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Neuromuscular Fundamentals: How Our Musculature is Controlled

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This book is rather unique in its approach and coverage. The approach is essentially that of an engineering textbook, emphasizing the quantitative aspects and highlighting the fundamentals and basic concepts involved. The coverage progresses in a logical and systematic manner from the subcellular, starting with the electrophysiology of the cell membrane, then proceeding to synapses, neurons, and muscle, before considering neuronal motor ensembles and the neuromuscular system as a whole. Simple, clear, and comprehensive explanations are given throughout. After an introductory chapter on some background material in biology, biophysics, and chemical kinetics, a substantial part of the book (Chapters 2-8) necessarily covers in considerable detail the basic components and processes that underlie the electrical and associated activities of the nervous system. The remaining chapters of the book (Chapters 9-13) focus on the neuromuscular system, starting with the structure of muscle cells, the generation of force by muscular contraction, and muscle receptors. The last chapter examines aspects of the control of movement, motor learning and memory, the maintenance of posture, and locomotion, and critically examines some of the theories that have been advanced to explain how movement is controlled. The book is intended for undergraduate or graduate students in the natural sciences, mathematics, or engineering who seek a deeper understanding of the fundamentals of neuroscience and the somatomotor system, in accordance with the aforementioned objectives. The book can serve as a textbook for a one-semester course on the neuromuscular system or as a reference in a more general course on neuroscience.

  • Provides a thorough analytical treatment of membrane electrophysiology, starting from the first principles

  • Emphasizes strongly the basic and fundamental concepts throughout

  • Discusses thoroughly the essential features and properties of the basic constituents of the nervous system, that is, neurons and synapses, including the neuromuscular junction

  • Explains the main aspects of posture, locomotion, and control of movement

  • Includes practice problems throughout the text and a solutions manual will be available for adopting professors

Nassir Sabah is professor of biomedical engineering in the electrical and computer engineering department at the American University of Beirut, Lebanon. He received his B.Sc. (Hons. Class I) and his M.Sc. in electrical engineering from the University of Birmingham, U.K., and his Ph.D. in biophysical sciences from the State University of New York (SUNY/Buffalo). He has served as Chairman of the Electrical Engineering Department, Director of the Institute of Computer Studies, and Dean of the Faculty of Engineering and Architecture at the American University of Beirut. In these capacities, he was responsible for the development of programs, curricula, and courses in electrical, biomedical, communications, and computer engineering.

Professor Sabah has extensive professional experience in the fields of electrical engineering, electronics, and computer systems, with more than 35 years’ teaching experience in neuroengineering, biomedical engineering, electronics, and electric circuits. He has over 100 technical publications, mainly in neurophysiology, biophysics, and biomedical instrumentation.

He has served on numerous committees and panels in Lebanon and the region. He is a Fellow of the Institution of Engineering and Technology (IET, U.K.), a member of the American Association for the Advancement of Science (AAAS), and a member of the American Society for Engineering Education (ASEE).

Author(s): Nassir H. Sabah
Publisher: CRC Press
Year: 2020

Language: English
Pages: 574
City: Boca Raton

Cover
Half Title
Title Page
Copyright Page
Dedication
Brief Contents
Table of Contents
Preface
Acknowledgments
Convention for Symbols
Author Biography
Chapter 1 Introduction: Background Material
Objective and Overview
1.1 Living Cells
1.1.1 Endoplasmic Reticulum
1.1.2 Mitochondria
1.1.3 Cytoskeleton
1.1.4 Endocytosis and Exocytosis
1.2 Neurons and Glia
1.2.1 Neurons
1.2.2 Axonal Transport
1.2.3 Glial Cells
1.3 Organization of the Nervous System
1.4 Diffusion, Fluxes, and Potentials
1.4.1 Chemical Potential
1.4.2 Electrochemical Potential
1.4.3 Permeability
1.5 Ionic Equilibriums
1.5.1 Osmotic Equilibrium
1.5.2 Basic Ionic Equilibrium
1.5.3 Equilibrium Voltage
1.5.4 Gibbs–Donnan Equilibrium
1.6 Chemical Kinetics
1.6.1 Reaction Rates
1.6.2 Order of Reactions
1.6.3 Reversible Reactions
1.6.4 Kinetic Models of Ion Channel Gating
Summary of Main Concepts
Chapter 2 The Cell Membrane in the Steady State
Objective and Overview
Learning Objectives
2.1 Structure of the Cell Membrane
2.1.1 Aqueous Pores
2.2 Electrical Properties of the Cell Membrane
2.2.1 Ionic Concentrations and Permeabilities
2.3 Ion Transporters
2.3.1 The Sodium-Potassium Pump
2.3.2 Uniporters and Cotransporters
2.4 Origin of the Resting Membrane Voltage
2.4.1 Membrane Voltage in the Steady State
2.5 Membrane Equivalent Circuit
2.5.1 Membrane Conductances
2.5.2 Generation of Electric Signals
2.6 Membrane Rectification
2.7 Membrane Reactance
2.7.1 Inductive Reactance
2.7.2 Capacitive Reactance
2.8 Semiconductor Analogy
Summary of Main Concepts
Chapter 3 Generation of the Action Potential
Objective and Overview
Learning Objectives
3.1 Generation of the Action Potential
3.2 The Hodgkin–Huxley Model
3.2.1 Voltage Clamp Technique and Basic Results
3.2.2 Mathematical Description
3.2.2.1 Potassium Conductance
3.2.2.2 Sodium Conductance
3.2.3 Kinetic Representation of the Hodgkin–Huxley Model
3.2.4 Na+ and K+ Channels
3.3 Properties of the Action Potential under Space Clamp
3.3.1 Active Response
3.3.2 Threshold
3.3.3 Strength-Duration Relationship
3.3.4 Effect of Temperature
3.3.5 Refractoriness
Summary of Main Concepts
Chapter 4 Propagation of the Action Potential
Objective and Overview
Learning Objectives
4.1.1 Cable Model
4.1.2 Solution of the Cable Equation
4.1.2.1 Response to a Current Step
4.1.2.2 Response to a Current Impulse
4.1.2.3 Wave Propagation
4.2.1 Quantitative Considerations
4.3 Properties of the Propagating Action Potential
4.3.1 Threshold
4.3.2 Effect of Temperature
4.3.3 Active vs. Passive Propagation
Summary of Main Concepts
Chapter 5 The Neuromuscular Junction
Objective and Overview
Learning Objectives
5.1 Structure
5.2 Sequence of Events
5.3 Statistics of Neurotransmitter Release
5.3.1 Spontaneous Release
5.3.2 Evoked Release
5.4 The ACh Receptor
5.4.1 Structure
5.4.2 Channel Kinetics
5.4.3 Channel Desensitization
5.5 Generation of the Muscle Action Potential
5.5.1 The Endplate Current
5.5.2 The Endplate Voltage
5.6 Interference with Normal Operation
5.6.1 ACh Agonists and Antagonists
Summary of Main Concepts
Appendix 5A Chapman–Kolmogorov Equation
Chapter 6 Synapses
Objective and Overview
Learning Objectives
6.1 Overview of Synapses
6.1.1 General
6.1.2 Neurotransmitters
6.1.2.1 Types of Neurotransmitters
6.1.2.2 Neurotransmitter Cycle
6.2 Fast Chemical Synapses
6.2.1 General
6.2.2 Fast Inhibitory Synapses
6.2.3 Fast Excitatory Synapses
6.3 Second-Messenger Systems
6.3.1 General Description
6.3.2 Neuromodulators
6.4 Presynaptic Inhibition and Facilitation
6.5 Synaptic Plasticity
6.5.1 Short-Term Synaptic Plasticity
6.5.2 Long-Term Synaptic Plasticity
6.5.2.1 Long-Term Potentiation
6.5.2.2 Long-Term Depression
6.5.3 Structural Changes in Dendritic Spines
6.5.4 Hebbian Synapses
6.6 Electrical Synapses
Summary of Main Concepts
Chapter 7 Neurons
Objective and Overview
Learning Objectives
7.1 Overview of Neurons
7.2 Triggering of Neuronal Spikes
7.2.1 Basic Synaptic Mechanisms
7.2.2 Synaptic Connections between Neurons
7.2.3 Nonsynaptic Mechanisms
7.2.4 Electrically Mediated Mechanisms
7.2.4.1 Gap Junctions
7.2.4.2 Field Potentials
7.3 Neuronal Ion Channels and Currents
7.3.1 Sodium Channels
7.3.2 Calcium Channels
7.3.3 Potassium Channels
7.3.4 Chloride Channels
7.3.5 Effects on Afterhyperpolarization and Afterdepolarization
7.4 Dendritic Responses
7.4.1 Synaptic Integration
7.4.2 Modulation of Synaptic Voltages
7.4.3 Backpropagation
7.4.4 Dendritic Spikes
7.4.5 Bistability in Dendrites
Summary of Main Concepts
Chapter 8 Neuronal Firing Patterns and Models
Objective and Overview
Learning Objectives
8.1 Neuronal Firing Patterns and Their Modulation
8.1.1 Neuronal Computation
8.1.2 Neuronal Excitability
8.1.3 Resonators and Integrators
8.1.4 Neuronal Firing Patterns
8.1.4.1 Regular Spiking Neurons
8.1.4.2 Intrinsically Bursting Neurons
8.1.4.3 Fast Rhythmic Bursting Neurons
8.1.4.4 Fast Spiking Interneurons
8.1.4.5 Low Threshold Spiking Interneurons
8.1.4.6 Late Spiking Interneurons
8.1.5 Rhythmic and Synchronized Firing
8.2 Neuronal Models
8.2.1 Dynamical Neuronal Models
8.2.1.1 Integrate-and-Fire Model
8.2.1.2 Resonate-and-Fire Model
8.2.1.3 Fast-Slow Reduced HH Model
8.2.1.4 Fitzhugh–Nagumo Model
8.2.1.5 Quadratic Model
8.2.1.6 Morris–Lecar Model
8.2.2 Biophysical Neuronal Models
8.2.2.1 Morphoelectrotonic Transformations
8.2.2.2 Compartmental Models
8.3 Models of Neuronal Networks
8.3.1 Compartmental Models
8.3.2 Firing Rate Models
Summary of Main Concepts
Chapter 9 Skeletal Muscle
Objective and Overview
Learning Objectives
9.1 Structure of Skeletal Muscle
9.1.1 Gross Structure
9.1.2 Microstructure
9.2 Contraction of Skeletal Muscle
9.2.1 Excitation-Contraction Coupling
9.2.2 ATP Synthesis
9.2.3 Heat Production
9.2.4 Muscle Fatigue
9.3 Organization of Muscle Fibers
9.3.1 Motor Unit
9.3.2 Muscle Fiber Types
9.3.3 Motoneuron-Muscle Fiber Interactions
9.3.4 Muscle Action
9.3.5 Muscle Architecture
9.4 Muscle Receptors
9.4.1 Golgi Tendon Organ
9.4.2 Muscle Spindle
9.4.2.1 Structure and General Properties
9.4.2.2 Sensory Responses
9.4.2.3 Fusimotor Effects
Summary of Main Concepts
Chapter 10 Functional Properties of Muscle
Objective and Overview
Learning Objectives
10.1 Types of Contraction
10.2 Twitch Contractions
10.2.1 Isometric Twitch
10.2.2 Isotonic Twitch
10.2.3 Summation of Contractions
10.2.4 Gradation of Muscular Contraction
10.3 Mechanics of Contraction
10.3.1 Length-Tension Relation
10.3.2 Force-Velocity Relation
10.3.3 Kinetics of Contraction
10.3.4 Mechanical Model
10.4 Pennate vs. Parallel Muscles
10.5 Cardiac Muscle
10.5.1 Cardiac Cells
10.5.2 Starling’s Law
10.5.3 Cardiac Action Potential
10.5.3.1 Non-Pacemaker Cardiocytes
10.5.3.2 Pacemaker Cardiocytes
10.6 Smooth Muscle
Summary of Main Concepts
Chapter 11 Spinal Cord and Reflexes
Objective and Overview
Learning Objectives
11.1 Gross Features
11.1.1 Vertebral Column
11.1.2 Peripheral Nerves
11.1.3 Neural Organization
11.2 Somatomotor Neurons
11.2.1 Motoneurons
11.2.1.1 General
11.2.1.2 Persistent Inward Current
11.2.1.3 Size Principle
11.2.2 Interneurons
11.2.2.1 Renshaw Cells
11.2.2.2 Ia Interneurons
11.2.2.3 Interneuronal Circuits
11.2.3 Modulatory Effects
11.3 Spinal Reflexes
11.3.1 General
11.3.2 Flexion Reflex
11.3.3 Stretch Reflex
11.3.4 Tendon Organ Reflex
11.3.5 Supraspinal Influences
11.4 Reflexes Elicited by Stimulation
11.4.1 H-Reflex
11.4.2 Tonic Vibration Reflex
Summary of Main Concepts
Chapter 12 Brain Motor Centers and Pathways
Objective and Overview
Learning Objectives
12.1 Hierarchical Organization
12.1.1 Higher Levels
12.2 Middle Hierarchical Level
12.2.1 General
12.2.2 Primary Motor Cortex
12.2.2.1 Descending Pathway
12.2.3 Basal Ganglia
12.2.4 Cerebellum
12.2.4.1 Gross Anatomy
12.2.4.2 Afferent and Efferent Connections
12.2.4.3 Cellular Organization and Features
12.2.4.4 Cerebellar Plasticity
12.2.4.5 Cerebellar Disorders
12.2.4.6 Cerebellar Function
12.2.5 Brainstem Nuclei and Descending Tracts
12.2.5.1 Red Nucleus
12.2.5.2 Reticular Nuclei
12.2.5.3 Vestibular Nuclei
12.2.5.4 Tectospinal Tract
12.2.5.5 C3-C4 Propriospinal System
Summary of Main Concepts
Chapter 13 Control of Movement and Posture
Objective and Overview
Learning Objectives
13.1 Aspects of Movement
13.1.1 Lever Action
13.1.2 Co-contraction of Antagonist Muscles
13.1.3 Feedforward and Feedback Control
13.1.4 Motor Coordination
13.1.5 Motor Equivalence
13.2 Motor Learning and Memory
13.3 Posture
13.3.1 Balance
13.3.2 Upright Posture
13.3.3 Postural Adjustments
13.4 Locomotion
13.4.1 Phases of Gait
13.4.2 Central Pattern Generators
13.4.3 Extra-Spinal Influences
13.5 The Equilibrium Point Hypothesis
13.5.1 General Views of Motor Control
13.5.2 Basics of the EP Hypothesis
13.5.3 Elaboration of the EP Hypothesis
13.5.4 Movement and Posture
13.5.5 Agonist-Antagonist Co-contraction
13.5.6 Motor Redundancy
Summary of Main Concepts
Bibliography and References
Index