Animal Locomotion (C Oabs T Oxford Animal Biology)

This book provides a synthesis of the physical, physiological, evolutionary, and biomechanical principles that underlie animal locomotion. An understanding and full appreciation of animal locomotion requires the integration of these principles. Toward this end, we provide the necessary introductory foundation that will allow a more in-depth understanding of the physical biology and physiology of animal movement. In so doing, we hope that this book will illuminate the fundamentals and breadth of these systems, while inspiring our readers to look more deeply into the scientific literature and investigate new features of animal movement. Several themes run through this book. The first is that by comparing the modes and mechanisms by which animals have evolved the capacity for movement, we can understand the common principles that underlie each mode of locomotion. A second is that size matters. One of the most amazing aspects of biology is the enormous spatial and temporal scale over which organisms and biological processes operate. Within each mode of locomotion, animals have evolved designs and mechanisms that effectively contend with the physical properties and forces imposed on them by their environment. Understanding the constraints of scale that underlie locomotor mechanisms is essential to appreciating how these mechanisms have evolved and how they operate. A third theme is the importance of taking an integrative and comparative evolutionary approach in the study of biology. Organisms share much in common. Much of their molecular and cellular machinery is the same. They also must navigate similar physical properties of their environment. Consequently, an integrative approach to organismal function that spans multiple levels of biological organization provides a strong understanding of animal locomotion. By comparing across species, common principles of design emerge. Such comparisons also highlight how certain organisms may differ and point to strategies that have evolved for movement in diverse environments. Finally, because convergence upon common designs and the generation of new designs result from historical processes governed by natural selection, it is also important that we ask how and why these systems have evolved. Cover 1 Animal Locomotion 4 Copyright 5 Table of Contents 6 Preface 10 List of Variables 12 Chapter 1. Physical and Biological Properties and Principles: Relatd to Animal Locomotion 14 1.1 Environmental media 14 1.1.1 Physical properties of media 14 1.1.2 Impact of physical media on locomotor function 15 1.2 Physics and energetics of movement 15 1.3 Biomechanics of locomotor support 16 1.3.1 Modes of loading 18 1.3.2 Safety factors 19 1.4 Scaling: the importance of size 20 1.4.1 Allometric equation 22 1.5 Dimensions and units 22 1.6 Summary 24 Additional reading 24 Chapter 2. Muscles and Skeletons: The Building Blocks of Animal Movement 25 2.1 Muscles 25 2.2 Molecular organization: mechanism of force generation and shortening 25 2.3 Levels of force generation and the isometric force-length relationship 27 2.4 Power, efficiency and the isotonic force-velocity relationship 29 2.5 “Work loops”: time varying force-length behavior of muscles 31 2.6 Excitation–contraction coupling and motor units 33 2.7 Muscle fiber types 35 2.8 Fiber architecture and its effects on muscle volume and energy use 38 2.9 Skeletons 40 2.10 The connection between muscle and skeleton 40 2.11 Vertebrate endoskeletons 41 2.12 Invertebrate exoskeletons 43 2.13 Hydrostatic skeletons 43 2.14 Skeletons as jointed lever systems 44 2.15 Summary 46 Additional reading 46 Chapter 3. Energetics of Locomotion 47 3.1 Linking cellular metabolism to locomotor energetics 47 3.1.1 Quantifying energy use: respirometry measurements of oxygen consumption or carbon dioxide production 48 3.2 Sources and time course of energy usage during exercise 48 3.2.1 Oxygen deficit, post-exercise oxygen recovery, and steady state metabolism 49 3.2.2 Sustainable activity, maximum aerobic metabolism, and metabolic scope 52 3.3 Endurance and fatigue 53 3.4 Energy costs across terrestrial locomotor speeds 53 3.4.1 Ectothermic versus endothermic energy patterns 54 3.4.2 Kangaroo and wallaby hopping: a remarkable relationship between energy cost and speed 56 3.4.3 Energy costs in the context of gait 57 3.5 Energy cost relative to body size 60 3.5.1 Modeling rates of muscle force development, activation-relaxation, and limb swing to estimate metabolic energy cost 62 3.6 Energy cost of incline running 65 3.7 Cost of swimming 66 3.8 Cost of flight 67 3.9 Locomotion costs compared 69 3.10 Intermittent exercise 71 3.11 Other adaptations for increased aerobic capacity 72 3.12 Summary 72 Additional reading 73 Chapter 4. Movement on Land 74 4.1 Biological wheels: why so few? 74 4.2 Limbs as propulsors: support and swing phases 75 4.3 Limb mechanical advantage and joint torques: interaction of limb posture and ground reaction force 77 4.4 Locomotor gaits 80 4.4.1 Walking 80 4.4.2 Trotting, running and hopping 81 4.4.3 Galloping 81 4.5 Stride frequency and stride length relative to speed and size 82 4.6 Spring-mass properties of running 84 4.6.1 Joint work in relation to steady versus non-steady movement 85 4.7 Maneuverability versus stability 86 4.8 Froude number and dynamic similarity 89 4.9 Inferring gait and speed of fossil animals 90 4.10 Mechanical work: potential and kinetic energy changes during terrestrial locomotion 90 4.10.1 Walking: body and limb movement as “inverted pendula” 91 4.10.2 Running, trotting, hopping and galloping: bouncing gaits 92 4.10.3 “SLIP” limb mechanics 93 4.11 Collisional mechanics of legged locomotion 93 4.12 Legged robotics 95 4.13 Limbless locomotion 95 4.14 Muscle work versus force economy 97 4.15 Tendon springs and muscle dampers 98 4.16 Summary 101 Additional reading 102 Chapter 5. Movement in Water 103 5.1 Thrust and drag 103 5.2 Inertia, viscosity and Reynolds number 104 5.3 Steady flow: drag and streamlines 106 5.3.1 Steady versus unsteady flow 108 5.4 Swimming fish, mammals and cephalopods: movement at high Re 108 5.4.1 Undulatory swimming 109 5.4.2 Caudal fin or fluke swimming 110 5.4.3 Tail shape: homocercal versus heterocercal tails 111 5.4.4 Pectoral, dorsal and anal fin swimming 114 5.5 Jet-based fluid propulsion 116 5.6 Movement at low Re: the reversibility of flow 117 5.6.1 Flagellar swimming 117 5.6.2 Ciliary swimming 120 5.6.3 Size considerations 120 5.7 Movement at intermediate Re: switching between paddles and rakes 121 5.8 Air-water interface: surface swimming, striding and sailing 121 5.8.1 Striding and sailing on the water surface 122 5.8.2 Running on the water surface at large size: integrating terrestrial and aquatic lifestyles 124 5.9 Biological robotics in and on water 125 5.10 Summary 125 Additional reading 126 Chapter 6. Movement in Air 127 6.1 Flight forces: lift, drag and thrust 128 6.1.1 Aspect ratio 131 6.1.2 Wing loading 131 6.2 Power requirements for steady flight 132 6.2.1 Profile and parasite drag 132 6.2.2 Induced drag: the cost of finite wings 134 6.3 Gliding flight 134 6.3.1 Soaring 137 6.4 Flapping flight 138 6.4.1 Kinematics 138 6.4.2 Changes in circulation and wake patterns with flight speed 140 6.4.3 Intermittent flight 142 6.4.4 Origin and evolution of flapping flight 143 6.5 Flight motors and wing anatomy 145 6.5.1 Vertebrate flight musculature 146 6.5.2 Avian pectoralis function: implications for power output during flight 147 6.5.3 Insect flight muscle mechanics 149 6.5.4 Thermal issues related to flight 151 6.6 Flight maneuvering and stability 152 6.6.1 Maneuvering flight 152 6.6.2 Flight stability and control 155 6.7 Unsteady aerodynamic mechanisms 156 6.8 Summary 159 Additional reading 159 Chapter 7. Jumping, Climbing and Suspensory Locomotion 160 7.1 Jumping 160 7.2 Jump take-offs and trajectories 161 7.3 Scaling of jumps 162 7.3.1 The role of muscle in jump scaling 162 7.3.2 Body weight and jump take-off 163 7.3.3 Limb length and jump scaling 163 7.4 Power enhancements to jump performance 165 7.4.1 Counter-movement jumping 166 7.4.2 Power amplification via rapid release of stored elastic energy 166 7.4.3 Extreme power amplification with springs and latches 168 7.5 Interactions with the substrate during jumping 169 7.6 Climbing and attachment mechanisms 171 7.6.1 Navigating branches 171 7.6.2 Static frictional gripping and claws 172 7.6.3 Locomoting with adhesion and friction 173 7.7 Suspensory locomotion 175 7.8 Inspiration for synthetic systems 176 7.9 Summary 176 Additional reading 177 Chapter 8. Neuromuscular Control of Movement 178 8.1 Sensory elements 178 8.1.1 Vertebrate sensory organs 178 Muscle spindles 179 Golgi tendon organs 181 8.1.2 Insect sensory organs 182 8.2 Sensorimotor integration via local reflex pathways 182 8.2.1 Vertebrate reflex pathways 183 8.2.2 Insect reflex pathways 185 8.3 Muscle recruitment in relation to functional demand: force, speed and endurance 187 8.3.1 Vertebrate motor recruitment 187 Orderly recruitment: the “size principle” 188 Motor unit distribution 189 Muscle synergies and global task control 190 8.3.2 Invertebrate motor recruitment 190 Invertebrate muscle fiber types 191 Examples of neuromotor organization in the locust and cockroach 192 Invertebrate muscle activation patterns in relation to speed 194 8.4 Reciprocal inhibition: a basic feature of sensorimotor neural circuits 195 8.5 Distributed control: the role of central pattern generators 196 8.6 Case examples of motor control 198 8.6.1 Mechanosensory and visual control of fly flight 199 8.6.2 Fish swimming: motor recruitment in variable temperatures 200 8.7 Summary 200 Additional reading 202 Chapter 9. Evolution of Locomotion 203 9.1 Large-scale trends in animal locomotion 203 9.1.1 Origins of flight 206 9.1.2 Evolution of legged terrestrial locomotion 208 9.2 From genes to locomotion 210 9.3 Comparative methods and animal locomotion 211 9.4 The relevance of evolution to robotics and bio-inspired design 213 9.5 Summary 215 Additional reading 216 Reference 218 Index 232 Animals Have Evolved Remarkable Biomechanical And Physiological Systems That Enable Their Rich Repertoire Of Motion. Animal Locomotion Offers A Fundamental Understanding Of Animal Motion Through A Broad Comparative And Integrative Approach, Including Basic Mathematics And Physics, Examination Of New And Enduring Literature, Consideration Of Classic And Cutting-edge Methods, And A Strong Emphasis On The Core Concepts That Consistently Ground The Dizzying Array Of Animal Movements. Across Scales And Environments, This Book Integrates The Biomechanics Of Animal Movement With The Physiology Of Animal Energetics And The Neural Control Of Locomotion. This Second Edition Has Been Thoroughly Revised, Incorporating New Content On Non-vertebrate Animal Locomotor Systems, Studies On Animal Locomotion That Have Inspired Robotic Designs, And A New Chapter On The Use Of Evolutionary Approaches In Locomotor Mechanisms And Performance. Animal Locomotion Is Suitable For Undergraduate And Graduate Level Students Taking Related Courses In Animal Physiology And Movement As Well As Researchers In The Fields Of Bio-inspired Robotics, Biomechanics, Physiology, And Evolutionary Biology. Physical And Biological Properties And Principles: Related To Animal Locomotion -- Muscles And Skeletons: The Building Blocks Of Animal Movement -- Energetics Of Locomotion -- Movement On Land -- Movement In Water -- Movement In Air -- Jumping, Climbing, And Suspensory Locomotion -- Neuromuscular Control Of Movement -- Evolution Of Locomotion. Andrew A. Biewener (charles P. Lyman Professor Of Biology, Director, Concord Field Station, Harvard University), Sheila N. Patek (associate Professor Of Biology, Duke University). First Edition Published In 2003--title Page Verso. Includes Bibliographical References (pages 205-217) And Index. « Animals have evolved remarkable biomechanical and physiological systems that enable their rich repertoire of motion. Animal Locomotion offers a fundamental understanding of animal movement through a broad comparative and integrative approach, including basic mathematics and physics, examination of new and enduring literature, consideration of classic and cutting-edge methods, and a strong emphasis on the core concepts that consistently ground the dizzying array of animal movements. Across scales and environments, this book integrates the biomechanics of animal movement with the physiology of animal energetics and the neural control of locomotion. This second edition has been thoroughly revised, incorporating new content on non-vertebrate animal locomotor systems, studies of animal locomotion that have inspired robotic designs, and a new chapter on the use of evolutionary approaches to locomotor mechanisms and performance. »-- Résumé de l'éditeur