Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 I. II. Chapter 6 - Introduction A. Discuss evolution of nervous system B. One of two communication methods within the body 1. Other is endocrine or hormonal 2. Modern concept is a neuroendocrine system 3. No evidence for a separate "mind" - all neural functions should be attributable to functions of brain cells and biochemistry 4. Figure 6-1, page 154 Structure and maintenance of neurons A. Basic unit of the nervous system is the neuron, Figure 6-2a, page 155. 1. Cell body - site of nucleus 2. Dendrites - branches of cell body 3. Axon - a single process extending from cell body a) Initial segment (trigger zone) also known as axon hillock b) Neural signals pass through cell body to axon hillock and then down axon - more later in chapter c) Axon may have branches called collaterals d) Axon ends in axon terminal or terminus (one terminus, two termini) e) Some axons have varicosities - more later f) Axons vary in length from more than a meter to quite short (1) Projection neurons - connect with distant parts of the nervous system or peripheral organs (2) Local circuit neurons - short and connect only with cells in immediate vicinity 4. Myelin - a fatty material formed from the plasma membrane of Schwann cells during development, Figure 6-2b & c, page 155. a) Derived from neural crest cells in embryo and migrate to peripheral neurons b) Nodes of Ranvier spaces between myelin-forming cells - more later Important the plasma membrane is primarily lipid which is a good resistor to electrical flow - more later d) In the CNS the myelin-forming cells are the oligodendroglia (a kind of glial cell) Axon transport a) Materials and some organelles have to be transported from cell body where they are synthesized to the terminus b) Transported on cytoskeleton c) Transport can go from terminus to cell body c) 5. Page 1 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 III. Functional classes of neurons, Figure 6-3, page 156, Table 6-1, page 157 A. Afferent neurons - part outside CNS 1. Have receptors at peripheral end, NOTE: this is a different use of the word receptor than used previously 2. Propagate electrochemical signals from receptor into brain or spinal cord 3. Do not have dendrites B. Efferent neurons 1. Propagate electrochemical signals from CNS out to effector cells such as muscle or glands 2. Efferent neurons go to effectors! 3. Cell body, dendrites, and a little of the axon is in the CNS most of the axon is in the periphery C. Interneurons - all within CNS 1. Integrators and signal changers 2. Ninety (99%) of all neurons 3. Number of interneurons in a reflex arc differ depending on the complexity D. For each afferent neuron entering CNS there are about 200,000 interneurons and 10 efferent neurons E. Synapse the connection by which two excitable cells communicate Figure 6-4, page 157 illustrates presynaptic/postsynaptic change 2. A neuron may have thousands of presynaptic neurons synapsing on its surface IV. Glial cells comprise 90% of the nervous system, the other 10% are neurons A. Also known as neuroglia B. These sustain neurons metabolically, support them physically and help regulate the ionic concentrations in the extracellular space C. Volume-wise: neurons are 50% and glial cells are 50% 1. D. E. V. Oligodendroglia myelin of axons in the CNS Astroglia multiple functions 1. Help regulate extracellular fluid 2. Sustain neurons metabolically 3. Migrate during development 4. Secrete growth factors F. Microglia may perform immune function in CNS Neural Growth and Regeneration A. Neurons or glial develop from neural ectoderm (neuroblasts, aka stem cells) and migrate to eventual adult site in controlled patterns 1. Growth cone, forms the tip of each extending axon and is involved in finding the correct route and final target for the process Page 2 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 a) As the axon grows, it is guided along the surfaces of other cells, most commonly glial cells b) Which route is followed depends largely on attracting, supporting, deflecting, or inhibiting influences exerted by several types of molecules. Some of these molecules, such as cell adhesion molecules (CAM), reside on the membranes of the glia and embryonic neurons. Others are soluble neurotropic factors (growth factors for neural tissue) in the extracellular fluid surrounding the growth cone or its distant target 2. Normally, after growth and projection of the axons, many of the newly formed neurons and synapses degenerate. As many as 50 to 70 percent of neurons die by apoptosis in some regions of the developing nervous system! Not known why. 3. Division of neuron precursors is largely complete before birth, and after early infancy new neurons are formed at a slower pace to replace those that die 4. Severed axons can repair themselves and regain significant function provided that the damage occurs outside the central nervous system and does not affect the neuron's cell body. a) After repairable injury, the axon segment now separated from the cell body degenerates. b) The proximal part of the axon (the stump still attached to the cell body) then gives rise to a growth cone, which grows out to the effector organ so that in some cases function is restored 5. Severed axons within the central nervous system attempt sprouting, but no significant regeneration of the axon occurs across the damaged site, and there are no well-documented reports of significant function return. 6. In humans, spinal injuries typically crush rather than cut the tissue, leaving the axons intact. a) In this case, a primary problem is self-destruction (apoptosis) of the nearby oligodendroglia b) When these cells die and their associated axons lose their myelin coat, the axons cannot transmit information effectively 7. Researchers are attempting a variety of measures to provide an environment that will support axonal regeneration in the central nervous system. They are creating tubes to support regrowth of the severed axons, VI. Basic Principles of Electricity A. Molecules may have a positive charge, negative charge or no charge (neutral) and macromolecules may have both + and - with a net of one or the other or neutral B. Electrical definitions 1. Opposite charges attract 2. Similar charges repel Page 3 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 a) 3. 4. The energy required to separate opposite charges or bring similar charges together is called potential or potential difference b) The unit of energy is voltage, a thousands of a volt is a millivolt (1000 mV = 1 volt) c) Figure 6-5, page 160 Movement of charged particles creates a current Resistance is the hindrance to current Ohm’s Law I = V/R V = voltage, R = resistance, I = current VII. The Resting Membrane Potential A. All cells measured thus far exhibit a potential difference between the inside of the cell and the outside - this is called the resting membrane potential and it is negative inside 1. Use analogy with battery 2. Measured with a voltmeter (microvoltmeter), Figure 6-6 a & b, page 161 3. Distribution of ions across cell membrane, Table 6-2, page 162 Ion Na+ ClK+ 4. 5. 6. Extracellular 150 110 5 Intracellular 15 10 140 The resting membrane potential is determined mainly by two factors a) The differences in ion concentration of the intracellular and extracellular fluids b) The permeabilities of the plasma membrane to the different ion species c) Figure 6-7, page 161 Diffusion potentials a) Sodium and potassium are most important in resting membrane potential b) Figure 6-8, page 162; membrane is impermeable to sodium ion but allows potassium to flow freely (1) The net result is a diffusion potential created by the concentration difference of potassium and the electrochemical potential (2) The equilibrium potential is achieved when the electrical force is equal and opposite to the chemical gradient (3) Figure 6-9, page 163; potential created by only sodium ion being freely diffusible Nernst equation → E = 60 log10 [C]o / [ C]I (for any ion) Page 4 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 7. B. Appendix D, page 791, Goldman equation for more than one ion determining potential The resting membrane potential 1. A potential difference can be realized in a resting nerve cell under the following conditions a) Potassium ion is much greater inside cell than outside b) Cell membrane is 50 to 75 times more permeable to potassium than to sodium c) Forces acting on potassium ion Figure 6-10, page 164 Forces acting on sodium ion Figure 6-11, page 164 2. Leakage can occur through membrane so that the actual potential difference is not at the theoretical potassium equilibrium potential a) The Na-K-ATPase pump compensates, Figure 6-12, page 165 b) Three sodiums out for every two potassiums back in known as an electrogenic pump but is usually a rather small contribution to the potential difference 3. Chloride ion a) In most cells chloride ion moves passively with the concentration and/or electrical gradient and do not contribute to determination of the resting membrane potential b) Some cells can actively pump chloride and this can contribute to resting membrane potential 4. Most of the negative charge in neurons is contributed to by the negatively charged proteins - they do not diffuse through the membrane readily or at all VIII. Graded Potentials and Action Potentials A. Two kinds of "disruptions" can occur in resting membrane potential graded potentials and action potentials 1. Terms - Table 6-3, page 166 2. Figure 6-13, page 165 a) Depolarized - potential toward zero b) Polarized - potential away from zero, repolarized refers to re-establishing the resting membrane potential c) Hyperpolarized - potential difference greater than original d) d) B. C. Overshoot inside of cell becomes positive Recording depolarization Figure 6-14, page 167 Graded (receptor potentials, synaptic potentials, end-plate potentials and pacemaker potentials) 1. Can be a depolarization or hyperpolarization, Figure 6-15, page 167 Page 5 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 2. D. Graded response - amplitude varies with conditions of the initiating events 3. Is conducted decrementally i.e., amplitude decreases with distance 4. Graded response; can be summed temporally or spatially 5. Has no threshold (explain with action potential) 6. Has no refractory period (explain with action potential) 7. Duration varies with initiating conditions (explain with action potential) 8. Initiated by stimulus (receptor), by neurotransmitter (synapse) or spontaneously (electrical) 9. Leakage of charge, Figure 6-16, page 167 Action potentials - Figure 6-17 A & B, page 168 1. Rapid alteration or transient change in the resting membrane potential is known as an action potential a) Cells that undergo such change are termed excitable - mainly neurons and muscle cells b) The action potential can also be propagated (conducted) long distances - later 2. Ionic basis of the action potential - "ionic flux theory" by Hodgkin, Huxley and Eccles (won Nobel Prize in 1963) a) An electric current opens sodium gates (channels) and sodium ion flows down its concentration and electrical gradient - the influx of sodium ion causes depolarization b) When the potential passes zero and approaches about +35 mV then the sodium gates close and potassium gates open - the efflux of potassium ion repolarizes the cell (after hyperpolarization) c) THE NA-K PUMP DOES NOT IMMEDIATELY REESTABLISH THE RESTING MEMBRANE POTENTIAL! (1) The resting membrane potential is reestablished by the efflux of potassium (2) If the Na-K pump is poisoned by oubain action potentials can still occur on a neuron for a long time (3) The pump acts only after many action potentials have been elicited and an imbalance in the ion distribution occurs very few ions are involved in a single action potential 3. Differences between Voltage-Gated Sodium and Potassium Channels a) Sodium channels open faster in response to a given voltage change b) Once activated, sodium channels close more rapidly c) Sodium channels inactivet, cycling through an inactive phase Page 6 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 4. 5. 6. 7. Mechanism of ion-channel changes a) The basis of the configurational changes in the gates or channels is not known b) In the case of sodium ion it is a positive feedback mechanism, Figure 6-189, page 169 c) Channels that respond to changes in membrane potential are termed voltage-sensitive channels d) Some nonneural cells are depolarized by the influx of Ca2+ e) Local anesthetics can prevent action potentials by preventing the channels from "opening" and "closing" - include novocaine & xylocaine f) Table 6-4, page 170 → Differences between voltage-gated sodium and potassium channels Threshold, Figure 6-19, page 170 a) The membrane potential at which the net movement of ions across the membrane first changes from outward to inward is the threshold b) The stimulus strength is called the threshold stimulus c) Subthreshold stimuli can occur but can’t elicit an action potential - they can summate, however d) A suprathreshold stimuli does not elicit a larger action potential since action potentials exhibit what is called ALL-OR-NONE, that is they have the same amplitude regardless of stimulus strength as long as the stimulus strength is at threshold or above e) The information conveyed by action potentials is analogous to an FM radio, frequency modulated, rather than an AM radio, amplitude modulated since the amplitude doesn’t change only the frequency and/or rate can convey information (THIS IS AN IMPORTANT CONCEPT!) Refractory periods a) Refractory means obstinate or unwilling b) The absolute refractory period follows an action potential during which another action potential can NOT be elicited no matter how strong the stimulus strength c) Following the absolute refractory period there is a relative refractory period during which an action potential can be elicited but at a strength of stimulus above threshold and decreasing with time Action potential propagation, Figure 6-20, page 172 a) An action potential at a given "active site" is accompanied by the flow of charged particles (ions) - this creates a local current b) The local current flow can alter the sodium gates in the adjacent "active site" and allow sodium to enter, Page 7 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 E. that initial influx then is followed by more gates opening, i.e. positive feedback c) The action potential can then be propagated down the axon in only one direction since the "active sites" are refractory - explain d) This propagation can be greatly speeded up by the presence of myelin, Figure 6-21, page 173 (1) The current is prevented from influencing the membrane by the insulating sheath except at the nodes of Ranvier (2) This kind of propagation is called saltatory conduction or "jumping" conduction (3) Range of conducting velocities: 1 mile/hour in small-diameter unmyelinated neurons to 400 mile/hour in large-diameter myelinated neurons (1 mph takes 4 sec to go head to toe but 10 milliseconds at 400 mph) 8. Initiation of action potentials a) Action potential are initiated in a variety of ways which will be discussed in later chapters b) Most are preceded by a graded potential of some kind - a pacemaker potential is an example See Table 6-5, page 174 for comparison with action potential SECTION C - SYNAPSES IX. X. Introduction to Synapses A. Synapses - Functionally and anatomically specialized connections between "excitable" cells 1. Neural-neural 2. Neural-muscular 3. Neural-glandular B. Estimate there are 1014 synapses in the CNS C. Activity is transferred from axon to the next cell via a neurotransmitter in chemical synapses 1. Excitatory synapses 2. Inhibitory synapses D. Neural relationships 1. Convergence and divergence - Figure 6-22 page 176 2. Diagrammatic representation of synapse – Figure 6-23, pag 176 3. Neurons function as neural integrators Functional Anatomy of Synapses, Figure 6-24 a, b, c, d, page 198-199 A. Two types of synapses: electrical and chemical a) Chemical synapses are most common in the nervous system b) There are electrical synapses which are not common and involve gap junctions Page 8 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 B. C. D. E. 2. Few electrical in mammals 3. Information from here on is about chemical synapses Chemical synapses, Figure 25, page 177 1. Axon ends in slight swelling, the axon terminal 2. Synaptic cleft separates two cells 3. In the presynaptic cell are vesicles containing neurotransmitters a) Action potentials arriving at terminus causes influx of calcium ion into the presynaptic terminus b) This triggers exocytosis - fusing of membrane of vesicle and plasma membrane at cleft c) Neurotransmitter is released and diffuses across cleft to postsynaptic membrane or subsynaptic membrane d) The neurotransmitter combines with appropriate receptors to effect receptor-operated channels, and G-protein receptors, Chapter 6 & 7 (1) More than one transmitter may be released (2) Second transmitter is called cotransmitter e) Transmission is only in one direction since the vesicles are on only one side and receptors are only on one side f) The receptors on the postsynaptic cell membrane is limited to the space right under the synapse in normal cell g) The events result in a synaptic delay Excitatory chemical synapses 1. Transmitter "causes" depolarization, Figure 6-26, page 178 2. Opens postsynaptic channels that are permeable to sodium, potassium and other small positively charged ions 3. The electrical and concentration gradients favor sodium influx whereas potassium has opposing forces 4. Called EPSP, Excitatory Post Synaptic Potential a) A graded potential with all the properties b) Brings axon hillock potential toward threshold Inhibitory chemical synapses 1. Transmitter "causes" hyperpolarization, Figure 6-27, page 179 2. Opens postsynaptic channels that are permeable to potassium, chloride and other small positively charged ions sodium channel are not effected 3. Called IPSP, Inhibitory Post Synaptic Potential a) A graded potential with all the properties b) Brings axon hillock potential away from threshold 4. Cells that actively transport chloride can hyperpolarize by increasing chloride permeability or an IPSP Inactivation Page 9 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 1. The neurotransmitter must be removed from cleft after performing its "role" 2. Transmitter can be inactivated enzymatically 3. Transmitter can diffuse away from the site 4. Transmitter can be actively transported back into the axon terminus or into glial cells in some cases XI. Activation of the Postsynaptic Cell A. The normal situation is not one-on-one transmission but rather many presynaptic neurons, both excitatory and inhibitory, that interact to produce action potentials 1. Figure 6-28, page 179 - shows interaction 2. Exhibit temporal and spatial summation occurs, Figure 6-29, page 180 3. Different parts of neuron can have different thresholds 4. Figure 6-30, page 180 - Excitatory and inhibitory current flows a) B. C. Excitatory positive charges flow into cell body b) Inhibitory positive charges flow out of cell Integration 1. In CNS may have influence of local currents on neighboring unmyelinated neurons 2. Can have influence from other substances, both "normal" and "abnormal" binding at receptors Synaptic Strength 1. The synapse can be effected by any event that effects the amount of transmitter released - more transmitter, more depolarization or hyperpolarization 2. Factors that can determine synaptic effectiveness, Table 66, page 183 a) Presynaptic factors (1) Availability of neurotransmitters (a) Availability of precursor molecules (b) Amount (or activity) of the ratelimiting enzyme in neurotransmitter synthesis pathway (2) Axon terminal membrane potential (3) Axon terminal residual calcium (4) Activation of membrane receptors on presynaptic terminal (a) Presynaptic (axon-axon) synapses (b) Autoreceptors (c) Other receptors (5) Certain drugs and diseases b) Postsynaptic factors (1) Immediate past history of electrical state of postsynaptic membrane, that is, facilitation Page 10 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 or inhibition from temporal or spatial summation (2) Effects of other transmitter-type chemicals acting on postsynaptic neuron (3) Certain drugs or diseases 3. Action of other neurons that form synapses on the axon terminus (presynaptic inhibition or facilitation), Figure 6-31 page 181 4. Modification of Synaptic Transmission by Drugs and Disease, Figure 6-32, page 182 a) Agonists - create similar response than normal transmitters b) Antagonists - block normal action XII. Neurotransmitters and Neuromodulators A. Neuromodulators are chemicals that act on neurons but are not quite transmitters - very vague in defining 1. Have long term effects 2. Amplify or dampen the effectiveness of ongoing synaptic activity by altering the neurons at nonsynaptic sites, i.e., extrasynaptic receptors 3. Some neuromodulators can be hormones B. Four classes of neurotransmitters or neuromodulators and one miscellaneous category, Table 6-7, page 205 1. Many of these chemicals do act also as hormones and paracrines 2. Generally speaking axons release only one transmitter and only that transmitter, there are exceptions in the number of transmitters released from a single axon 3. A single transmitter may produce EPSPs at one synapse but IPSPs at another - i.e. the response is a property of the subsynaptic membrane receptors and not the messenger C. Acetylcholine (Ach) 1. Probably most common transmitter, certainly the best known and first described - found in neural-muscular synapses and in brain 2. Two types of Ach receptors a) Nicotinic - receptors respond to nicotine (1) Example of a receptor that contains the channel (2) Receptor is an allosteric protein which changes configuration in presence of Ach b) Muscarinic - receptors respond to muscarine (1) Couple with G-proteins (2) Alter enzyme activity and ion channels 3. Choline+CoA in cytoplasm of axon terminus and stored in vesicles (CoA is derived from pantothenic acid 4. Inactivated by acetylcholinesterase Page 11 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 5. D. E. F. Degeneration of cholinergic system in brain can lead to Alzheimer's disease Biogenic amines R-NH2 (most derived from amino acid tyrosine) 1. Catecholamines, Figure 6-34, page 206 a) Norepinephrine and epinephrine (1) Receptors are adrenergic or noradrenergic (2) Alpha-adrenergic receptors and Betaadrenergic receptors are influenced by different drugs b) Actively transported back into presynaptic cell and this stops activity c) Distributed widely d) Slow time-courses - may act on neurons being acted upon by other more rapidly acting transmitters e) Found in autonomic nervous system f) Dopamine - cocaine interferes with dopamine reuptake 2. Serotonin a) Produced from tryptophan in CNS b) Important in neural pathways controlling states of consciousness and mood (sleep) c) Found also in platelets Amino acid neurotransmitters 1. Act directly as transmitters 2. Include glycine, glutamate and aspartate a) Glycine can act as inhibitory transmitter b) Aspartate and glutamate are among most potent transmitters in CNS - mostly excitatory 3. GABA, gamma-aminobutyric acid, is not an alpha amino acid but is an amino acid 4. GABA is major inhibitory transmitter in the brain Neuropeptides 1. More than eighty-five have been thus far described in neural tissue although they have been found and described to function as hormones or paracrine agents in nonneural tissue 2. Derived from large proteins synthesized in cell body and are attached to ribosomes a) Prohormones are packaged into vesicles and transported in neuron where needed (terminus or varicosities) b) When needed, the hormones are cleaved off by peptidases c) Some are called polyproteins d) The neurons are called peptidergic and usually are cosecreted with another type of neurotransmitter 3. Types derived from prohormones or preprohormones Page 12 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 a) b) G. XIII. A. B. XIV. A. B. C. Endorphins (endogenous morphines), dynorphins and enkephalins are known as opioid peptides small, very short lived and for the most part are analgesic in nature Substance P - peptide that is in a group of transmitters called tachykinins for neurons that have to do with pain (nociceptive) and may be the link between the nervous system and immunity system Miscellaneous 1. Nitric oxide serves as neurotransmitter (neural-neural and neural-effector) a) Produced from arginine in a cell and that triggers cGMP activity in effector cell b) Plays a role in learning, development, drug tolerance, penile erection and sensory and motor modulation among others c) Is found outside the CNS and PNS in the cardiovascular and immune systems d) Basis of action of nitroglycerin 2. Carbon monoxide – as above 3. Adenosine and ATP neurotransmitter (fast acting) Neuroeffector Communication Other type of synapses other than neural-neural Covered in subsequent chapters Structure of the Nervous System Terms 1. "Nerve" is a collection of many neurons 2. A nerve in the CNS is called a pathway, tract or when connecting, a commissure 3. In the PNS, groups of neuron cell bodies is called a ganglia 4. the CNS, groups of neuron cell bodies is called a nuclei 5. Pathways, Figure 6-35, page 210 a) Long neural pathways b) Multineuronal or multisynaptic pathways Categorized anatomically and physiologically and are divided into two main categories anatomically (CNS & PNS) 1. Central nervous system a) Brain and brainstem b) Spinal cord 2. Peripheral nervous system a) Spinal nerves b) Cranial nerves Central nervous system: brain 1. Six subdivisions - Figure 6-34, page 189 a) Cerebrum b) Diencephalon Page 13 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 2. 3. 4. 5. c) Midbrain d) Pons e) Medulla oblongata f) Cerebellum Cerebral ventricles a) Four interconnected containing cerebrospinal fluid b) Figure 6-36, page 191 Brainstem - stalk of brain that relays all information between spinal cord, cerebrum and cerebellum (to cerebellum via cerebellar peduncles) a) Midbrain b) Pons c) Medulla oblongata d) Reticular formation - also known as reticular core and reticular activating system {RAS}, (old terms) (1) Absolutely essential for life (2) Runs length of brain stem through all three parts (3) Houses cardiovascular, respiratory, swallowing and vomiting centers discussed later (4) Sends output and receives from almost all parts of nervous system Branches to: (a) Thalamus (b) Base of forebrain (c) Cerebral cortex (d) Connects cerebellum and spinal cord (reticulospinal pathway) (5) Ten of the 12 cranial nerves have nuclei in the reticular formation Cerebellum - has a cortex and cerebellar nuclei (chiefly involved with skeletal muscle function) a) Connected to brainstem by cerebellar peduncles b) Does not initiate or directly coordinate activity c) Receives input from muscles, joints, skin, eyes and ears among others d) Acts as "quality control center", telling higher brain centers what muscles have done Forebrain - what is left after brainstem and cerebellum are excluded a) Cerebrum - right and left cerebral hemispheres, Figure 6-35, page 190 (1) Hemispheres are connected by axon bundles known as commissures, the corpus callosum being the largest Page 14 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 (2) (3) Areas within a hemisphere are connected by association fibers Outer portion is cerebral cortex, 3 mm thick and is the most sophisticated part with cortical neurons of two basic types Pyramidal cells (major output to the rest of the nervous system) (b) Nonpyramidal cells (c) Brings together afferent (sensory) information for perception and control of the motor system (4) Divided into lobes - frontal, parietal, occipital and temporal, Figure 6-41 page 214 (5) Subcortical areas lie underneath cortex (a) Basal ganglia - role in movement and some complex behavior (b) Substantia nigra - movement (Parkinson's disease and dopamine) b) Diencephalon (a) Thalamus - relay and integrating station for sensory input to cortex and is important in motor control; part of reticular core is found here (b) Hypothalamus - lies below thalamus, very important part of brain that is responsible for the integration and control of many systems (c) Limbic system - associated with learning and emotional behavior; not really a separate part of brain but made up of connections from other parts of brain, Figure 6-36, page 191 Central nervous system: spinal cord 1. Slender cylinder, Figure 6-37, page 193 and Figure 6-38, page 194 a) Grey matter - central region filled with interneurons b) White matter - surrounding, color due to myelin sheaths 2. The interneurons form tracts and pathways a) Afferent neurons enter the CNS via the dorsal roots and form the dorsal root ganglia which are the cell bodies of the afferent neurons b) Efferent neurons leave the CNS via the ventral roots (cell bodies in CNS) and form spinal nerves - spinal nerves are in pairs from each side of the spinal cord Major parts of brain: summary - Table 6-8, page 192 Peripheral nervous system (a) D. E. F. Page 15 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 1. 2. 3. Bundles of neurons are called nerves a) Thirty-one (31) pairs of spinal nerves b) Twelve (12) pairs of cranial nerves (discuss misnaming of optic nerve here) - Table 6-9, page 195 (do not memorize the cranial nerves - that’s for anatomy!) c) Most are myelinated, some not, however d) The nerves are divided into afferent and efferent Table 6-10, page 195 Peripheral nervous system: afferent division a) Sensory neurons with cell bodies in ganglia outside CNS b) A second process, the central process, relays information into the CNS where it branches c) Afferent neurons are sometimes called primary afferents or first-order neurons Peripheral nervous system: efferent division, divided into somatic and autonomic nervous systems, please NOTE autonomic not automatic - Figure 6-39, page 196 a) Differences between somatic and autonomic nervous systems, Table 6-11, page 195 b) Somatic c) All the fibers going from the CNS to skeletal muscles d) Cell bodies are in groups in brain or spinal cord e) Leave CNS and pass, without intermediate synapsing, to effector organs (skeletal muscle) (they eventually synapse on effector organ) f) The neurotransmitter is acetylcholine g) Are normally called motor neurons h) Autonomic, Figure 6-40, page 197 (1) Innervate cardiac muscle, smooth (involuntary) muscle, glands, and enteric nervous system of gastrointestinal tract (2) Sympathetic thoracolumbar (a) (b) Figure 6-41, page 198 Sympathetic trunk and spinal cord The sympathetic anatomy or wiring allows this portion of the autonomic nervous system to act as a unit which is, of course, adaptive (3) (4) Parasympathetic craniosacral Have two neurons and one synapse in periphery - the synapses are in autonomic ganglia (sympathetic trunk is most prominent) (5) Preganglion before ganglion postganglion after ganglion Page 16 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 (6) Parasympathetic preganglionic & postganglionic transmitter is acetylcholine (cholinergic fibers) in all cases; the sympathetic transmitter in preganglionic fibers is acetylcholine and in postganglionic cells is norepinephrine (adrenergic fibers) with one exception, Figure 6-42, page 199, Table 6-12, page 198 (7) Nicotinic, muscarine, alpha-adrenergic and beta-adrenergic receptors on autonomic fibers, Table 6-12, page 219 i) Dual innervation, Figure 6-44, page 218 (1) Many organs get sympathetic and parasympathetic innervation (2) some ability to control autonomic responses is possible in humans and experimental animals - basis of biofeedback Do not attempt to memorize Table 6-13, pages 200 now, it will be covered in subsequent chapters - merely review for appreciation of dual innervation THE BRAIN HAS TOP PRIORITY FOR OXYGEN AND GLUCOSE! XV. Blood Supply, Blood-Brain Barrier Phenomena and Cerebrospinal Fluid A. The brain is a very fragile organ and must have a constant blood supply with oxygen and glucose (little glycogen stores) 1. Little glycogen stores 2. Brain is 2% of body weight but receives 15% of blood supply 3. Brain covering between bony tissue soft neural tissue meninges a) dura mater - next to bone b) arachnoid - middle c) pia mater - next to brain B. Blood-brain barrier – protects immunologically privileged area 1. This is an anatomical and physiological phenomenon that separates the brain tissue from the plasma of the blood. a) The key are the endothelial cells which compose the capillary walls (1) They have tight junctions binding them together (2) The tight junctions prevent most things from moving from plasma into brain cerebrospinal fluid (3) The astrocytes which surround the capillaries are not a major part of the bloodbrain barrier b) Lipid soluble molecules cross barrier readily nicotine, ethanol, heroin Page 17 of 18 Chapter 6 - Neural Control Mechanisms Human Physiology, pages 153-204 c) d) Picture the wall of the capillary as having two unit membranes (1) Toward brain is called antiluminal membrane (2) Toward lumen of capillary is called luminal membrane (3) The two sides of the system are not symmetrical in the nature of the carriers that are there for some substances Water soluble molecules cross less readily or not at all Glucose main energy source, carried by specific membrane carrier (D-glucose was readily transported, L-glucose was not), there many glucose carriers available in the endothelial membrane (2) For glucose, the carriers are symmetrical and "work" down a concentration gradient no extra energy needed (facilitated diffusion) e) Amino acids - three categories [large neutral (10), small neutral and basic and acidic] f) Metabolic blood-brain barrier - enzymes in the endothelial cell modify substance and render them incapable of entering brain, example is L-DOPA g) Effect of sugar infusion 2. The chemical composition of the extracellular fluid of the brain is carefully regulated Cerebrospinal fluid - Figure 6-43, page 201 1. Secreted by the choroid plexus 2. Is found in the ventricles of the brain 3. Circulates and is reabsorbed into the blood stream from the top surface of the brain through one-way valves in large veins 4. It is also carefully regulated (1) C. Page 18 of 18