 Pearson Education, Inc. Memory and Learning The formation of memories is an example of neuronal plasticity Short-term memory is accessed via the hippocampus The hippocampus also plays a role in forming long-term memory, which is later stored in the cerebral cortex Some consolidation of memory is thought to occur during sleep © 2014 Pearson Education, Inc. Long-Term Potentiation In the vertebrate brain, a form of learning called long-term potentiation (LTP) involves an increase in the strength of synaptic transmission LTP involves glutamate receptors If the presynaptic and postsynaptic neurons are stimulated at the same time, the set of receptors present on the postsynaptic membranes changes © 2014 Pearson Education, Inc. Figure 49.21 PRESYNAPTIC NEURON Glutamate NMDA receptor (open) POSTSYNAPTIC NEURON Ca2+ Na+ Mg2+ Stored AMPA receptor 1 NMDA receptor (closed) (a) Synapse prior to long-term potentiation (LTP) 3 (b) Establishing LTP 3 1 2 Depolarization (c) Synapse exhibiting LTP © 2014 Pearson Education, Inc. 4 Action potential 2 Figure 49.21a PRESYNAPTIC NEURON Glutamate NMDA receptor (open) POSTSYNAPTIC NEURON Ca2+ Na+ Mg2+ Stored AMPA receptor NMDA receptor (closed) (a) Synapse prior to long-term potentiation (LTP) © 2014 Pearson Education, Inc. Figure 49.21b 1 3 (b) Establishing LTP © 2014 Pearson Education, Inc. 2 Figure 49.21c 3 1 2 Depolarization (c) Synapse exhibiting LTP © 2014 Pearson Education, Inc. 4 Action potential Concept 49.5: Many nervous system disorders can be explained in molecular terms Disorders of the nervous system include schizophrenia, depression, drug addiction, Alzheimer’s disease, and Parkinson’s disease These are a major public health problem Genetic and environmental factors contribute to diseases of the nervous system To distinguish between genetic and environmental variables, scientists often carry out family studies © 2014 Pearson Education, Inc. Figure 49.22 40 Genes shared with relatives of person with schizophrenia 12.5% (3rd-degree relative) 25% (2nd-degree relative) 50% (1st-degree relative) 100% 30 20 Relationship to person with schizophrenia © 2014 Pearson Education, Inc. Identical twin Fraternal twin Child Full sibling Parent Half sibling Grandchild Nephew/niece Individual, general population 0 Uncle/aunt 10 First cousin Risk of developing schizophrenia (%) 50 Schizophrenia About 1% of the world’s population suffers from schizophrenia Schizophrenia is characterized by hallucinations, delusions, and other symptoms Evidence suggests that schizophrenia affects neuronal pathways that use dopamine as a neurotransmitter © 2014 Pearson Education, Inc. Depression Two broad forms of depressive illness are known In major depressive disorder, patients have a persistent lack of interest or pleasure in most activities Bipolar disorder is characterized by manic (highmood) and depressive (low-mood) phases Treatments for these types of depression include drugs such as Prozac, which increase the activity of biogenic amines in the brain © 2014 Pearson Education, Inc. The Brain’s Reward System and Drug Addiction The brain’s reward system rewards motivation with pleasure Some drugs are addictive because they increase activity of the brain’s reward system These drugs include cocaine, amphetamine, heroin, alcohol, and tobacco Drug addiction is characterized by compulsive consumption and an inability to control intake © 2014 Pearson Education, Inc. Addictive drugs enhance the activity of the dopamine pathway Drug addiction leads to long-lasting changes in the reward circuitry that cause craving for the drug As researchers expand their understanding of the brain’s reward system, there is hope that their insights will lead to better prevention and treatment of drug addiction © 2014 Pearson Education, Inc. Figure 49.23 Nicotine stimulates dopaminereleasing VTA neuron. Inhibitory neuron Dopaminereleasing VTA neuron Opium and heroin decrease activity of inhibitory neuron. Cocaine and amphetamines block removal of dopamine from synaptic cleft. Cerebral neuron of reward pathway © 2014 Pearson Education, Inc. Reward system response Alzheimer’s Disease Alzheimer’s disease is a mental deterioration characterized by confusion and memory loss Alzheimer’s disease is caused by the formation of neurofibrillary tangles and amyloid plaques in the brain There is no cure for this disease though some drugs are effective at relieving symptoms © 2014 Pearson Education, Inc. Figure 49.24 Amyloid plaque © 2014 Pearson Education, Inc. Neurofibrillary tangle 20 µm Parkinson’s Disease Parkinson’s disease is a motor disorder caused by death of dopamine-secreting neurons in the midbrain It is characterized by muscle tremors, flexed posture, and a shuffling gait There is no cure, although drugs and various other approaches are used to manage symptoms © 2014 Pearson Education, Inc. Figure 49.UN04 Dopa decarboxylase L-dopa © 2014 Pearson Education, Inc. Dopamine Circadian cycle period (hours) Figure 49.UN02 Wild-type hamster 𝜏 hamster Wild-type hamster with SCN from 𝜏 hamster 𝜏 hamster with SCN from wild-type hamster 24 23 22 21 20 19 Before procedures © 2014 Pearson Education, Inc. After surgery and transplant Figure 49.UN05 Brain Nerve net Hydra (cnidarian) © 2014 Pearson Education, Inc. Spinal cord (dorsal nerve cord) Sensory ganglia Salamander (vertebrate) Figure 49.UN06 CNS PNS VENTRICLE Ependymal cell Cilia Astrocyte Oligodendrocyte Schwann cells Capillary © 2014 Pearson Education, Inc. Neuron Microglial cell Figure 49.UN07 Cerebrum Forebrain Thalamus Hypothalamus Pituitary gland Midbrain Pons Hindbrain © 2014 Pearson Education, Inc. Medulla oblongata Cerebellum Spinal cord Figure 49.UN08 © 2014 Pearson Education, Inc.