THE NERVOUS SYSTEM CHAPTER 28 EVOLUTION OF THE ANIMAL NERVOUS SYSTEM • The nervous system consists of neurons and supporting cells. • Association neurons (or interneurons) are located in the brain and spinal cord of vertebrates, together called the central nervous system (CNS). • They help provide more complex reflexes and, in the case of the brain, higher associative functions, including learning and memory. EVOLUTION OF THE ANIMAL NERVOUS SYSTEM • Sensory neurons that carry impulses from sensory receptors to the CNS. • Motor neurons that carry impulses away from the the CNS to effectors (muscles and glands). • Together, the motor and sensory neurons constitute the peripheral nervous system (PNS) of vertebrates. Association neuron Cell body Cell body Axon 2 Dendrites 3 Cell body Axon Motor neuron Axon 1 Sensory neuron Direction of conduction Dendrites ORGANIZATION OF THE VERTEBRATE NERVOUS SYSTEM Central nervous system “Integration center’’ (Association neurons of brain and spinal cord) Motor nervous system Sensory nervous system (Efferent nerves) (Afferent nerves) Muscles Organs Glands Sensory receptors Peripheral nervous system EVOLUTION OF THE ANIMAL NERVOUS SYSTEM • Sponges are the only major phylum of animals that lack nerves. • Sponges respond minimally to stimuli and do not send messages from one part of the body to another. EVOLUTION OF THE ANIMAL NERVOUS SYSTEM • The simplest nervous systems occur among cnidarians. • Cnidarian neurons are linked to one another in a web, or nerve net. • There is no associative activity and little coordination. • Any motion that results is called a reflex because it is an automatic consequence of stimulation. EVOLUTION OF THE ANIMAL NERVOUS SYSTEM • The first associative activity in nervous systems is seen in free living flatworms, phylum Platyhelminthes. • These animals have a ladderlike nervous system with two nerve cords. • The two cords converge at an enlarged association area that functions as a primitive brain. EVOLUTION OF THE ANIMAL NERVOUS SYSTEM • A series of evolutionary changes lead from the flatworms to the vertebrate nervous system: • More sophisticated sensory mechanisms. • Differentiation into central and peripheral nervous systems. • Differentiation of sensory and motor areas. • Increased complexity of association. • Elaboration of the brain. Nerve net 1 Cnidarian Associative neurons Nerve cords 2 Flatworm Central nervous system Brain Peripheral nerves 3 Earthworm Brain Ventral nerve cords Giant axon 5 Arthropod 4 Mollusk NEURONS GENERATE NERVE IMPULSES • All neurons have the same basic structure: cell body, dendrites, and axon. Cell body Nucleus Dendrites Schwann cell Schwann cell Axon Nucleus Axon (a) Myelin sheath Node of Ranvier Schwann cell Axon (b) Myelin sheath NEURONS GENERATE NERVE IMPULSES • Most neurons depend upon support from neuroglial cells. • Schwann cells and oligodendrocytes are important supporting cells that envelop the axons of many neurons with a sheath of fatty material called myelin. • Myelin acts as an electrical insulator. NEURONS GENERATE NERVE IMPULSES • Schwann cells produce myelin in the PNS while oligodendrocytes produce myelin in the CNS. • The myelin is wrapped around the axon as a myelin sheath comprised of multiple layers. • The myelin sheath is interrupted at intervals, leaving unmyelinated gaps called nodes of Ranvier. • Nerve impulses jump from node to node, greatly increasing the speed of transmission. • In multiple sclerosis and Tay-Sachs disease, the myelin sheath degenerates. NEURONS GENERATE NERVE IMPULSES • When a neuron is “at rest,” active transport channels (sodium-potassium pumps) in the neuron transport Na+ out of the cell and K+ ions in. • The result is to make the outside of the membrane more positive than the inside, a condition called the resting membrane potential. NEURONS GENERATE NERVE IMPULSES • Neurons are constantly expending energy to maintain the resting membrane potential. • The voltage difference between the neuron interior and exterior is –70 millivolts. • The resting potential is the starting point for a nerve impulse. NEURONS GENERATE NERVE IMPULSES • A nerve impulse travels along the axon and dendrites as electrical current caused by ions moving in and out of the neuron through voltage-gated channels. • These membrane channels open and close in response to electrical voltage changes. • The impulse starts when pressure or other sensory inputs disturb a neuron’s plasma membrane, causing Na+ channels to open. NEURONS GENERATE NERVE IMPULSES • When Na+ channels open, Na+ floods into the neuron from the outside. • For a brief moment, the inside of the neuron is “depolarized,” becoming more positive. • The open Na+ channels in the small patch of depolarized membrane remain open for only one half of a millisecond. • If the voltage change of the depolarization is great enough, it causes nearby voltage-gated Na+ and K+ channels to open. NEURONS GENERATE NERVE IMPULSES 1 Na+ channel • The Na+ channels open first, which starts a wave of depolarization moving down the neuron. • This moving local reversal of voltage is called an action potential. • An action potential follows an allor-none law: a large enough depolarization produces either a full action potential or none at all. K+ Voltage-gatedK+ channel channels + + + + – – – – Na+ + + + – – – – Na–K pump– – – – – + Na+ + + + + + Na+ At the resting membrane potential, the inside of the axon is negatively charged because the sodium-potassium pump keeps a higher concentration of Na+ outside. Voltage-gated ion channels are closed, but there is some leakage of K+. 2 K+ – – – + + + + + + + + + – – – – – – Na+ + + + – – – – – – – – – + + + + + + Na+ In response to a stimulus, the membrane depolarizes: Voltagegated Na+ channels open, Na+ flows into the cell, and the inside becomes more positive. 3 K+ K+ + + – – – + + + + – – + + + – – – – Na+ Na+ – – + + + – – – – + + –+ – – + + + + Na+ K The local change in voltage opens adjacent voltage-gated Na+ channels, and an action potential is produced. 4 K+ K+ K+ + + + + – – – + + – – – – + + + – – Na+ – – – – + + + – – + + + + – – – + + K+ As the action potential travels farther down the axon, voltage-gated Na+ channels close and K+ channels open, allowing K+ to flow out of the cell and restoring the negative charge inside the cell. Ultimately, the sodium-potassium pump restores the resting membrane potential. NEURONS GENERATE NERVE IMPULSES • The K+ voltage-gated channels open after a slight delay, causing K+ to flow out of the cell • this makes the interior of the neuron more negative, causing the voltage-gated Na+ channels to close. • The period of time after an action potential has passed but before the resting potential is restored is called the refractory period. http://youtu.be/XdCrZm_JAp0 THE SYNAPSE • Axons do not actually make direct contact with other neurons or with other cells. • A narrow gap called the synaptic cleft, separates the axon tip and the target neuron or tissue. • This junction of an axon with another cell is called a synapse. • The membrane on the axon side is called the presynaptic membrane. • The membrane on the receiving side is called the postsynaptic membrane. A SYNAPSE BETWEEN TWO NEURONS Synaptic cleft Axon of presynaptic cell Postsynaptic cell Presynaptic vesicles Presynaptic membrane Postsynaptic membrane © Dennis Kunkel/Phototake THE SYNAPSE • Signals from an axon are carried across the synapse by chemical messengers called neurotransmitters. • These chemicals are packaged in tiny sacs, or vesicles, at the tip of the axon. • When a nerve impulse reaches the end of an axon, it causes the vesicles to release the neurotransmitters into the synaptic cleft. • Chemically-gated channels on the postsynaptic membrane then respond by allowing ions to enter. EVENTS AT THE SYNAPSE Neurotransmitter Gate closed Axon terminal Chemically-gated ion channel Mitochondria Synaptic vesicles Synaptic cleft Postsynaptic cell skeletal muscle) (a) (b) THE SYNAPSE • The vertebrate nervous system uses dozens of different kinds of neurotransmitters. • They fall into two classes, depending on whether they excite or inhibit the postsynaptic cell. • In an excitatory synapse, the chemically-gated channel is usually a Na+ channel. • This can lead to an action potential. • In an inhibitory synapse, the chemically-gated channel is usually a K+ or Cl– channel. • This prevents an action potential. THE SYNAPSE • An individual nerve cell can possess both excitatory and inhibitory synapses. • Integration must occur in which the various excitatory and inhibitory electrical effects tend to cancel or reinforce one another. • If the result of the integration is a large enough depolarization, an action potential will fire. INTEGRATION Axon hillock Axon (right): © E.R. Lewis, YY Zeevi, T.E. Everhart, U. of California/Biological Photo Service A single motor neuron in the spinal cord may have as many as 50,000 synapses! THE SYNAPSE • Examples of neurotransmitters: • Acetylcholine (Ach) • Used at the synapse between a nerve and a muscle fiber, called a neuromuscular junction. • It is used in excitatory synapses in skeletal muscle and in inhibitory synapses in cardiac muscle. • Glycine and GABA • These are inhibitory neurotransmitters, especially in neural controls of body movements. THE SYNAPSE • Biogenic amines are a group of neurotransmitters including: • Dopamine, important in controlling body movements. • Norepinephrine and epinephrine, which are involved in the autonomic nervous system. • Serotonin, which is involved in sleep regulation and other emotional states. ADDICTIVE DRUGS ACT ON CHEMICAL SYNAPSES • Emotional states (mood, pleasure, pain, etc.) are determined by particular groups of neurons that use special sets of Serotonin neurotransmitters and neuromodulators. • Many researchers think that depression results from a Prozac shortage of serotonin. blocks • Prozac, an antidepressant, reabsorption inhibits the reabsorption of serotonin. Receptor ADDICTIVE DRUGS ACT ON CHEMICAL SYNAPSES • Nerve cells are particularly prone to the loss of sensitivity when exposed to a chemical signal for a long time. • If receptor proteins within synapses are exposed to high levels of neurotransmitters for prolonged periods, the nerve cell often responds by inserting fewer receptor proteins into the membrane. ADDICTIVE DRUGS ACT ON CHEMICAL SYNAPSES • When receptor proteins in the pleasure pathways of the brain are exposed to high levels of dopamine due to cocaine, for example, the nerve cells respond by lowering the number of receptor proteins. • With so few receptors, the drug user needs the drug to maintain even normal nerve activity levels. • This is addiction, the physiological adaptation of the nervous system due to drug abuse. KEY BIOLOGICAL PROCESS: DRUG ADDICTION 1 Neurotransmitter 2 3 4 Transporter protein Synapse Receptor protein At a normal synapse, neurotransmitters are quickly recycled by transporter proteins, so the firing rate of receptor proteins stays low. Drug molecule Drug molecules like cocaine bind to the transporters and block recycling, so the level of neurotransmitters rises, and the firing rate increases. The receiving neuron “turns down the volume” by lowering the number of receptors, so the firing rate returns to normal. If the cocaine is removed, the level of neurotransmitters falls to normal, too low to fire the reduced number of receptors. EVOLUTION OF THE VERTEBRATE BRAIN • The brain is the most complex vertebrate organ ever to evolve, and it can perform a bewildering variety of complex functions. EVOLUTION OF THE VERTEBRATE BRAIN • The basic organization of the vertebrate brain can be seen in the brains of primitive fishes. Thalamus Cerebrum Optic tectum Cerebellum Olfactory bulb Spinal cord Optic chiasm Medulla oblongata Hindbrain (Rhombencephalon) Pituitary Hypothalamus Midbrain (Mesencephalon) Forebrain (Prosencephalon) EVOLUTION OF THE VERTEBRATE BRAIN • The brain is divided into three regions that are found in differing proportions in all vertebrates: • 1) The hindbrain, or rhombencephalon - The largest portion of the brain in fishes. • 2) The midbrain, or mesencephalon - Devoted primarily to processing visual information in fishes. • 3) The forebrain, or prosencephalon - Concerned mainly with olfaction (smell) in fishes. • Plays a far more dominant role in neural processing in terrestrial vertebrates than in fishes. THE EVOLUTION OF THE VERTEBRATE BRAIN Shark Frog Crocodile Cat Spinal cord Medulla oblongata Human Cerebellum Optic tectum Midbrain Cerebrum Olfactory tract Bird EVOLUTION OF THE VERTEBRATE BRAIN • In sharks and other fishes, the hindbrain is predominant, and the rest of the brain serves primarily to process sensory information. • In amphibians and reptiles, the forebrain is far larger, and it contains a larger cerebrum devoted to associative activity. • In birds, which evolved from reptiles, the cerebrum is even more pronounced. • In mammals, the cerebrum is the largest portion of the brains. HOW THE BRAIN WORKS • The cerebrum is the center for thought and association and makes up about 85% of the weight of the brain. • The cerebrum is divided into right and left halves, called cerebral hemispheres. • Much of the neural activity of the cerebrum occurs within a thin, gray outer layer called the cerebral cortex. Thalamus Corpus striatum Cerebral cortex Corpus callosum Skull Pineal gland Pons Hypothalamus Pituitary gland Reticular formation Medulla oblongata Cerebellum Spinal cord HOW THE BRAIN WORKS • The cerebral cortex is gray because it is densely packed with cell bodies. • The wrinkles on the surface of the cerebral cortex increase its surface area (and number of cell bodies) threefold. • Underneath the cortex is a solid white region of myelinated nerve fibers that shuttle information between the cortex and the rest of the brain. HOW THE BRAIN WORKS • Different areas of the cerebral cortex control different body activities. Frontal lobe Visual association area Cerebrum Vision Occipital lobe Smell Temporal lobe Medulla oblongata Balance and coordination Primary visual area Cerebellum Spinal cord HOW THE BRAIN WORKS • The right and left cerebral hemispheres are linked by bundles of neurons called tracts. • The tracts serve as information highways telling each half of the brain what the other half is doing. • These tracts cross over so that each half of the brain controls muscles and glands on the opposite side of the body. HOW THE BRAIN WORKS • Beneath the cerebrum are the thalamus and hypothalamus. Hypothalamus • The thalamus is the major site of sensory processing in the brain. • The hypothalamus Amygdala integrates internal activities. Hippocampus • It controls centers in the brain stem that regulate the internal environment of the body, such as heartbeat, temperature, blood pressure, and respiration rate. • It also directs secretions of the pituitary gland. Thalamu HOW THE BRAIN WORKS • Extending back from the base of the brain is a structure known as the cerebellum. • It controls balance, posture, and muscular coordination. • It is best developed in birds to control the complicated balance and coordination associated with flight. HOW THE BRAIN WORKS • The brain stem connects the rest of the brain to the spinal cord and includes: • the midbrain • pons • medulla oblongata • The brain stem contains nerves that control functions, such as breathing, swallowing, digestion, heartbeat, and diameter of blood vessels. HOW THE BRAIN WORKS • The two hemispheres of the cerebrum are responsible for different activities. • Language is lateralized to one dominant hemisphere, usually the left. • The nondominant hemisphere is adept at spatial reasoning. DIFFERENT BRAIN REGIONS CONTROL VARIOUS ACTIVITIES THE SPINAL CORD • The spinal cord is a cable of neurons extending from the brain down through the backbone. • The spinal cord is surrounded and protected by a series of bones called the vertebrae. • Spinal nerves pass out to the body from between the vertebrae. THE VERTEBRATE NERVOUS SYSTEM Cerebrum Grey matter White matter Femoral nerve Sciatic nerve Tibial nerve Sacral nerves Lumbar nerves Thoracic nerves Cerebellum Cervical nerves Thalamusand hypothalamus (surroundedby the cerebrum) Spinal cord VOLUNTARY AND AUTONOMIC NERVOUS SYSTEMS • The motor pathways of the PNS can be further divided into: • The somatic (voluntary) nervous system • Relays commands to the skeletal muscles • The autonomic (involuntary) nervous system • Relays commands to the smooth muscles of the body and to cardiac muscle THE DIVISIONS OF THE VERTEBRATE NERVOUS SYSTEM Nervous system Central nervous system Brain Peripheral nervous system Spinal cord Sensory pathways Somatic (voluntary) nervous system Sympathetic division Motor pathways Autonomic (involuntary) nervous system Parasympathetic division VOLUNTARY AND AUTONOMIC NERVOUS SYSTEMS • Motor neurons of the voluntary system stimulate muscles to contract in two ways. • First, muscles are stimulated to contract in response to conscious commands. • Second, muscles can be stimulated as part of reflexes (does not require conscious control. VOLUNTARY AND AUTONOMIC NERVOUS SYSTEMS SPINAL CORD White Gray matter matter • A reflex produces a rapid motor response to a stimulus because a sensory neuron passes information Monosynaptic synapse directly to a motor neuron. Motor • Useful for the body to neuron react particularly quicklyQuadriceps muscle in time of danger. (effector) • Many reflexes never reach the brain. Dorsalroot ganglion Sensory neuron Stretch receptor Patella Patellar tendon VOLUNTARY AND AUTONOMIC NERVOUS SYSTEMS • Neurons of the autonomic nervous system carry messages from the CNS that keep the body going even when it is not active. • The CNS uses this system to maintain the body’s homeostasis. • It is comprised of two elements that act in opposition to one another: • Sympathetic nervous system that dominates in times of stress. • Parasympathetic nervous system that does the opposite and conserves energy.