Biology 141 – Human Anatomy and Physiology
Text: Marieb and Hoehn, 8th edn – 2010 Instructor: J i H i i Mi h ll I Jamie Heisig-Mitchell Lecture Outline: Chapters 10 - 14

Skeletal Muscles: Functional Groups

Prime movers

Provide the major force for producing a specific movement


Antagonists g

Oppose or reverse a particular movement

Skeletal Muscles: Functional Groups

 

Add force to a movement Reduce undesirable or unnecessary movement

4. 4


Synergists that immobilize a bone or muscle’s origin



Muscle Mechanics: Lever Systems

Components of a lever system
 Lever—rigid

bar (bone) that moves on a fixed point or fulcrum (joint)  Effort—force (supplied by muscle contraction) applied to a lever to move a resistance (load)  Load—resistance (bone + tissues + any added weight) moved by the effort

Effort x length of effort arm = load x length of load arm (force x distance) = (resistance x distance)
Effort 10 kg

0.25 cm Effort 25 cm Fulcrum Load

10 x 25 = 1000 x 0.25 250 = 250

1000 kg Load


(a) Mechanical advantage with a power lever

Figure 10.2a

Effort 100 kg Effort Load 25 cm 50 cm Fulcrum Fulcrum 50 kg Load

100 x 25 = 50 x 50 2500 = 2500

(b) Mechanical disadvantage with a speed lever

Figure 10.2b



Classes of Lever Systems

First class
 Fulcrum

between load and effort

(a) First-class lever Arrangement of the elements is load-fulcrum-effort Load Effort

Fulcrum Load

Fulcrum Example: scissors


Figure 10.3a (1 of 2)

(a) First-class lever Arrangement of the elements is load-fulcrum-effort




In the body: A first-class lever system raises your head off your chest. The posterior neck muscles provide the effort, the atlanto-occipital joint is the fulcrum, and the weight to be lifted is the facial skeleton.
Figure 10.3a (2 of 2)


3b (1 of 2) (b) Second-class lever Arrangement of the elements is fulcrum-load-effort Effort Load Fulcrum In the body: Second-class leverage is exerted when you stand on tip-toe. and the weight of the body is the load. the joints of the ball of the foot are the fulcrum. Figure 10. The effort is exerted by the calf muscles pulling upward on the heel.3b (2 of 2) 4 .3/20/2012 Classes of Lever Systems  Second class  Load between fulcrum and effort (b) Second-class lever Arrangement of the elements is fulcrum-load-effort Load Fulcrum Load Effort Effort Example: wheelbarrow Fulcrum Figure 10.

the fulcrum is the elbow joint.3c (1 of 2) (c) Third-class lever Arrangement of the elements is load-effort-fulcrum Effort Load Fulcrum In the body: Flexing the forearm by the biceps brachii muscle exemplifies third-class leverage.3c (2 of 2) 5 . and the load is the hand and distal end of the forearm. Figure 10.3/20/2012 Classes of Lever Systems  Third class  Effort applied between fulcrum and load (c) Third-class lever Arrangement of the elements is load-effort-fulcrum Load Effort Fulcrum Load Fulcrum Effort Example: tweezers or forceps Figure 10. The effort is exerted on the proximal radius of the forearm.

Integration g  Interpretation of sensory input 3.1 Divisions of the Nervous System  Central nervous system (CNS)  Brain  Integration and spinal cord and command center  Peripheral nervous system (PNS)  Paired spinal and cranial nerves carry messages to and from the CNS 6 . Sensory input  Information gathered by sensory receptors about internal and external changes 2. Motor output  Activation of effector organs (muscles and glands) produces a response Sensory input Integration Motor output Figure 11.3/20/2012 Functions of the Nervous System 1.

etc 7 . urination. cardiac muscle. and joints Visceral afferent fibers—convey impulses from visceral fibers convey organs Transmits impulses from the CNS to effector organs 2. and glands Two functional subdivisions v   Sympathetic – “Fight or Flight” Parasympathetic . Sensory (afferent) division   Somatic afferent fibers—convey impulses from skin. Motor (efferent) division  Motor Division of PNS 1. Autonomic (involuntary) nervous system (ANS)    Visceral motor nerve fibers Regulates smooth muscle.3/20/2012 Peripheral Nervous System (PNS)  Two functional divisions 1. Somatic (voluntary) nervous system  Conscious control of skeletal muscles Motor Division of PNS 2. defecation.digestion. skeletal muscles.

smooth muscles.2 Histology of Nervous Tissue  Two principal cell types 1. and glands Parasympathetic division Conserves energy Promotes housekeeping functions during rest Sympathetic motor fiber of ANS Structure Function Sensory (afferent) division of PNS Motor (efferent) division of PNS Heart Parasympathetic motor fiber of ANS Bladder Figure 11. Neurons—excitable cells that transmit electrical signals Histology of Nervous Tissue 2.3/20/2012 Central nervous system (CNS) Brain and spinal cord Integrative and control centers Peripheral nervous system (PNS) Cranial nerves and spinal nerves Communication lines between the CNS and the rest of the body Sensory (afferent) division Somatic and visceral sensory nerve fibers Conducts impulses from receptors to the CNS Motor (efferent) division Motor nerve fibers Conducts impulses from the CNS to effectors (muscles and glands) Somatic sensory fiber Skin Somatic nervous system Somatic motor (voluntary) Conducts impulses from the CNS to skeletal muscles Visceral sensory fiber Stomach Skeletal muscle Motor fiber of somatic nervous system Sympathetic division Mobilizes body systems during activity Autonomic nervous system (ANS) Visceral motor (involuntary) Conducts impulses from the CNS to cardiac muscles. Neuroglia (glial cells)—supporting cells:       Astrocytes (CNS) Microglia (CNS) Ependymal cells (CNS) Oligodendrocytes (CNS) Satellite cells (PNS) Schwann cells (PNS) 8 .

synaptic endings. and highly branched glial cells Cling to neurons. versatile. and capillaries Support and brace neurons Astrocytes     Help determine capillary permeability Guide migration of young neurons Control the chemical environment Participate in information processing i th b i P ti i t i i f ti i in the brain Capillary Neuron Astrocyte (a) Astrocytes are the most abundant CNS neuroglia. Figure 11.3/20/2012 Astrocytes    Most abundant.3a 9 .

3/20/2012 Microglia    Small.3b Ependymal Cells   Range in shape from squamous to columnar May be ciliated  Line  Separate the central cavities of the brain and spinal column the CNS interstitial fluid from the cerebrospinal fluid in the cavities 10 . ovoid cells with thorny processes Migrate toward injured neurons Phagocytize microorganisms and neuronal debris Neuron Microglial cell (b) Microglial cells are defensive cells in the CNS. Figure 11.

3/20/2012 Fluid-filled cavity Ependymal cells Brain or spinal cord tissue (c) Ependymal cells line cerebrospinal fluid-filled cavities.3c Oligodendrocytes   Branched cells Processes wrap CNS nerve fibers. forming insulating myelin sheaths Myelin sheath Process of oligodendrocyte Nerve N fibers (d) Oligodendrocytes have processes that form myelin sheaths around CNS nerve fibers. Figure 11.3d 11 . Figure 11.

3e Neurons (Nerve Cells)  Special characteristics:  Long-lived ( 100 years or more) few exceptions  High metabolic rate—depends on continuous supply of oxygen and glucose d l  Plasma membrane functions in:  Amitotic—with  Electrical  Cell-to-cell signaling interactions during development 12 .3/20/2012 Satellite Cells and Schwann Cells  Satellite cells  Surround neuron cell bodies in the PNS  Schwann cells (neurolemmocytes)  Surround peripheral nerve fibers and form myelin sheaths  Vital to regeneration of damaged peripheral nerve fibers Satellite cells Cell body of neuron Schwann cells (forming myelin sheath) Nerve fib N fiber (e) Satellite cells and Schwann cells (which form myelin) surround neurons in the PNS. Figure 11.

3/20/2012 Cell Body (Perikaryon or Soma)     Biosynthetic center of a neuron Spherical nucleus with nucleolus Well-developed Golgi apparatus Rough R h ER called Nissl bodies (chromatophilic ll d Ni l b di ( h t hili substance) Cell Body (Perikaryon or Soma)    Network of neurofibrils (neurofilaments) Axon hillock—cone-shaped area from which axon arises Clusters of cell bodies are called nuclei in the CNS CNS.4b Schwann cell Neurilemma (one interTerminal node) branches 13 . ganglia in the PNS Dendrites (receptive regions) Cell body (biosynthetic center and receptive region) Nucleolus Axon (impulse generating and conducting region) Nucleus Nissl bodies Axon hillock (b) Impulse direction Node of Ranvier Axon terminals (secretory region) Figure 11.

and diffusely branched Receptive (input) region of a neuron Convey electrical signals toward the cell body as graded potentials The Axon    One axon per cell arising from the axon hillock Long axons (nerve fibers) Occasional branches (axon collaterals) 14 .3/20/2012 Processes   Dendrites and axons Bundles of processes are called  Tracts  Nerves in the CNS in the PNS Dendrites    Short. tapering.

enzymes  Retrograde—toward  Examples: the cell body organelles to be degraded. signal molecules. viruses.3/20/2012 The Axon   Numerous terminal branches (telodendria) Knoblike axon terminals (synaptic knobs or boutons)  Secretory  Release region of neuron neurotransmitters to excite or inhibit other cells Axons: Function   Conducting region of a neuron Generates and transmits nerve impulses (action potentials) away from the cell body Axons: Function  Molecules and organelles are moved along axons by motor molecules in two directions:  Anterograde—toward  Examples: axonal terminal mitochondria. membrane components. and bacterial toxins 15 .

3/20/2012 Myelin Sheath   Segmented protein-lipoid sheath around most long or large-diameter axons It functions to:  Protect  Increase and electrically insulate the axon speed of nerve impulse transmission Myelin Sheaths in the PNS  Schwann cells wraps many times around the axon  Myelin sheath—concentric layers of Schwann cell membrane  Neurilemma—peripheral bulge of Schwann cell p p g cytoplasm Myelin Sheaths in the PNS  Nodes of Ranvier  Myelin  Sites sheath gaps between adjacent Schwann cells where axon collaterals can emerge 16 .

5a Unmyelinated Axons   Thin nerve fibers are unmyelinated One Schwann cell may incompletely enclose 15 or more unmyelinated axons Myelin Sheaths in the CNS     Formed by processes of oligodendrocytes. The tight membrane wrappings surrounding the axon form the myelin sheath. not the whole cells Nodes of Ranvier are present No neurilemma Thinnest fibers are unmyelinated 17 . wrapping its plasma membrane loosely around it in successive layers. Figure 11. Neurilemma Myelin sheath (a) Myelination of a nerve fiber (axon) 3 The Schwann cell cytoplasm is forced from between the membranes. Schwann cell nucleus 2 The Schwann cell then rotates around the axon.3/20/2012 Schwann cell plasma membrane Schwann cell cytoplasm Axon 1 A Schwann cell envelopes an axon.

Bipolar—1 axon and 1 dendrite  18 .. Figure 11. retinal neurons 2. e.3/20/2012 Myelin sheath Process of oligodendrocyte Nerve N fibers (d) Oligodendrocytes have processes that form myelin sheaths around CNS nerve fibers.g.3d White Matter and Gray Matter  White matter  Dense collections of myelinated fibers neuron cell bodies and unmyelinated fibers  Gray matter  Mostly Structural Classification of Neurons  Three types: 1. Multipolar—1 axon and several dendrites   Most abundant Motor neurons and interneurons Rare.

short process that has two branches:   Peripheral process—more distal branch.1 (2 of 3) 19 .3/20/2012 Structural Classification of Neurons 3. often associated with a sensory receptor Central process branch entering the CNS process—branch Table 11.1 (1 of 3) Table 11. Unipolar (pseudounipolar)—single.

3/20/2012 Functional Classification of Neurons  Three types: 1. Sensory (afferent)  Transmit impulses from sensory receptors toward the CNS Carry impulses from the CNS to effectors 2.1 (3 of 3) 20 . most are entirely within the CNS Table 11. Interneurons (association neurons)  Shuttle signals through CNS pathways. Motor (efferent)  Functional Classification of Neurons 3.

the system has potential energy Definitions    Voltage (V): measure of potential energy generated by separated charge Potential difference: voltage measured between two points p Current (I): the flow of electrical charge (ions) between two points 21 .3/20/2012 Neuron Function    Neurons are highly irritable Respond to adequate stimulus by generating an action potential (nerve impulse) Impulse is always the same regardless of stimulus Principles of Electricity     Opposite charges attract each other Energy is required to separate opposite charges across a membrane Energy is liberated when the charges move toward one another If opposite charges are separated.

Leakage (nongated) channels—always open Role of Membrane Ion Channels 2. Gated channels (three types):    Chemically gated (ligand-gated) channels—open with binding of a specific neurotransmitter Voltage-gated channels—open and close in response to changes in membrane potential Mechanically gated channels—open and close in response to channels open physical deformation of receptors 22 .3/20/2012 Definitions    Resistance (R): hindrance to charge flow (provided by the plasma membrane) Insulator: substance with high electrical resistance Conductor: substance with low electrical resistance Role of Membrane Ion Channels   Proteins serve as membrane ion channels Two main types of ion channels 1.

6 Gated Channels  When gated channels are open:  Ions diffuse quickly across the membrane along their electrochemical gradients  Along chemical concentration gradients from higher concentration to lower concentration  Along electrical gradients toward opposite electrical charge  Ion flow creates an electrical current and voltage changes across the membrane Resting Membrane Potential (Vr)  Potential difference across the membrane of a resting cell  Approximately –70 mV in neurons (cytoplasmic side of membrane is negatively charged relative to outside) in ionic makeup of ICF and ECF permeability of the plasma membrane  Generated by: G db  Differences  Differential 23 . (b) Voltage-gated ion channels open and close in response to changes in membrane voltage.3/20/2012 Receptor Na+ Neurotransmitter chemical attached to receptor Na+ Na+ Na+ Chemical binds Membrane voltage changes h K+ Open Closed Open K+ Closed (a) Chemically (ligand) gated ion channels open when the appropriate neurotransmitter binds to the receptor. Figure 11. allowing (in this case) simultaneous movement of Na+ and K+.

3/20/2012 Resting Membrane Potential  Differences in ionic makeup has lower concentration of Na+ and Cl– than ECF  ICF has higher concentration of K+ and negatively charged proteins (A–) than ECF  ICF Resting Membrane Potential  Differential permeability of membrane to A–  Slightly permeable to Na+ (through leakage channels)  75 times more permeable to K+ (more leakage p ( g channels)  Freely permeable to Cl–  Impermeable Resting Membrane Potential   Negative interior of the cell is due to much greater diffusion of K+ out of the cell than Na+ diffusion into the cell Sodium-potassium pump stabilizes the resting p p p g membrane potential by maintaining the concentration gradients for Na+ and K+ 24 .

8 Membrane Potentials That Act as Signals  Membrane potential changes when:  Concentrations  Permeability of ions across the membrane change of membrane to ions changes  Changes in membrane potential are signals used to receive.. Suppose a cell has only K+ channels. Outside cell The Na+ concentration is higher outside the cell. K+ (140 mM ) Inside cell Na+ (15 mM ) The permeabilities of Na+ and K+ across the membrane are different. Na+ entry through leakage channels reduces the negative membrane potential slightly. let’s add a pump to compensate for leaking ions. integrate and send information Membrane Potentials That Act as Signals  Two types of signals  Graded  Action potentials short-distance signals signals of axons  Incoming potentials  Long-distance 25 . resulting in the resting membrane potential. K+ K+ Na+ Cell interior –70 mV Figure 11.. K+ leakage channels K+ K+ Na+-K+ ATPases (pumps) maintain the concentration gradients of Na+ and K+ across the membrane. Na+-K+ ATPases (pumps) maintain the concentration gradients. K+ (5 mM ) Na+ (140 mM ) The K+ concentration is higher inside the cell. let’s add some Na+ channels to our cell. K+ K+ K+ K+ Cell interior –90 mV Na+ Now.. K+ loss through abundant leakage channels establishes a negative membrane potential.3/20/2012 The concentrations of Na+ and K+ on each side of the membrane are different. K K+ Na+ Cell interior –70 mV Na+-K+ pump K+ K+ Na+ Finally..

9a Changes in Membrane Potential  Hyperpolarization  An increase in membrane potential (away from zero) of the membrane becomes more negative than the resting potential  Reduces the probability of producing a nerve impulse  Inside 26 . Figure 11. the inside becoming less negative (more positive).3/20/2012 Changes in Membrane Potential  Depolarization A reduction in membrane potential (toward zero) of the membrane becomes less negative than the resting potential  Increases the probability of producing a nerve impulse  Inside Depolarizing stimulus Inside positive Inside negative Depolarization Resting potential Time (ms) (a) Depolarization: The membrane potential moves toward 0 mV.

10a 27 .3/20/2012 Hyperpolarizing stimulus Resting potential Hyperpolarization Time (ms) (b) Hyperpolarization: The membrane potential increases. Figure 11. localized changes in membrane potential Depolarizations or hyperpolarizations Graded potential spreads as local currents change the membrane potential of adjacent regions Stimulus Depolarized region Plasma membrane (a) Depolarization: A small patch of the membrane (red area) has become depolarized. Figure 11. the inside becoming more negative.9b Graded Potentials    Short-lived.

postsynaptic potentials    Magnitude varies di l ( d d) with stimulus M i d i directly (graded) i h i l strength Decrease in magnitude with distance as ions flow and diffuse through leakage channels Short-distance signals Memb brane potential (mV) Active area (site of initial depolarization) –70 Resting potential Distance (a few mm) (c) Decay of membrane potential with distance: Because current is lost through the “leaky” plasma membrane. graded potentials are short-distance signals.g. Figure 11..3/20/2012 (b) Spread of depolarization: The local currents (black arrows) that are created depolarize adjacent membrane areas and allow the wave of depolarization to spread. Consequently. Figure 11.10c 28 . the voltage declines with distance from the stimulus (the voltage is decremental).10b Graded Potentials  Occur when a stimulus causes gated ion channels to open  E. generator potentials. receptor potentials.

3/20/2012 Action Potential (AP)     Brief reversal of membrane potential with a total amplitude of ~100 mV Occurs in muscle cells and axons of neurons Does not decrease in magnitude over distance Principal means of long-distance neural communication The big picture 1 Resting state 2 Depolarization 3 Repolarization Membrane pote ential (mV) 3 4 Hyperpolarization 2 Action potential Threshold 1 4 1 Time (ms) Figure 11.11 (1 of 5) Generation of an Action Potential  Resting state  Only  All leakage channels for Na+ and K+ are open gated Na+ and K+ channels are closed 29 .

3/20/2012 Properties of Gated Channels  Properties of gated channels  Each  Na+ channel has two voltage-sensitive gates gates gates Closed at rest. open with depolarization Open at rest. block channel once it is open  Activation  Inactivation  Properties of Gated Channels    Each K+ channel has one voltage-sensitive gate Closed at rest Opens slowly with depolarization Depolarizing Phase    Depolarizing local currents open voltage-gated Na+ channels Na+ influx causes more depolarization At threshold (–55 to –50 mV) positive feedback ( 55 50 leads to opening of all Na+ channels. and a reversal of membrane polarity to +30mV (spike of action potential) 30 .

11 (2 of 5) Membrane po otential (mV) 3 Action potential Na+ permeability K+ permeability 2 1 4 Time (ms) 1 31 . allowing excessive K+ efflux  This causes after-hyperpolarization of the membrane (undershoot)  Some The AP is caused by permeability changes in the plasma membrane Relative membran permeability ne Figure 11.3/20/2012 Repolarizing Phase  Repolarizing phase  Na+ channel slow inactivation gates close permeability to Na+ declines to resting levels  Slow voltage-sensitive K+ gates open  K+ exits the cell and internal negativity is restored  Membrane Hyperpolarization  Hyperpolarization K+ channels remain open.

3/20/2012 Role of the Sodium-Potassium Pump  Repolarization  Restores  Does the resting electrical conditions of the neuron not restore the resting ionic conditions  Ionic redistribution back to resting conditions is restored by the thousands of sodium-potassium pumps Propagation of an Action Potential    Na+ influx causes a patch of the axonal membrane to depolarize Local currents occur Na+ channels toward the point of origin are inactivated and not affected by the local currents Propagation of an Action Potential    Local currents affect adjacent areas in the forward direction Depolarization opens voltage-gated channels and triggers an AP gg Repolarization wave follows the depolarization wave 32 .

Action potential peak is at the recording electrode. Membrane at the recording electrode is still hyperpolarized.12c 33 . Action potential has not yet reached the recording electrode. Resting potential Peak of action potential Hyperpolarization Figure 11. Figure 11.12a Voltage at 2 ms (b) Time = 2 ms.12b Voltage at 4 ms (c) Time = 4 ms. Figure 11.3/20/2012 Voltage at 0 ms Recording electrode (a) Time = 0 ms. Action potential peak is past the recording electrode.

action potentials either happen completely. or not at all Coding for Stimulus Intensity  All action potentials are alike and are independent of stimulus intensity  How does the CNS tell the difference between a weak stimulus and a strong one?   Strong stimuli can generate action potentials more often than weaker stimuli The CNS determines stimulus intensity by the frequency of impulses 34 .strong enough to push the membrane potential toward and beyond threshold p y AP is an all-or-none phenomenon .3/20/2012 Threshold  At threshold:  Membrane  Na+ is depolarized by 15 to 20 mV permeability increases  N influx exceeds K+ efflux Na  The positive feedback cycle begins Threshold    Subthreshold stimulus .weak local depolarization that does not reach threshold Threshold stimulus .

3/20/2012 Action potentials Threshold Stimulus Time (ms) Figure 11.13 Absolute Refractory Period    Time from the opening of the Na+ channels until the resetting of the channels Ensures that each AP is an all-or-none event Enforces one-way transmission of nerve impulses Absolute refractory period Depolarization (Na+ enters) Relative refractory period Repolarization (K+ leaves) After-hyperpolarization Stimulus Time (ms) Figure 11.14 35 .

3/20/2012 Relative Refractory Period  Follows the absolute refractory period Na+ channels have returned to their resting state K+ channels are still open  Repolarization is occurring  Most  Some   Threshold for AP generation is elevated Exceptionally strong stimulus may generate an AP Conduction Velocity   Conduction velocities of neurons vary widely Effect of axon diameter  Larger diameter fibers have less resistance to local current flow and have faster impulse conduction conduction in unmyelinated axons is slower than saltatory conduction in myelinated axons  Effect of myelination  Continuous Conduction Velocity  Effects of myelination  Myelin  Saltatory sheaths insulate and prevent leakage of charge conduction in myelinated axons is about 30 times faster  V lt Voltage-gated t d  APs Na+ channels are l t d at the nodes N h l located t th d appear to jump rapidly from node to node 36 .

Myelin sheath Figure 11. Voltage-gated Stimulus ion channel (b) In an unmyelinated axon. Stimulus Myelin sheath Node of Ranvier 1 mm (c) In a myelinated axon. voltage decays because current leaks across the membrane. as on a dendrite. APs are generated only in the nodes of Ranvier and appear to jump rapidly from node to node. including interferons and Copazone:  Hold symptoms at bay complications p  Reduce disability  Reduce 37 . speech disturbances. loss of muscular control. Conduction is slow because movements of ions and of the gates of channel proteins take time and must occur before voltage regeneration occurs. voltage-gated Na+ and K+ channels regenerate the action potential at each point along the axon. myelin keeps current in axons (voltage doesn’t decay much). weakness. so voltage does not decay.15 Multiple Sclerosis (MS)      An autoimmune disease that mainly affects young adults Symptoms: visual disturbances.3/20/2012 Stimulus Size of voltage (a) In a bare plasma membrane (without voltage-gated channels). and urinary incontinence Myelin sheaths in the CNS become nonfunctional scleroses Shunting d h Sh i and short-circuiting of nerve i i ii f impulses occurs l Impulse conduction slows and eventually ceases Multiple Sclerosis: Treatment  Some immune system–modifying drugs.

myelinated somatic sensory and motor fibers  Group B fibers p  Intermediate diameter. or an effector cell 38 .3/20/2012 Nerve Fiber Classification  Nerve fibers are classified according to:  Diameter  Degree  Speed Sp of myelination of conduction Nerve Fiber Classification  Group A fibers  Large diameter. unmyelinated ANS fibers The Synapse  A junction that mediates information transfer from one neuron:  To  To another neuron. lightly myelinated ANS fibers  Group C fibers  Smallest diameter.

16 39 .3/20/2012 The Synapse   Presynaptic neuron—conducts impulses toward the synapse Postsynaptic neuron—transmits impulses away from the synapse y p Types of Synapses    Axodendritic—between the axon of one neuron and the dendrite of another Axosomatic—between the axon of one neuron and the soma of another Less common types:  Axoaxonic (axon to axon) (dendrite to dendrite)  Dendrosomatic (dendrite to soma)  Dendrodendritic Axodendritic synapses Axosomatic synapses Dendrites Cell body Axoaxonic synapses (a) Axon Axon Axosomatic synapses (b) Cell body (soma) of postsynaptic neuron Figure 11.

which contains synaptic vesicles  Receptor region on the postsynaptic neuron Synaptic Cleft   Fluid-filled space separating the presynaptic and postsynaptic neurons Prevents nerve impulses from directly passing from one neuron to the next 40 . and may be unidirectional or bidirectional  Are important in:  Embryonic  Some nervous tissue brain regions Chemical Synapses   Specialized for the release and reception of neurotransmitters Typically composed of two parts  Axon terminal of the presynaptic neuron.3/20/2012 Electrical Synapses  Less common than chemical synapses  Neurons are electrically coupled (joined by gap junctions)  Communication is very rapid.

3/20/2012 Synaptic Cleft  Transmission across the synaptic cleft:  Is a chemical event (as opposed to an electrical one) release. and binding of neurotransmitters  Ensures unidirectional communication between neurons  Involves Information Transfer    AP arrives at axon terminal of the presynaptic neuron and opens voltage-gated Ca2+ channels Synaptotagmin protein binds Ca2+ and promotes fusion of synaptic vesicles with axon membrane y p Exocytosis of neurotransmitter occurs Information Transfer   Neurotransmitter diffuses and binds to receptors (often chemically gated ion channels) on the postsynaptic neuron Ion channels are opened. g y inhibitory event (graded potential) 41 . causing an excitatory or p . diffusion.


Chemical synapses transmit signals from one neuron to another using neurotransmitters.
Presynaptic neuron Presynaptic neuron

Postsynaptic neuron 1 Action potential arrives at axon terminal. 2 Voltage-gated Ca2+ channels open and Ca2+ enters the axon terminal.

Mitochondrion Ca2+ Ca2+ Ca2+ Ca2+ Synaptic cleft Synaptic vesicles

3 Ca2+ entry causes neurotransmittercontaining synaptic vesicles to release their contents by exocytosis. 4 Neurotransmitter diffuses across the synaptic cleft and binds to specific receptors on the postsynaptic membrane.

Axon terminal

Postsynaptic neuron

Ion movement Graded potential Reuptake

Enzymatic degradation

Diffusion away from synapse

5 Binding of neurotransmitter opens ion channels, resulting in graded potentials. 6 Neurotransmitter effects are terminated by reuptake through transport proteins, enzymatic degradation, or diffusion away from the synapse.

Figure 11.17

Termination of Neurotransmitter Effects

Within a few milliseconds, the neurotransmitter effect is terminated
 Degradation  Reuptake p

by enzymes by astrocytes or axon terminal y y  Diffusion away from the synaptic cleft

Synaptic Delay

Neurotransmitter must be released, diffuse across the synapse, and bind to receptors Synaptic delay—time needed to do this (0.3–5.0 ms) ) Synaptic delay is the rate-limiting step of neural transmission



Postsynaptic Potentials
 

Graded potentials Strength determined by:
 

Amount of neurotransmitter released Time the neurotransmitter is in the area

Types of postsynaptic potentials
1. 2.

EPSP—excitatory postsynaptic potentials IPSP—inhibitory postsynaptic potentials

Table 11.2 (1 of 4)

Table 11.2 (2 of 4)



Table 11.2 (3 of 4)

Table 11.2 (4 of 4)

Excitatory Synapses and EPSPs

Neurotransmitter binds to and opens chemically gated channels that allow simultaneous flow of Na+ and K+ in opposite directions Na+ influx is greater that K+ efflux, causing a net g , g depolarization EPSP helps trigger AP at axon hillock if EPSP is of threshold strength and opens the voltage-gated channels


Neurotransmitter binding opens K+ or Cl– channels. Neurotransmitter binding opens chemically gated ion channels.18a Inhibitory Synapses and IPSPs    Neurotransmitter binds to and opens channels for K+ or Cl– Causes a hyperpolarization (the inner surface of membrane becomes more negative) g ) Reduces the postsynaptic neuron’s ability to produce an action potential Membra ane potential (mV) Threshold An IPSP is a local hyperpolarization of the postsynaptic membrane and drives the neuron away from AP threshold. Stimulus Time (ms) (a) Excitatory postsynaptic potential (EPSP) Figure 11.18b 45 .3/20/2012 Membra ane potential (mV) Threshold An EPSP is a local depolarization of the postsynaptic membrane that brings the neuron closer to AP threshold. allowing the simultaneous passage of Na+ and K+. Stimulus Time (ms) (b) Inhibitory postsynaptic potential (IPSP) Figure 11.

Excitatory synapse 1 (E1) Excitatory synapse 2 (E2) Inhibitory synapse (I1) Figure 11.19a. (a) No summation: 2 stimuli separated in time cause EPSPs that do not add together.3/20/2012 Integration: Summation    A single EPSP cannot induce an action potential EPSPs can summate to reach threshold IPSPs can also summate with EPSPs. canceling each other out Integration: Summation  Temporal summation  One or more presynaptic neurons transmit impulses in rapid-fire order neuron is stimulated by a large number of terminals at the same time  Spatial summation p  Postsynaptic E1 E1 Threshold of axon of postsynaptic neuron Resting potential E1 E1 Time E1 E1 Time (b) Temporal summation: 2 excitatory stimuli close in time cause EPSPs that add together. b 46 .

3/20/2012 E1 E1 E2 I1 E1 + E2 Time (c) Spatial summation: 2 simultaneous stimuli at different locations cause EPSPs that add together. I1 Time E1 + I1 (d) Spatial summation of EPSPs and IPSPs: Changes in membane potential can cancel each other out. Figure 11. d Integration: Synaptic Potentiation    Repeated use increases the efficiency of neurotransmission Ca2+ concentration increases in presynaptic terminal and ostsynaptic neuron Brief high-frequency stimulation partially depolarizes the postsynaptic neuron Chemically gated channels (NMDA receptors) allow Ca2+ entry  Ca2+ activates kinase enzymes that promote more effective responses to subsequent stimuli  Integration: Presynaptic Inhibition   Release of excitatory neurotransmitter by one neuron may be inhibited by the activity of another neuron via an axoaxonic synapse Less neurotransmitter is released and smaller EPSPs are formed 47 .19c.

3/20/2012 Neurotransmitters    Most neurons make two or more neurotransmitters. and epinephrine Serotonin and histamine  Indolamines    Broadly distributed in the brain Play roles in emotional behaviors and the biological clock 48 . which are released at different stimulation frequencies 50 or more neurotransmitters have been identified Classified by chemical structure and by function Chemical Classes of Neurotransmitters  Acetylcholine (Ach)  Released at neuromuscular junctions and some ANS neurons  Synthesized by enzyme choline acetyltransferase  Degraded by the enzyme acetylcholinesterase (AChE) Chemical Classes of Neurotransmitters  Biogenic amines include:  Catecholamines  Dopamine. norepinephrine (NE).

3/20/2012 Chemical Classes of Neurotransmitters  Amino acids include:  GABA—Gamma  Glycine  Aspartate  Glutamate ()-aminobutyric acid Chemical Classes of Neurotransmitters  Peptides (neuropeptides) include:  Substance  P Mediator of pain signals Act as natural opiates. reduce pain perception  Endorphins   Gut-brain  peptides Somatostatin and cholecystokinin Chemical Classes of Neurotransmitters  Purines such as ATP:  Act in both the CNS and PNS fast or slow responses  Induce Ca2+ influx in astrocytes  Provoke pain sensation  Produce 49 .

3/20/2012 Chemical Classes of Neurotransmitters  Gases and lipids  Nitric oxide (NO) on demand the intracellular receptor guanylyl cyclase to cyclic  Synthesized  Activates GMP  Involved in learning and memory  Carbon monoxide (CO) is a regulator of cGMP in the brain Chemical Classes of Neurotransmitters  Gases and lipids  Endocannabinoids  Lipid  Bind soluble. synthesized on demand from membrane lipids with G protein–coupled receptors in the brain  Involved in learning and memory Functional Classification of Neurotransmitters  Neurotransmitter effects may be excitatory (depolarizing) and/or inhibitory (hyperpolarizing) Determined by the receptor type of the postsynaptic neuron GABA and glycine are usually inhibitory  Glutamate is usually excitatory  Acetylcholine  Excitatory at neuromuscular junctions in skeletal muscle  Inhibitory in cardiac muscle   50 .

Channel-linked receptors G protein-linked receptors 51 . 2.3/20/2012 Neurotransmitter Actions  Direct action  Neurotransmitter binds to channel-linked receptor and opens ion channels  Promotes rapid responses  Examples: ACh and amino acids Neurotransmitter Actions  Indirect action  Neurotransmitter binds to a G protein-linked receptor and acts through an intracellular second messenger  Promotes long-lasting effects  Examples: biogenic amines. and dissolved gases Neurotransmitter Receptors  Types 1. neuropeptides.

20a of ligand (ACh in this case).3/20/2012 Channel-Linked (Ionotropic) Receptors      Ligand-gated ion channels Action is immediate and brief Excitatory receptors are channels for small cations Na influx N + i fl contributes most t d t ib t t to depolarization l i ti Inhibitory receptors allow Cl– influx or K+ efflux that causes hyperpolarization Ion flow blocked Ligand Ions flow Closed ion channel Open ion channel (a) Channel-linked receptors open in response to binding Figure 11. and often prolonged and widespread Examples: muscarinic ACh receptors and those that bind biogenic amines and neuropeptides 52 . slow. complex. G Protein-Linked (Metabotropic) Receptors    Transmembrane protein complexes Responses are indirect.

GDP 2 Receptor 3 G protein 4 Adenylate activates G activates cyclase converts protein. cAMP activates enzymes. y . 5c cAMP activates specific genes. cyclic AMP. Adenylate cyclase Closed ion channel Open ion channel Receptor G protein 5a cAMP changes membrane permeability y p g g by opening or closing ion channels. g . e. cyclic GMP.g. adenylate ATP to cAMP cyclase.17b 53 . 5b Nucleus Active enzyme (b) G-protein linked receptors cause formation of an intracellular second messenger (cyclic AMP in this case) that brings about the cell’s response.. (2nd messenger).3/20/2012 G Protein-Linked Receptors: Mechanism    Neurotransmitter binds to G protein–linked receptor G protein is activated Activated G protein controls production of second messengers. diacylglycerol or Ca2+ G Protein-Linked Receptors: Mechanism  Second messengers  Open or close ion channels kinase enzymes  Phosphorylate channel proteins p y p  Activate genes and induce protein synthesis  Activate 1 Neurotransmitter (1st messenger) binds and activates receptor. y . g. Figure 11.

and nerve cell adhesion molecules or N-CAMs) N Neurotropins that attract or repel the growth cone h l h h  Nerve growth factor (NGF). which keeps the neuroblast alive   Astrocytes provide physical support and cholesterol essential for construction of synapses Cell Death  About 2/3 of neurons die before birth  Death results in cells that fail to make functional synaptic contacts  Many cells also die due to apoptosis (programmed cell death) during development 54 . integrin.3/20/2012 Developmental Aspects of Neurons   The nervous system originates from the neural tube and neural crest formed from ectoderm The neural tube becomes the CNS Neuroepithelial cells of the neural tube undergo differentiation to form cells needed for development  Cells (neuroblasts) become amitotic and migrate  Neuroblasts sprout axons to connect with targets and become neurons  Axonal Growth  Growth cone at tip of axon interacts with its environment via: Cell surface adhesion proteins (laminin.

3/20/2012 Central Nervous System (CNS)   CNS consists of the brain and spinal cord Cephalization  Evolutionary development of the rostral (anterior) p portion of the CNS  Increased number of neurons in the head  Highest level is reached in the human brain Embryonic Development   Neural plate forms from ectoderm Neural plate invaginates to form a neural groove and neural folds Embryonic Development   Neural groove fuses dorsally to form the neural tube Neural tube gives rise to the brain and spinal cord 55 .

3/20/2012 Embryonic Development  Anterior end of the neural tube gives rise to three primary brain vesicles  Prosencephalon—forebrain  Mesencephalon—midbrain p  Rhombencephalon—hindbrain Embryonic Development  Primary vesicles give rise to five secondary brain vesicles  Telencephalon and diencephalon arise from the forebrain  Mesencephalon remains undivided  Metencephalon and myelencephalon arise from the hindbrain Embryonic Development   Telencephalon  cerebrum (two hemispheres with cortex. 56 . hypothalamus. white matter. and basal nuclei) Diencephalon  thalamus. epithalamus. and retina p .

2c-e Effect of Space Restriction on Brain Development    Midbrain flexure and cervical flexure cause forebrain to move toward the brain stem Cerebral hemispheres grow posteriorly and laterally y Cerebral hemisphere surfaces crease and fold into convolutions 57 .3/20/2012 Embryonic Development     Mesencephalon  brain stem (midbrain) Metencephalon  brain stem (pons) and cerebellum Myelencephalon  brain stem (medulla oblongata) Central canal of the neural tube enlarges to form fluid-filled ventricles (c) Secondary brain vesicles (d) Adult brain structures (e) Adult neural canal regions Telencephalon Cerebrum: cerebral hemispheres (cortex. retina Brain stem: midbrain Lateral ventricles Diencephalon Third ventricle Mesencephalon Metencephalon Cerebral aqueduct Brain stem: pons Cerebellum Fourth ventricle Myelencephalon Brain stem: medulla oblongata Spinal cord Central canal Figure 12. basal nuclei) Diencephalon (thalamus. epithalamus). white matter. hypothalamus.

p . ) Cerebellum Regions and Organization of the CNS  Spinal cord  Central  External cavity surrounded by a gray matter core white matter composed of myelinated fiber tracts Regions and Organization of the CNS  Brain  Similar  Nuclei pattern with additional areas of gray matter in cerebellum and cerebrum C Cortex of cerebellum and cerebrum 58 . 2. pons. 3.3/20/2012 Regions and Organization of the CNS  Adult brain regions 1. Cerebral hemispheres Diencephalon Brain stem (midbrain. 4. and medulla) ( .

4 Ventricles of the Brain   Connected to one another and to the central canal of the spinal cord Lined by ependymal cells Ventricles of the Brain  Contain cerebrospinal fluid  Two C-shaped lateral ventricles in the cerebral hemispheres  Third ventricle in the diencephalon  Fourth ventricle in the hindbrain. develops from the lumen of the neural tube 59 .3/20/2012 Central cavity Migratory pattern of neurons Cerebrum Cerebellum Region of cerebellum Cortex of gray matter Inner gray matter Outer white matter Gray matter Central cavity Inner gray matter Outer white matter Brain stem Gray matter Central cavity Outer white matter Spinal cord Inner gray matter Figure 12. dorsal to the pons.

shallow grooves (sulci). understanding Each hemisphere connects to contralateral side of the body There is lateralization of cortical function in the hemispheres 60 . memory storage. sensory perception.3/20/2012 Cerebral Hemispheres  Surface markings Ridges (gyri). communication. voluntary motor initiation. and deep grooves (fissures)  Five lobes  Frontal  Parietal  Temporal  Occipital  Insula  Cerebral Hemispheres  Surface markings Central sulcus  Separates the precentral gyrus of the frontal lobe and the postcentral gyrus of the parietal lobe  Longitudinal fissure  Separates the two hemispheres  Transverse cerebral fissure  Separates the cerebrum and the cerebellum  Cerebral Cortex      Thin (2–4 mm) superficial layer of gray matter 40% of the mass of the brain Site of conscious mind: awareness.

skilled.3/20/2012 Functional Areas of the Cerebral Cortex  The three types of functional areas are:  Motor areas—control voluntary movement areas—conscious awareness of sensation  Association areas—integrate diverse information g v  Sensory  Conscious behavior involves the entire cortex Motor Areas     Primary (somatic) motor cortex Premotor cortex Broca’s area Frontal F t l eye fi ld field Primary Motor Cortex     Large pyramidal cells of the precentral gyri Long axons  pyramidal (corticospinal) tracts Allows conscious control of precise. voluntary movements Motor homunculi: upside-down caricatures representing the motor innervation of body regions 61 .

3/20/2012 Posterior Motor map in precentral gyrus Motor Anterior Toes Jaw Tongue Swallowing Primary motor cortex (precentral gyrus) Figure 12. or patterned motor skills Coordinates simultaneous or sequential actions Involved in the planning of movements that depend on sensory feedback Broca’s Area     Anterior to the inferior region of the premotor area Present in one hemisphere (usually the left) A motor speech area that directs muscles of the tongue Is active as one prepares to speak 62 .9 Premotor Cortex     Anterior to the precentral gyrus Controls learned. repetitious.

skeletal muscles. and joints Capable of spatial discrimination: identification of body region being stimulated 63 .3/20/2012 Frontal Eye Field   Anterior to the premotor cortex and superior to Broca’s area Controls voluntary eye movements Sensory Areas     Primary somatosensory cortex Somatosensory association cortex Visual areas Auditory areas     Olfactory cortex Gustatory cortex Visceral sensory area Vestibular cortex Primary Somatosensory Cortex    In the postcentral gyri Receives sensory information from the skin.

9 Somatosensory Association Cortex    Posterior to the primary somatosensory cortex Integrates sensory input from primary somatosensory cortex Determines size texture and relationship of parts size. texture. of objects being felt Visual Areas  Primary visual (striate) cortex  Extreme  Most posterior tip of the occipital lobe of it is buried in the calcarine sulcus  Receives visual information from the retinas v v 64 .3/20/2012 Posterior Sensory Anterior Sensory map in postcentral gyrus Genitals Primary somatosensory cortex (postcentral gyrus) Intraabdominal Figure 12.

and movement)  Complex processing involves entire posterior half of the hemispheres Auditory Areas  Primary auditory cortex  Superior  Interprets margin of the temporal lobes information from inner ear as pitch.. color. along with the olfactory bulbs and tracts  (Remainder of the rhinencephalon in humans is part of the limbic system)  Region of conscious awareness of odors 65 .3/20/2012 Visual Areas  Visual association area  Surrounds  Uses the primary visual cortex past visual experiences to interpret visual stimuli (e.g. and location posterior to the primary auditory cortex memories of sounds and permits perception of  Auditory association area A d  Located  Stores sounds OIfactory Cortex   Medial aspect of temporal lobes (in piriform lobes) Part of the primitive rhinencephalon. loudness. form.

upset stomach or full bladder Vestibular Cortex   Posterior part of the insula and adjacent parietal cortex Responsible for conscious awareness of balance (p (position of the head in space) p ) 66 ..3/20/2012 Gustatory Cortex   In the insula Involved in the perception of taste Visceral Sensory Area   Posterior to gustatory cortex Conscious perception of visceral sensations.g. e.

and personality Contains working memory needed for judgment judgment. recall. persistence. including the premotor cortex Allow us to give meaning to information received received. and conscience Development depends on feedback from social environment 67 .3/20/2012 Multimodal Association Areas    Receive inputs from multiple sensory areas Send outputs to multiple areas. store it as memory. and decide on action to take Multimodal Association Areas  Three parts  Anterior association area (prefrontal cortex) association area  Limbic association area  Posterior Anterior Association Area (Prefrontal Cortex)     Most complicated cortical region Involved with intellect. compare it to previous experience. reasoning. cognition.

parietal. and occipital lobes Plays a role in recognizing patterns and faces and localizing us in space g p Involved in understanding written and spoken language (Wernicke’s area) Limbic Association Area   Part of the limbic system Provides emotional impact that helps establish memories Lateralization of Cortical Function  Lateralization  Division of labor between hemispheres  Cerebral dominance  Designates the hemisphere dominant for language (left hemisphere in 90% of people) 68 .3/20/2012 Posterior Association Area    Large region in temporal.

intuition.3/20/2012 Lateralization of Cortical Function  Left hemisphere  Controls language. math. visual-spatial skills intuition and artistic skills skills.  Left and right hemispheres communicate via fiber tracts in the cerebral white matter Cerebral White Matter   Myelinated fibers and their tracts Responsible for communication  Commissures (in corpus callosum)—connect gray matter of the two hemispheres p  Association fibers—connect different parts of the same hemisphere  Projection fibers—(corona radiata) connect the hemispheres with lower brain or spinal cord Basal Nuclei (Ganglia)   Subcortical nuclei Consists of the corpus striatum  Caudate  Lentiform nucleus nucleus (putamen + globus pallidus)  Functionally associated with the subthalamic nuclei (diencephalon) and the substantia nigra (midbrain) 69 . and logic  Right hemisphere  Insight Insight.

3/20/2012 Functions of Basal Nuclei  Though somewhat elusive. named for their location Nuclei project and receive fibers from the cerebral cortex 70 . the following are thought to be functions of basal nuclei  Influence  Help p muscular control regulate attention and cognition g g  Regulate intensity of slow or stereotyped movements  Inhibit antagonistic and unnecessary movements Diencephalon  Three paired structures  Thalamus  Hypothalamus  Epithalamus p  Encloses the third ventricle Thalamus      80% of diencephalon Superolateral walls of the third ventricle Connected by the interthalamic adhesion (intermediate mass) Contains several nuclei.

and relays information Afferent impulses from all senses and all parts of the body Impulses from the hypothalamus for regulation of emotion and visceral function  Impulses from the cerebellum and basal nuclei to help direct the motor cortices    Mediates sensation. rate and force of heartbeat. and memory Hypothalamus   Forms the inferolateral walls of the third ventricle Contains many nuclei  Example:  Paired mammillary bodies anterior nuclei  Olfactory relay stations  Infundibulum—stalk that connects to the pituitary gland Hypothalamic Function   Autonomic control center for many visceral functions (e. edits. learning. digestive tract motility) Center for emotional response: Involved in p perception of pleasure. cortical arousal.g. fear.3/20/2012 Thalamic Function   Gateway to the cerebral cortex Sorts. and rage and in biological rhythms and drives 71 . motor activities. blood pressure..

water balance.3/20/2012 Hypothalamic Function     Regulates body temperature. food intake. forms roof of the third ventricle Pineal gland—extends from the posterior border and secretes melatonin  Melatonin—helps regulate sleep-wake cycles Brain Stem  Three regions  Midbrain  Pons  Medulla M oblongata g 72 . and thirst Regulates sleep and the sleep cycle Controls release of hormones by the anterior pituitary Produces posterior pituitary hormones Epithalamus   Most dorsal portion of the diencephalon.

3/20/2012 Brain Stem     Similar structure to spinal cord but contains embedded nuclei Controls automatic behaviors necessary for survival Contains fiber tracts connecting higher and lower neural centers Associated with 10 of the 12 pairs of cranial nerves Midbrain   Located between the diencephalon and the pons Cerebral peduncles  Contain pyramidal motor tracts between third and fourth ventricles  Cerebral aqueduct  Channel Midbrain Nuclei   Nuclei that control cranial nerves III (oculomotor) and IV (trochlear) Corpora quadrigemina—domelike dorsal protrusions   Superior colliculi—visual reflex centers Inferior colliculi—auditory relay centers   Substantia nigra—functionally linked to basal nuclei Red nucleus—relay nuclei for some descending motor pathways and part of reticular formation 73 .

and VII (facial) Some nuclei of the reticular formation Nuclei that help maintain normal rhythm of breathing Fourth ventricle Superior cerebellar peduncle Trigeminal main sensory nucleus Trigeminal motor nucleus Middle cerebellar peduncle Trigeminal nerve (V) Medial lemniscus (b) Pons Reticular formation Pontine nuclei Fibers of pyramidal tract Figure 12. VI (abducens).3/20/2012 Tectum Periaqueductal gray matter Oculomotor nucleus (III) Medial lemniscus Red nucleus Substantia nigra Fibers of pyramidal tract (a) Midbrain Dorsal Superior colliculus Cerebral aqueduct Reticular formation Ventral Crus cerebri of cerebral peduncle Figure 12.16b 74 .16a Pons   Forms part of the anterior wall of the fourth ventricle Fibers of the pons   Connect higher brain centers and the spinal cord Relay impulses between the motor cortex and the cerebellum    Origin of cranial nerves V (trigeminal).

.3/20/2012 Medulla Oblongata      Joins spinal cord at foramen magnum Forms part of the ventral wall of the fourth ventricle Contains a choroid plexus of the fourth ventricle Pyramids—two ventral longitudinal ridges formed y g g by pyramidal tracts Decussation of the pyramids—crossover of the corticospinal tracts Medulla Oblongata     Inferior olivary nuclei—relay sensory information from muscles and joints to cerebellum Cranial nerves VIII. and XII are associated with the medulla Vestibular nuclear complex—mediates responses that maintain equilibrium Several nuclei (e.g. X. nucleus cuneatus and nucleus gracilis) relay sensory information Medulla Oblongata   Autonomic reflex centers Cardiovascular center  Cardiac center adjusts force and rate of heart contraction  Vasomotor center adjusts blood vessel diameter for blood pressure regulation 75 .

with pontine centers Medulla Oblongata  Additional centers regulate  Vomiting  Hiccuping  Swallowing S g  Coughing  Sneezing Hypoglossal nucleus (XII) Dorsal motor nucleus of vagus (X) Inferior cerebellar peduncle Reticular formation Fourth ventricle Choroid plexus Solitary nucleus Vestibular nuclear complex (VIII) Cochlear nuclei (VIII) Nucleus ambiguus Inferior olivary nucleus Pyramid Figure 12.3/20/2012 Medulla Oblongata  Respiratory centers  Generate  Control respiratory rhythm rate and depth of breathing.16c Lateral nuclear group Medial nuclear group Raphe nucleus Medial lemniscus (c) Medulla oblongata 76 .

posterior.17b 77 . and flocculonodular   Folia—transversely oriented gyri Arbor vitae—distinctive treelike pattern of the cerebellar white matter Anterior lobe Cerebellar cortex Arbor vitae Cerebellar peduncles • Superior • Middle • Inferior Medulla oblongata (b) Flocculonodular lobe Posterior lobe Choroid plexus of fourth ventricle Figure 12.3/20/2012 The Cerebellum    11% of brain mass Dorsal to the pons and medulla Subconsciously provides precise timing and appropriate patterns of skeletal muscle contraction Anatomy of the Cerebellum   Two hemispheres connected by vermis Each hemisphere has three lobes  Anterior.

17d Cerebellar Peduncles   All fibers in the cerebellum are ipsilateral Three paired fiber tracts connect the cerebellum to the brain stem  Superior peduncles connect the cerebellum to the midbrain db  Middle peduncles connect the pons to the cerebellum  Inferior peduncles connect the medulla to the cerebellum Cerebellar Processing for Motor Activity     Cerebellum receives impulses from the cerebral cortex of the intent to initiate voluntary muscle contraction Signals from proprioceptors and visual and equilibrium pathways continuously “inform” the cerebellum of the body’s b d ’ position and momentum d Cerebellar cortex calculates the best way to smoothly coordinate a muscle contraction A “blueprint” of coordinated movement is sent to the cerebral motor cortex and to brain stem nuclei 78 .3/20/2012 Anterior lobe Posterior lobe (d) Vermis Figure 12.


Cognitive Function of the Cerebellum
 

Recognizes and predicts sequences of events during complex movements Plays a role in nonmotor functions such as word association and puzzle solving p g

Functional Brain Systems

Networks of neurons that work together and span wide areas of the brain
 Limbic  Reticular

system formation

Limbic System

Structures on the medial aspects of cerebral hemispheres and diencephalon Includes parts of the diencephalon and some cerebral structures that encircle the brain stem



Septum pellucidum Diencephalic structures of the limbic system •Anterior thalamic nuclei (flanking 3rd ventricle) •Hypothalamus •Mammillary body Corpus callosum

Fiber tracts connecting limbic system structures •Fornix •Anterior commissure Cerebral structures of the y limbic system •Cingulate gyrus •Septal nuclei •Amygdala •Hippocampus •Dentate gyrus •Parahippocampal gyrus

Olfactory bulb
Figure 12.18

Limbic System

Emotional or affective brain
 Amygdala—recognizes

angry or fearful facial expressions, assesses danger, and elicits the fear response  Cingulate gyrus plays a role in expressing emotions gyrus—plays via gestures, and resolves mental conflict

Puts emotional responses to odors
 Example:

skunks smell bad

Limbic System: Emotion and Cognition

The limbic system interacts with the prefrontal lobes, therefore:
 We

can react emotionally to things we consciously understand to be happening  We are consciously aware of emotional richness in our lives

Hippocampus and amygdala—play a role in memory



Reticular Formation

Three broad columns along the length of the brain stem
 Raphe

nuclei ( g (large cell) group of nuclei )g p  Lateral (small cell) group of nuclei
 Medial

Has far-flung axonal connections with hypothalamus, thalamus, cerebral cortex, cerebellum, and spinal cord

Reticular Formation: RAS and Motor Function

RAS (reticular activating system)
 Sends

impulses to the cerebral cortex to keep it conscious and alert  Filters out repetitive and weak stimuli (~99% of all stimuli!)  Severe injury results in permanent unconsciousness (coma)

Reticular Formation: RAS and Motor Function

Motor function
 Helps  Reticular

control coarse limb movements autonomic centers regulate visceral motor functions
 Vasomotor  Cardiac  Respiratory




Radiations to cerebral cortex

Visual impulses Reticular formation Ascending general sensory tracts (touch, pain, temperature)

Auditory impulses Descending motor projections to spinal cord
Figure 12.19

Electroencephalogram (EEG)

Records electrical activity that accompanies brain function Measures electrical potential differences between various cortical areas

Brain Waves
   

Patterns of neuronal electrical activity Generated by synaptic activity in the cortex Each person’s brain waves are unique Can be C b grouped into four classes based on di t f l b d frequency measured as Hertz (Hz)



Types of Brain Waves

Alpha waves (8–13 Hz)—regular and rhythmic, lowamplitude, synchronous waves indicating an “idling” brain Beta waves (14–30 Hz)—rhythmic, less regular waves occurring when mentally alert Theta waves (4–7 Hz)—more irregular; common in children and uncommon in adults Delta waves (4 Hz or less)—high-amplitude waves seen in deep sleep and when reticular activating system is damped, or during anesthesia; may indicate brain damage

1-second interval

Alpha waves—awake but relaxed

Beta waves—awake, alert

Theta waves—common in children

Delta waves—deep sleep (b) Brain waves shown in EEGs fall into four general classes.

Figure 12.20b

Brain Waves: State of the Brain

Change with age, sensory stimuli, brain disease, and the chemical state of the body EEGs used to diagnose and localize brain lesions, tumors, infarcts, infections, abscesses, and epileptic , , , , p p lesions A flat EEG (no electrical activity) is clinical evidence of death


and severe biting of the tongue  Tonic-clonic (grand mal) seizures (g )  Victim Control of Epilepsy   Anticonvulsive drugs Vagus nerve stimulators implanted under the skin of the chest can keep electrical activity of the brain from becoming chaotic g 84 .3/20/2012 Epilepsy    A victim of epilepsy may lose consciousness. or petit mal  Mild seizures seen in young children where the expression goes blank loses consciousness. bones are often broken due to intense contractions. fall stiffly. may experience loss of bowel and bladder control. and have uncontrollable jerking Epilepsy is not associated with intellectual impairments p Epilepsy occurs in 1% of the population Epileptic Seizures  Absence seizures.

judgment. fainting or syncopy) is a signal that brain function is impaired Consciousness  Clinically defined on a continuum that grades behavior in response to stimuli  Alertness  Drowsiness  Stupor  Coma ( (lethargy) gy) Sleep   State of partial unconsciousness from which a person can be aroused by stimulation Two major types of sleep (defined by EEG patterns)  Nonrapid  Rapid eye movement (NREM) eye movement (REM) 85 .g. logic.) (memory. etc ) Loss of consciousness (e.3/20/2012 Consciousness     Conscious perception of sensation Voluntary initiation and control of movement Capabilities associated with higher mental processing (memory logic judgment etc..

and sleepwalking may occur. most dreaming occurs. and then REM sleep begins abruptly p g p y Awake REM: Skeletal muscles (except ocular muscles and diaphragm) are actively inhibited. NREM stage 2: Irregular EEG with sleep spindles (short high. Figure 12. arousal is difficult. NREM stage 1: Relaxation begins. g p The suprachiasmatic and preoptic nuclei of the hypothalamus time the sleep cycle A typical sleep pattern alternates between REM and NREM sleep 86 .amplitude bursts).21a (a) Typical EEG patterns Sleep Patterns     Alternating cycles of sleep and wakefulness reflect a natural circadian (24-hour) rhythm RAS activity is inhibited during.3/20/2012 Sleep   First two stages of NREM occur during the first 30– 45 minutes of sleep Fourth stage is achieved in about 90 minutes. night terrors. but RAS also mediates. dreaming sleep . arousal is easy. EEG shows alpha waves. NREM stage 3: Sleep deepens. vital signs decline. NREM stage 4: EEG is dominated by delta waves. theta and delta waves appear. arousal is more difficult. bed-wetting.

21b Importance of Sleep      Slow-wave sleep (NREM stages 3 and 4) is presumed to be the restorative stage People deprived of REM sleep become moody and depressed REM sleep may be a reverse learning process where superfluous information is purged from the brain Daily sleep requirements decline with age Stage 4 sleep declines steadily and may disappear after age 60 Sleep Disorders  Narcolepsy  Lapsing abruptly into sleep from the awake state  Insomnia  Chronic inability to obtain the amount or quality of sleep needed cessation of breathing during sleep  Sleep apnea  Temporary 87 .3/20/2012 Awake REM Stage 1 Stage 2 Non Stage 3 REM Stage 4 Time (hrs) (b) Typical progression of an adult through one night’s sleep stages Figure 12.

limited to seven or p y g . eight pieces of information  Long-term memory (LTM) has limitless capacity Outside stimuli General and special sensory receptors Afferent inputs Temporary storage (buffer) in cerebral cortex Automatic memory Data selected for transfer Data permanently lost Forget F t Short-term memory (STM) Forget Data transfer influenced by: Retrieval Excitement Rehearsal Association of old and new data Long-term memory (LTM) Figure 12.22 Data unretrievable 88 . or working memory)— temporary holding of information.3/20/2012 Language  Language implementation system  Basal nuclei area and Wernicke’s area (in the association cortex on the left side) A l Analyzes i incoming word sounds i d d  Produces outgoing word sounds and grammatical structures  Broca’s  Corresponding areas on the right side are involved with nonverbal language components Memory   Storage and retrieval of information Two stages of storage  Short-term memory (STM.

motor memory. motivated. Includes procedural (skills) memory. surprised. and emotional memory 89 . Declarative memory (factual knowledge)    Explicit information Related to our conscious thoughts and our language ability Stored in LTM with context in which it was learned Categories of Memory 2. and aroused  Rehearsal—repetition and practice  Association—tying new information with old memories  Automatic memory—subconscious information stored in LTM Categories of Memory 1. hard to unlearn y g. Nondeclarative memory     Less conscious or unconscious Acquired through experience and repetition Best remembered by doing.3/20/2012 Transfer from STM to LTM  Factors that affect transfer from STM to LTM  Emotional state—best if alert.

) ACh Basal forebrain Prefrontal cortex ACh Figure 12.3/20/2012 Brain Structures Involved in Declarative Memory   Hippocampus and surrounding temporal lobes function in consolidation and access to memory ACh from basal forebrain is necessary for memory formation and retrieval Thalamus Touch Hearing Vision Taste Smell Basal forebrain Prefrontal cortex Hippocampus Sensory input Thalamus (a) Declarative memory circuits Association cortex Medial temporal lobe (hippocampus. etc.23a Brain Structures Involved in Nondeclarative Memory  Procedural memory  Basal nuclei relay sensory and motor inputs to the thalamus and premotor cortex  Dopamine from substantia nigra is necessary   Motor memory—cerebellum Emotional memory—amygdala 90 .

opening calcium channels in postsynaptic p .3/20/2012 Sensory and motor inputs Association cortex Basal nuclei Thalamus Premotor cortex Dopamine Substantia nigra Premotor cortex Thalamus Basal nuclei Substantia nigra (b) Procedural (skills) memory circuits Figure 12. or LTP) is crucial Neurotransmitter (glutamate) binds to NMDA receptors. p g p y p terminal 91 .23b Molecular Basis of Memory  During learning: Altered mRNA is synthesized and moved to axons and dendrites  Dendritic spines change shape  Extracellular proteins are deposited at synapses involved in LTM  Number and size of presynaptic terminals may increase  More neurotransmitter is released by presynaptic neurons  Molecular Basis of Memory   Increase in synaptic strength (long-term potentiation.

3/20/2012 Molecular Basis of Memory   Calcium influx triggers enzymes that modify proteins of the postsynaptic terminal and presynaptic terminal (via release of retrograde messengers) Enzymes trigger postsynaptic gene activation for synthesis of synaptic proteins. in presence of CREB (cAMP response-element binding protein) and BDNF (brain-derived neurotrophic factor) Protection of the Brain     Bone (skull) Membranes (meninges) Watery cushion (cerebrospinal fluid) Blood-brain b i Bl d b i barrier Meninges     Cover and protect the CNS Protect blood vessels and enclose venous sinuses Contain cerebrospinal fluid (CSF) Form partitions i th skull F titi in the k ll 92 .

3/20/2012 Meninges  Three layers  Dura mater mater  Pia mater  Arachnoid Superior sagittal sinus itt l i Subdural space Subarachnoid space Skin of scalp Periosteum Bone of skull Periosteal Dura Meningeal mater Arachnoid mater Pia mater Arachnoid villus Blood vessel Falx cerebri (in longitudinal fissure only) Figure 12.24 Dura Mater   Strongest meninx Two layers of fibrous connective tissue (around the brain) separate to form dural sinuses 93 .

3/20/2012 Dura Mater  Dural septa limit excessive movement of the brain  Falx cerebri—in the longitudinal fissure.25a Arachnoid Mater     Middle layer with weblike extensions Separated from the dura mater by the subdural space Subarachnoid space contains CSF and blood vessels Arachnoid villi protrude into the superior sagittal sinus and permit CSF reabsorption 94 . attached to crista galli  Falx cerebelli—along the vermis of the cerebellum  Tentorium cerebelli—horizontal dural fold over cerebellum and in the transverse fissure Superior sagittal sinus Straight sinus Crista galli of the ethmoid bone Pituitary gland Falx cerebri Tentorium cerebelli Falx cerebelli (a) Dural septa Figure 12.

3/20/2012 Pia Mater  Layer of delicate vascularized connective tissue that clings tightly to the brain Cerebrospinal Fluid (CSF)  Composition  Watery  Less solution protein and different ion concentrations than plasma  Constant volume Cerebrospinal Fluid (CSF)  Functions  Gives buoyancy to the CNS organs the CNS from blows and other trauma N Nourishes the brain and carries chemical signals g  Protects 95 .

Figure 12. 2 CSF flows through the ventricles and into the subarachnoid space via the median and lateral apertures. oxygen.26a Third ventricle Cerebral aqueduct Lateral aperture Fourth ventricle Median aperture Central canal of spinal cord (a) CSF circulation 2 96 .) (b) CSF formation by choroid plexuses Cavity of ventricle Figure 12. Cl–. 4 CSF is absorbed into the dural venous sinuses via the arachnoid villi. 3 CSF flows through the subarachnoid space.26b Superior sagittal sinus Choroid plexus Interventricular foramen 4 Arachnoid villus Subarachnoid space Arachnoid mater Meningeal dura mater Periosteal dura mater 1 3 Right lateral ventricle (deep to cut) Choroid plexus of fourth ventricle 1 CSF is produced by the choroid plexus of each ventricle. Mg2+.3/20/2012 Choroid Plexuses     Produce CSF at a constant rate Hang from the roof of each ventricle Clusters of capillaries enclosed by pia mater and a layer of ependymal cells Ependymal cells use ion pumps to control the composition of the CSF and help cleanse CSF by removing wastes Ependymal cells Capillary Connective tissue of pia mater Section of choroid plexus Wastes and unnecessary solutes absorbed CSF forms as a filtrate containing glucose. Some CSF flows through the central canal of the spinal cord. etc. vitamins. and ions (Na+.

3a 97 .3/20/2012 Blood-Brain Barrier   Helps maintain a stable environment for the brain Separates neurons from some bloodborne substances Blood-Brain Barrier  Composition  Continuous  Basal endothelium of capillary walls lamina  Feet of astrocytes y  Provide signal to endothelium for the formation of tight junctions Capillary Neuron Astrocyte (a) Astrocytes are the most abundant CNS neuroglia. Figure 11.

e. where it is necessary to monitor the chemical composition of the blood Homeostatic Imbalances of the Brain  Traumatic brain injuries  Concussion—temporary  Contusion—permanent alteration in function damage S Subdural or subarachnoid hemorrhage—may force g y brain stem through the foramen magnum.g. and anesthetics  Absent in some areas. resulting in death  Cerebral edema—swelling of the brain associated with traumatic head injury Homeostatic Imbalances of the Brain  Cerebrovascular accidents (CVAs)(strokes) Blood circulation is blocked and brain tissue dies.g. or sensory and speed deficits  Transient ischemic attacks (TIAs)—temporary episodes of reversible cerebral ischemia  Tissue plasminogen activator (TPA) is the only approved treatment for stroke  98 . vomiting center and the hypothalamus. including alcohol.. blockage of a cerebral artery by a blood clot  Typically leads to hemiplegia. e.. nicotine.3/20/2012 Blood-Brain Barrier: Functions  Selective barrier  Allows  Allows nutrients to move by facilitated diffusion any fat-soluble substances to pass.

there are two clusters of neuroblasts  Alar plate—will become interneurons. axons form white matter of cord  Basal plate—will become motor neurons.28 99 . axons will grow to effectors  Neural crest cells form the dorsal root ganglia sensory neurons. axons grow into the dorsal aspect of the cord Dorsal root ganglion: sensory neurons from neural crest Alar plate: interneurons White matter Basal plate: motor neurons Neural tube cells Central cavity Figure 12.3/20/2012 Homeostatic Imbalances of the Brain  Degenerative brain disorders Alzheimer’s disease (AD): a progressive degenerative disease of the brain that results in dementia  Parkinson’s disease: degeneration of the dopaminereleasing neurons of th substantia nigra l i f the b t ti i  Huntington’s disease: a fatal hereditary disorder caused by accumulation of the protein huntingtin that leads to degeneration of the basal nuclei and cerebral cortex  The Spinal Cord: Embryonic Development  By week 6.

meninges.3/20/2012 Spinal Cord  Location  Begins  Ends at the foramen magnum as conus medullaris at L1 vertebra two-way communication to and from the brain spinal reflex centers  Functions  Provides  Contains Spinal Cord: Protection    Bone. anchors the spinal cord to the coccyx . p y 100 . and CSF Cushion of fat and a network of veins in the epidural space between the vertebrae and spinal dura mater CSF in subarachnoid space Spinal Cord: Protection   Denticulate ligaments: extensions of pia mater that secure cord to dura mater Filum terminale: fibrous extension from conus medullaris.

with the bony vertebral arches removed. The dura mater and arachnoid mater are cut open and reflected laterally. encloses central canal 101 .3/20/2012 Cervical enlargement Dura and arachnoid mater Lumbar enlargement Conus medullaris Cauda equina Filum terminale Cervical spinal nerves Thoracic spinal nerves Lumbar spinal nerves Sacral spinal nerves (a) The spinal cord and its nerve roots. Figure 12.29a Spinal Cord  Spinal nerves  31 pairs  Cervical and lumbar enlargements  The nerves serving the upper and lower limbs emerge here collection of nerve roots at the inferior end of the vertebral canal  Cauda equina  The Cross-Sectional Anatomy  Two lengthwise grooves divide cord into right and left halves  Ventral  Dorsal (anterior) median fissure (p (posterior) median sulcus )  Gray commissure—connects masses of gray matter.

3/20/2012 Epidural space (contains fat) Subdural space Subarachnoid space (contains CSF) Pia mater Arachnoid mater Dura mater Spinal meninges Bone of vertebra Dorsal root ganglion Body of vertebra (a) Cross section of spinal cord and vertebra Figure 12.31a Dorsal median sulcus Dorsal funiculus White Ventral funiculus columns Lateral funiculus Dorsal root ganglion Spinal nerve Dorsal root (fans out into dorsal rootlets) Ventral root (derived from several ventral rootlets) Gray commissure Dorsal horn Gray Ventral horn matter Lateral horn Central canal Ventral median fissure Pia mater Arachnoid mater Spinal dura mater (b) The spinal cord and its meningeal coverings Figure 12.31b Gray Matter     Dorsal horns—interneurons that receive somatic and visceral sensory input Ventral horns—somatic motor neurons whose axons exit the cord via ventral roots Lateral horns (only in thoracic and lumbar regions) – sympathetic neurons Dorsal root (spinal) gangia—contain cell bodies of sensory neurons 102 .

3/20/2012 Dorsal root (sensory) Dorsal root ganglion Somatic sensory neuron Visceral sensory neuron Visceral motor neuron Somatic motor neuron Spinal nerve Ventral root (motor) Ventral horn (motor neurons) Dorsal horn (interneurons) Interneurons receiving input from somatic sensory neurons Interneurons receiving input from visceral sensory neurons Visceral motor (autonomic) neurons Somatic motor neurons Figure 12.32 White Matter     Consists mostly of ascending (sensory) and descending (motor) tracts Transverse tracts (commissural fibers) cross from one side to the other Tracts are located in three white columns (funiculi on each side—dorsal (posterior). and ventral (anterior) Each spinal tract is composed of axons with similar functions Pathway Generalizations     Pathways decussate (cross over) Most consist of two or three neurons (a relay) Most exhibit somatotopy (precise spatial relationships) Pathways are paired symmetrically (one on each side of the spinal cord or brain) 103 . lateral.

3/20/2012 Ascending tracts Fasciculus gracilis Dorsal white Fasciculus cuneatus column Dorsal spinocerebellar tract Ventral spinocerebellar tract Lateral spinothalamic tract Ventral spinothalamic tract Descending tracts Ventral white commissure Lateral reticulospinal tract Lateral corticospinal tract Rubrospinal tract Medial reticulospinal tract Ventral corticospinal tract Vestibulospinal tract Tectospinal tract Figure 12.33 Ascending Pathways   Consist of three neurons First-order neuron  Conducts impulses from cutaneous receptors and p p proprioceptors p  Branches diffusely as it enters the spinal cord or medulla  Synapses with second-order neuron Ascending Pathways  Second-order neuron  Interneuron  Cell body in dorsal horn of spinal cord or medullary nuclei  Axons extend to thalamus or cerebellum 104 .

3/20/2012 Ascending Pathways  Third-order neuron  Interneuron  Cell  Axon body in thalamus extends to somatosensory cortex y Ascending Pathways  Two pathways transmit somatosensory information to the sensory cortex via the thalamus  Dorsal  Spinothalamic p column-medial lemniscal pathways pathways p y  Spinocerebellar tracts terminate in the cerebellum Dorsal Column-Medial Lemniscal Pathways   Transmit input to the somatosensory cortex for discriminative touch and vibrations Composed of the paired fasciculus cuneatus and fasciculus gracilis in the spinal cord and the medial g p lemniscus in the brain (medulla to thalamus) 105 .

34b 106 .34a Axon of first-order neuron Muscle spindle (proprioceptor) (a) Spinocerebellar pathway Anterolateral Pathways   Lateral and ventral spinothalamic tracts Transmit pain. temperature. and coarse touch impulses within the lateral spinothalamic tract Primary somatosensory cortex Axons of third-order neurons Thalamus Cerebrum Midbrain Cerebellum Pons Lateral spinothalamic t L t l i th l i tract t (axons of second-order neurons) Medulla oblongata Pain receptors Cervical spinal cord Lumbar spinal cord (b) Axons of first-order neurons Temperature receptors Spinothalamic pathway Figure 12.3/20/2012 Primary somatosensory cortex Axons of third-order neurons Thalamus Cerebrum Midbrain Cerebellum Dorsal spinocerebellar tract (axons of second-order second order neurons) Pons Medial l M di l lemniscus (t i (tract) t) (axons of second-order neurons) Nucleus gracilis Nucleus cuneatus Medulla oblongata Fasciculus cuneatus (axon of first-order sensory neuron) Joint stretch receptor (proprioceptor) Cervical spinal cord Fasciculus gracilis (axon of first-order sensory neuron) Lumbar spinal cord Touch receptor Dorsal column–medial lemniscal pathway Figure 12.

3/20/2012 Spinocerebellar Tracts   Ventral and dorsal tracts Convey information about muscle or tendon stretch to the cerebellum Descending Pathways and Tracts  Deliver efferent impulses from the brain to the spinal cord  Direct  Indirect pathways—pyramidal tracts p pathways—all others y Descending Pathways and Tracts  Involve two neurons: 1. Upper motor neurons  Pyramidal cells in primary motor cortex Ventral horn motor neurons Innervate skeletal muscles 2. Lower motor neurons   107 .

and all motor pathways except pyramidal pathways Also called the multineuronal pathways 108 .3/20/2012 The Direct (Pyramidal) System    Impulses from pyramidal neurons in the precentral gyri pass through the pyramidal (corticospinal)l tracts Axons synapse with interneurons or ventral horn y p motor neurons The direct pathway regulates fast and fine (skilled) movements Pyramidal cells (upper motor neurons) Primary motor cortex Internal capsule Cerebrum Midbrain Cerebellum Pons Cerebral peduncle Ventral corticospinal tract Pyramids Decussation of pyramid Lateral corticospinal tract Skeletal muscle Medulla oblongata Cervical spinal cord Lumbar spinal cord (a) Pyramidal (lateral and ventral corticospinal) pathways Somatic motor neurons (lower motor neurons) Figure 12.35a Indirect (Extrapyramidal) System   Includes the brain stem motor nuclei.

and eye movements that follow objects  Muscles Indirect (Extrapyramidal) System    Reticulospinal and vestibulospinal tracts—maintain balance Rubrospinal tracts—control flexor muscles Superior colliculi and tectospinal tracts mediate head movements in response to visual stimuli Cerebrum Red nucleus Midbrain Cerebellum Pons Rubrospinal tract Medulla oblongata Cervical spinal cord (b) Rubrospinal tract Figure 12. and regulate:  Axial muscles that maintain balance and posture controlling coarse movements g  Head. neck.3/20/2012 Indirect (Extrapyramidal) System  These pathways are complex and multisynaptic.35b 109 .

there is no voluntary or involuntary control of muscles  Muscles atrophy Spinal Cord Trauma  Spastic paralysis—damage to upper motor neurons of the primary motor cortex  Spinal neurons remain intact.3/20/2012 Spinal Cord Trauma  Functional losses  Parasthesias  Sensory loss  Paralysis  Loss of motor function Spinal Cord Trauma  Flaccid paralysis—severe damage to the ventral root or ventral horn cells  Impulses do not reach muscles. muscles are stimulated by reflex activity  No voluntary control of muscles 110 .

3/20/2012 Spinal Cord Trauma  Transection  Cross sectioning of the spinal cord at any level in total motor and sensory loss in regions inferior to the cut  Paraplegia—transection between T1 and L1  Quadriplegia—transection in the cervical region  Results Poliomyelitis     Destruction of the ventral horn motor neurons by the poliovirus Muscles atrophy Death may occur due to paralysis of respiratory muscles or cardiac arrest Survivors often develop postpolio syndrome many years later. as neurons are lost Amyotrophic Lateral Sclerosis (ALS)      Also called Lou Gehrig’s disease Involves progressive destruction of ventral horn motor neurons and fibers of the pyramidal tract Symptoms—loss of the ability to speak. swallow. attack by the immune system. or both 111 . and breathe Death typically occurs within five years Linked to glutamate excitotoxicity.

3/20/2012 Developmental Aspects of the CNS     CNS is established during the first month of development Gender-specific areas appear in both brain and spinal cord. alcohol and opiates). or infection can harm the developing CNS Smoking decreases oxygen in the blood.g. depending on presence or absence of fetal testosterone Maternal exposure to radiation. but these are not significant in healthy individuals until they reach their 80s Shrinkage of brain accelerates in old age g g Excessive use of alcohol causes signs of senility unrelated to the aging process 112 . drugs (e.. which can lead to neuron death and fetal brain damage Developmental Aspects of the CNS    The hypothalamus is one of the last areas of the CNS to develop Visual cortex develops slowly over the first 11 weeks Neuromuscular coordination progresses in superiorto-inferior and proximal-to-distal directions along with myelination Developmental Aspects of the CNS    Age brings some cognitive declines.

1 Sensory Receptors    Specialized to respond to changes in their environment (stimuli) Activation results in graded potentials that trigger nerve impulses p Sensation (awareness of stimulus) and perception (interpretation of the meaning of the stimulus) occur in the brain 113 .3/20/2012 Peripheral Nervous System (PNS)  All neural structures outside the brain  Sensory receptors nerves and associated ganglia M Motor endings g  Peripheral Central nervous system (CNS) Peripheral nervous system (PNS) Sensory (afferent) division Motor (efferent) division Somatic nervous system Autonomic nervous system (ANS) Sympathetic division Parasympathetic division Figure 13.

taste. excessive pressure.. stretch.g. pressure. pressure. smell..g. and temperature Most special sense organs 114 . extreme heat or cold. ll changes in blood chemistry) Nociceptors—sensitive to pain-causing stimuli (e. pain. retina) Chemoreceptors—respond to chemicals ( Ch d h i l (e.g.3/20/2012 Classification of Receptors  Based on:  Stimulus  Location  Structural S type complexity p y Classification by Stimulus Type      Mechanoreceptors—respond to touch. vibration. Exteroceptors    Respond to stimuli arising outside the body Receptors in the skin for touch. and itch Thermoreceptors—sensitive to changes in temperature Photoreceptors—respond to light energy (e. inflammatory chemicals) Classification by Location 1.

tissue stretch. pressure. equilibrium. Complex receptors (special sense organs)  Vision. pain. hearing. joints. Proprioceptors   Respond to stretch in skeletal muscles. smell. tendons. Interoceptors (visceroceptors)   Respond to stimuli arising in internal viscera and blood vessels Sensitive to chemical changes. Simple receptors for general senses:   Tactile sensations (touch.3/20/2012 Classification by Location 2. ligaments. temperature. and connective tissue coverings of bones and muscles Inform the brain of one’s movements Classification by Structural Complexity 1. stretch. and taste 2. and temperature changes Classification by Location 3. and muscle sense Unencapsulated (free) or encapsulated dendritic endings 115 . vibration).

3/20/2012 Unencapsulated Dendritic Endings  Thermoreceptors  Cold  Heat receptors (10–40ºC). in superficial dermis receptors (32–48ºC). in deeper dermis Unencapsulated Dendritic Endings  Nociceptors  Respond to: from damaged tissue outside the range of thermoreceptors  Pinching  Chemicals  Capsaicin  Temperatures Unencapsulated Dendritic Endings  Light touch receptors  Tactile  Hair (Merkel) discs follicle receptors 116 .

1 Encapsulated Dendritic Endings  All are mechanoreceptors Meissner’s (tactile) corpuscles—discriminative touch Pacinian (lamellated) corpuscles—deep pressure and vibration  Ruffini endings—deep continuous pressure  Muscle spindles—muscle stretch  Golgi tendon organs—stretch in tendons  Joint kinesthetic receptors—stretch in articular capsules   Table 13.1 117 .3/20/2012 Table 13.

3.3/20/2012 From Sensation to Perception    Survival depends upon sensation and perception Sensation: the awareness of changes in the internal and external environment Perception: the conscious interpretation of those stimuli Sensory Integration   Input comes from exteroceptors. 2. proprioceptors. and interoceptors Input is relayed toward the head. Receptor level—the sensor receptors Circuit level—ascending pathways Perceptual level—neuronal circuits in the cerebral p v cortex 118 . but is processed along the way g y Sensory Integration  Levels of neural integration in sensory systems: 1.

3/20/2012 3 Perceptual level (processing in cortical sensory centers) Motor cortex Somatosensory cortex Thalamus 2 Reticular formation Pons Circuit level Ci it l l (processing in Spinal ascending pathways) cord Cerebellum Medulla Free nerve endings (pain. warmth) Muscle spindle 1 Receptor level (sensory reception Joint and transmission kinesthetic to CNS) receptor Figure 13. cold. the receptor potential and generator potential are the same thing stimulus  receptor/generator potential in afferent neuron  action potential at first node of Ranvier 119 .2 Processing at the Receptor Level    Receptors have specificity for stimulus energy Stimulus must be applied in a receptive field Transduction occurs  Stimulus energy is converted into a graded potential called a receptor potential Processing at the Receptor Level  In general sense receptors.

and smell nociceptors and most proprioceptors  Tonic receptors adapt slowly or not at all  Examples: 120 .3/20/2012 Processing at the Receptor Level  In special sense organs: stimulus  receptor potential in receptor cell  release of neurotransmitter  generator potential in first-order sensory neuron  action potentials (if threshold is reached) Adaptation of Sensory Receptors  Adaptation is a change in sensitivity in the presence of a constant stimulus  Receptor  Receptor p membranes become less responsive potentials decline in frequency or stop p q y p Adaptation of Sensory Receptors  Phasic (fast-adapting) receptors signal the beginning or end of a stimulus  Examples: receptors for pressure. touch.

g.g. sweet or sour ( g. the melody in a piece of music) 121 ..3/20/2012 Processing at the Circuit Level   Pathways of three neurons conduct sensory impulses upward to the appropriate brain regions First-order neurons  Conduct impulses from the receptor level to the secondorder neurons in the CNS Transmit impulses to the thalamus or cerebellum Conduct impulses from the thalamus to the somatosensory cortex (perceptual level)  Second-order neurons   Third-order neurons  Processing at the Perceptual Level   Identification of the sensation depends on the specific location of the target neurons in the sensory cortex Aspects of sensory perception: Perceptual detection—ability to detect a stimulus (requires summation of impulses)  Magnitude estimation—intensity is coded in the frequency of impulses  Spatial discrimination—identifying the site or pattern of the stimulus (studied by the two-point discrimination test)  Main Aspects of Sensory Perception    Feature abstraction—identification of more complex aspects and several stimulus properties Quality discrimination—the ability to identify submodalities of a sensation (e.. tastes) Pattern recognition—recognition of familiar or significant patterns in stimuli (e.

K+. and bradykinin Impulses travel on fibers that release neurotransmitters glutamate and substance P Some pain impulses are blocked by inhibitory endogenous opioids Structure of a Nerve   Cordlike organ of the PNS Bundle of myelinated and unmyelinated peripheral axons enclosed by connective tissue Structure of a Nerve  Connective tissue coverings include:  Endoneurium—loose connective tissue that encloses axons and their myelin sheaths  Perineurium—coarse connective tissue that bundles fibers into fascicles  Epineurium—tough fibrous sheath around a nerve 122 . ATP. acids.3/20/2012 Perception of Pain     Warns of actual or impending tissue damage Stimuli include extreme pressure and temperature. histamine.

visceral) (Chapter 14) 123 . somatic) (Chapter 12) ganglia (motor.3/20/2012 Endoneurium Perineurium Epineurium Axon Myelin sheath Fascicle Blood vessels Figure 13.3b (b) Classification of Nerves    Most nerves are mixtures of afferent and efferent fibers and somatic and autonomic (visceral) fibers Pure sensory (afferent) or motor (efferent) nerves are rare Types of fibers in mixed nerves:   Somatic afferent and somatic efferent Visceral afferent and visceral efferent  Peripheral nerves classified as cranial or spinal nerves Ganglia  Contain neuron cell bodies associated with nerves  Dorsal  Autonomic root ganglia (sensory.

Figure 13.4 (2 of 4) 124 . axon will regenerate Involves coordinated activity among: Macrophages—remove d b i M h debris Schwann cells—form regeneration tube and secrete growth factors  Axons—regenerate damaged part    CNS oligodendrocytes bear growth-inhibiting proteins that prevent CNS fiber regeneration Endoneurium Schwann cells Droplets of myelin 1 The axon becomes fragmented at f t d t the injury site. Fragmented axon Site of nerve damage Figure 13.3/20/2012 Regeneration of Nerve Fibers    Mature neurons are amitotic If the soma of a damaged nerve is intact.4 (1 of 4) Schwann cell Macrophage 2 Macrophages clean out the dead axon distal a on to the injury.

or filaments. Single enlarging axon filament Figure 13. Fine axon sprouts or filaments Figure 13.3/20/2012 Aligning Schwann cells form regeneration tube 3 Axon sprouts. grow through a regeneration tube formed by Schwann cells.4 (3 of 4) Schwann cell Site of new myelin sheath formation f ti 4 The axon regenerates and a new myelin sheath forms.4 (4 of 4) Levels of Motor Control    Segmental level Projection level Precommand level 125 .

3/20/2012 Precommand Level (highest) • Cerebellum and basal nuclei • Programs and instructions (modified by feedback) Feedback Projection Level (middle) • Motor cortex (pyramidal system) and brain stem nuclei (vestibular. etc.) • Convey instructions to spinal cord motor neurons and send a copy of that information to higher levels Segmental Level (lowest) • Spinal cord • Contains central pattern generators (CPGs) Internal feedback Sensory input Reflex activity Motor output Figure 13. red. oft-repeated motor activity Projection Level  Consists of:  Upper motor neurons that direct the direct (pyramidal) system to produce voluntary skeletal muscle movements  Brain stem motor areas that oversee the indirect (extrapyramidal) system to control reflex and CPG CPGcontrolled motor actions  Projection motor pathways keep higher command levels informed of what is happening 126 .13a (a) Levels of motor control and their interactions Segmental Level    The lowest level of the motor hierarchy Central pattern generators (CPGs): segmental circuits that activate networks of ventral horn neurons to stimulate specific groups of muscles p g p Controls locomotion and specific. reticular formation.

3/20/2012 Precommand Level  Neurons in the cerebellum and basal nuclei  Regulate  Precisely motor activity start or stop movements  Coordinate movements with posture  Block unwanted movements  Monitor muscle tone  Perform unconscious planning and discharge in advance of willed movements Precommand Level  Cerebellum  Acts on motor pathways through projection areas of the brain stem  Acts on the motor cortex via the thalamus  Basal nuclei  Inhibit various motor centers under resting conditions Reflexes   Inborn (intrinsic) reflex: a rapid. p . involuntary.  Example: driving skills 127 . predictable motor response to a stimulus Learned (acquired) reflexes result from practice or repetition.

3/20/2012 Reflex Arc  Components of a reflex arc (neural path) 1. 5. 2. Receptor—site of stimulus action Sensory neuron—transmits afferent impulses to the CNS Integration center—either monosynaptic or polysynaptic region within the CNS Motor neuron—conducts efferent impulses from the integration center to an effector organ Effector—muscle fiber or gland cell that responds to the efferent impulses by contracting or secreting Stimulus Skin 1 Receptor 2 Sensory neuron 3 Integration center 4 Motor neuron 5 Effector Interneuron Spinal cord (in cross section) Figure 13. 4.14 Spinal Reflexes  Spinal somatic reflexes  Integration  Effectors center is in the spinal cord are skeletal muscle  Testing of somatic reflexes is important clinically to assess the condition of the nervous system 128 . 3.

3/20/2012 Stretch and Golgi Tendon Reflexes  For skeletal muscle activity to be smoothly coordinated. proprioceptor input is necessary  Muscle spindles inform the nervous system of the length of the muscle  Golgi tendon organs inform the brain as to the amount of tension in the muscle and tendons Muscle Spindles   Composed of 3–10 short intrafusal muscle fibers in a connective tissue capsule Intrafusal fibers  Noncontractile in their central regions (lack myofilaments)  Wrapped with two types of afferent endings: primary sensory endings of type Ia fibers and secondary sensory endings of type II fibers Muscle Spindles   Contractile end regions are innervated by gamma () efferent fibers that maintain spindle sensitivity Note: extrafusal fibers (contractile muscle fibers) y p ( ) are innervated by alpha () efferent fibers 129 .

increasing the rate of APs.3/20/2012 Secondary sensory endings (type II fiber) Efferent (motor) fiber to muscle spindle  Efferent (motor) fiber to extrafusal muscle fibers Extrafusal muscle fiber Intrafusal muscle fibers Primary sensory endings (type Ia fiber) Muscle spindle Connective tissue capsule Sensory fiber Golgi tendon organ Tendon Figure 13. 2. Action potentials (APs) are generated at a constant rate in the associated sensory (la) fiber. contract thereby stretching the spindle  Stretch causes an increased rate of impulses in Ia fibers Muscle spindle Intrafusal muscle fiber Primary sensory (la) nerve fiber Extrafusal muscle fiber Time (a) Unstretched muscle. Stretching activates the muscle spindle. b 130 . Time (b) Stretched muscle. External stretch of muscle and muscle spindle Internal stretch of muscle spindle:  Activating the  motor neurons stimulates the ends to contract. Figure 13.15 Muscle Spindles  Excited in two ways: 1.16a.

d Stretch Reflexes   Maintain muscle tone in large postural muscles Cause muscle contraction in response to increased muscle length (stretch) 131 . in length.3/20/2012 Muscle Spindles    Contracting the muscle reduces tension on the muscle spindle Sensitivity would be lost unless the muscle spindle is y p shortened by impulses in the  motor neurons – coactivation maintains the tension and sensitivity of the spindle during muscle contraction Time Time (c) Only motor (d) . The muscle spindle Muscle spindle tension is mainbecomes slack and no tained and it can APs are fired. Both extrafusal and Only the extrafusal intrafusal muscle muscle fibers contract. fibers contract.Coactivation. Figure 13. It is still signal changes unable to signal further length changes. neurons activated.16c.

Figure 13. the associated sensory neurons (blue) transmit afferent impulses neurons (blue) transmit afferent impulses at higher frequency to the spinal cord. 3b Efferent impulses of alpha motor 3b Efferent impulses of alpha motor neurons to antagonist muscles are neurons to antagonist muscles are reduced (reciprocal inhibition). inhibition of its antagonist. which resists or reverses the stretch. the stretched muscle p p . (quadriceps) contracts and the antagonists (hamstrings) relax Stretched muscle spindles initiate a stretch reflex. at higher frequency to the spinal cord. Afferent fibers also synapse with interneurons (green) that inhibit motor synapse with interneurons (green) that inhibit motor neurons (purple) controlling antagonistic muscles. which excite extrafusal fibers of the stretched muscle. which excite extrafusal fibers motor neurons (red).17 (1 of 2) Sensory Sensory neuron neuron 132 . neurons (purple) controlling antagonistic muscles. reduced (reciprocal inhibition). Cell body of Cell body of sensory neuron sensory neuron Initial stimulus Initial stimulus (muscle stretch) (muscle stretch) Spinal cord Spinal cord Muscle spindle Muscle spindle Antagonist muscle Antagonist muscle 3a Efferent impulses of alpha motor neurons 3a Efferent impulses of alpha motor neurons cause the stretched muscle to contract. Afferent fibers also of the stretched muscle.3/20/2012 Stretch Reflexes  How a stretch reflex works:  Stretch  Ia activates the muscle spindle sensory neurons synapse directly with  motor neurons in the spinal cord   motor neurons cause the stretched muscle to contract  All stretch reflexes are monosynaptic and ipsilateral Stretch Reflexes   Reciprocal inhibition also occurs . The events by which muscle stretch is damped The events by which muscle stretch is damped 1 When muscle spindles are activated 1 When muscle spindles are activated by stretch. cause the stretched muscle to contract. 2 The sensory neurons synapse directly with alpha 2 The sensory neurons synapse directly with alpha motor neurons (red). the associated sensory by stretch. Stretched muscle spindles initiate a stretch reflex. causing contraction of the stretched muscle and causing contraction of the stretched muscle and inhibition of its antagonist.Ia fibers synapse with interneurons that inhibit the  motor neurons of antagonistic muscles Example: In the patellar reflex. which resists or reverses the stretch.

3a The motor neurons (red) send activating impulses to the quadriceps causing it to contract. 3b The interneurons (green) make inhibitory synapses with ventral horn neurons (purple) that prevent the antagonist muscles (hamstrings) from resisting the contraction of the quadriceps. where synapses occur with motor neurons and interneurons.3/20/2012 The patellar (knee-jerk) reflex—a specific example of a stretch reflex 2 Quadriceps (extensors) 1 3a 3b 3b Patella Spinal cord (L2–L4) Muscle spindle Hamstrings (flexors) Patellar ligament 1 Tapping the patellar ligament excites muscle spindles in the quadriceps. extending the knee. 2 Afferent impulses (blue) travel to the spinal cord. Figure 13.17 (2 of 2) + – Excitatory synapse Inhibitory synapse Golgi Tendon Reflexes    Polysynaptic reflexes Help to prevent damage due to excessive stretch Important for smooth onset and termination of muscle contraction Golgi Tendon Reflexes  Produce muscle relaxation (lengthening) in response to tension Contraction or passive stretch activates Golgi tendon organs Afferent impulses are transmitted to spinal cord  Contracting muscle relaxes and the antagonist contracts (reciprocal activation)  Information transmitted simultaneously to the cerebellum is used to adjust muscle tension   133 .

Golgi tendon organs are activated. Figure 13. impulses to antagonist muscle cause it to contract. 2 Afferent fibers synapse with interneurons in the spinal cord. Muscle relaxes. Interneurons Quadriceps (extensors) Spinal S i l cord d Golgi tendon organ Hamstrings (flexors) 3a Efferent impulses + Excitatory synapse – Inhibitory synapse 3b Efferent to muscle with stretched tendon are damped. reducing tension.3/20/2012 1 Quadriceps strongly contracts.18 Flexor and Crossed-Extensor Reflexes  Flexor (withdrawal) reflex  Initiated  Causes by a painful stimulus automatic withdrawal of the threatened body and polysynaptic part  Ipsilateral Flexor and Crossed-Extensor Reflexes  Crossed extensor reflex  Occurs with flexor reflexes in weight-bearing limbs to maintain balance  Consists of an ipsilateral flexor reflex and a contralateral extensor reflex  The  The stimulated side is withdrawn (flexed) contralateral side is extended 134 .

Site of reciprocal activation: At the same time.19 Superficial Reflexes   Elicited by gentle cutaneous stimulation Depend on upper motor pathways and cord-level reflex arcs Superficial Reflexes  Plantar reflex  Stimulus: stroking lateral aspect of the sole of the foot downward flexion of the toes  Tests for function of corticospinal tracts p  Response: 135 .3/20/2012 + Excitatory synapse – Inhibitory synapse Interneurons Afferent fiber Efferent fibers Efferent fibers Extensor inhibited Flexor stimulated Arm movements Flexor inhibited Extensor stimulated Site of stimulus: a noxious stimulus causes a flexor reflex on the same side. Figure 13. the extensor muscles on the opposite side are activated. withdrawing that limb.

indicates corticospinal or motor cortex damage  Response: Superficial Reflexes  Abdominal reflexes  Cause contraction of abdominal muscles and movement of the umbilicus in response to stroking of the skin  Vary in intensity from one person to another  Absent when corticospinal tract lesions are present Developmental Aspects of the PNS  Spinal nerves branch from the developing spinal cord and neural crest cells  Supply both motor and sensory fibers to developing muscles to help direct their maturation  Cranial nerves innervate muscles of the head 136 .3/20/2012 Superficial Reflexes  Babinski’s sign  Stimulus: as above dorsiflexion of hallux and fanning of toes  Present in infants due to incomplete myelination p y  In adults.

3/20/2012 Developmental Aspects of the PNS    Distribution and growth of spinal nerves correlate with the segmented body plan Sensory receptors atrophy with age and muscle tone lessens due to loss of neurons. decreased . and slower central processing Peripheral nerves remain viable throughout life unless subjected to trauma 137 . numbers of synapses per neuron.

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