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Understanding Mammalian Locomotion will formally introduce the emerging perspective of collision dynamics in mammalian terrestrial locomotion and explain how it influences the interpretation of form and functional capabilities. The objective is to bring the reader interested in the function and mechanics of mammalian terrestrial locomotion to a sophisticated conceptual understanding of the relevant mechanics and the current debate ongoing in the field.
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Understanding Mammalian Locomotion will formally introduce the emerging perspective of collision dynamics in mammalian terrestrial locomotion and explain how it influences the interpretation of form and functional capabilities. The objective is to bring the reader interested in the function and mechanics of mammalian terrestrial locomotion to a sophisticated conceptual understanding of the relevant mechanics and the current debate ongoing in the field.
Produktdetails
- Produktdetails
- Verlag: John Wiley & Sons / Wiley
- Seitenzahl: 432
- Erscheinungstermin: 19. Januar 2016
- Englisch
- Abmessung: 261mm x 179mm x 27mm
- Gewicht: 848g
- ISBN-13: 9780470454640
- ISBN-10: 0470454644
- Artikelnr.: 34439397
- Verlag: John Wiley & Sons / Wiley
- Seitenzahl: 432
- Erscheinungstermin: 19. Januar 2016
- Englisch
- Abmessung: 261mm x 179mm x 27mm
- Gewicht: 848g
- ISBN-13: 9780470454640
- ISBN-10: 0470454644
- Artikelnr.: 34439397
John E.A. Bertram is a Professor in the Department of Cell Biology and Anatomy, Cumming School of Medicine, and adjunct Professor in the Department of Comparative Biology and Experimental Medicine, Faculty of Veterinary Medicine, at the University of Calgary in Calgary, AB, Canada
List of Contributors xv Preface xvii Chapter 1 Concepts Through Time:
Historical Perspectives on Mammalian Locomotion 1 John E. A. Bertram 1.1
Introduction 1 1.2 The ancients and the contemplation of motion 2 1.3 The
European Renaissance and foundations of the age of discovery 3 1.4 The era
of technological observation 5 1.5 Physiology and mechanics of terrestrial
locomotion - cost and consequences 7 1.6 Comparative studies of gait 10 1.6
Re?]interpreting the mechanics: a fork in the road, or simply seeing the
other side of the coin? 13 1.7 The biological source of cost 13 1.8 The
physical source of cost (with biological consequences) - the road less
traveled 14 1.9 Conclusions 21 References 21 Chapter 2 Considering Gaits:
Descriptive Approaches 27 John E. A. Bertram 2.1 Introduction 27 2.2
Defining the fundamental gaits 28 2.3 Classifying and comparing the
fundamental gaits 30 2.4 Symmetric gaits 32 2.5 A symmetric gaits 34 2.6
Beyond "Hildebrand plots" 40 2.7 Statistical classification 43 2.8 Neural
regulation and emergent criteria 45 2.9 Mechanical measures as descriptions
of gaits 47 2.10 Conclusion 47 References 48 Chapter 3 Muscles as Actuators
51 Anne K. Gutmann and John E. A. Bertram 3.1 Introduction 51 3.2 Basic
muscle operation 52 3.2.1 Sliding filament theory - the basis for
cross?]bridge theory 52 3.2.2 Basic cross?]bridge theory 52 3.2.3
Multi?]state cross?]bridge models 57 3.3 Some alternatives to cross?]bridge
theory 59 3.4 Force production 60 3.4.1 Isometric force production 60 3.4.2
Non?]isometric force production 63 3.5 The Hill?]type model 66 3.6
Optimizing work, power, and efficiency 68 3.7 Muscle architecture 70 3.7.1
The sarcomere as the fundamental contractile unit 70 3.7.2 Muscle geometry
70 3.7.3 Elastic energy storage and return 72 3.7.4 Damping/energy
dissipation 72 3.8 Other factors that influence muscle performance 73 3.8.1
Fiber type 73 3.9 A ctivation and recruitment 75 3.10 What does muscle do
best? 76 References 76 Chapter 4 Concepts in Locomotion: Levers, Struts,
Pendula and Springs 79 John E. A. Bertram 4.1 Introduction 79 4.2 The limb:
How details can obscure functional role 83 4.3 Limb function in stability
and the concept of the "effective limb" 85 4.3.1 Considering the mechanisms
of stability 85 4.3.2 The role of the effective limb 88 4.4 Levers and
struts 89 4.5 Ground reaction force in gaits 92 4.5.1 Trot 94 4.5.2 Walk 96
4.5.3 Gallop 97 4.6 The consequence of applied force: CoM motion, pendula
and springs 98 4.7 Energy exchange in locomotion - valuable or inevitable?
102 4.8 Momentum and energy in locomotion: dynamic fundamentals 103 4.9
Energy - lost unless recovered, or available unless lost? 104 References
105 Chapter 5 Concepts in Locomotion: Wheels, Spokes, Collisions and
Insight from the Center of Mass 111 John E. A. Bertram 5.1 Introduction 111
5.2 Understanding brachiation: an analogy for terrestrial locomotion 112
5.3 Bipedal walking: inverted pendulum or inverted "collision?]limiting
brachiator analog"? 117 5.4 Basic dynamics of the step?]to?]step transition
in bipedal walking 120 5.5 Subtle dynamics of the step?]to?]step transition
in bipedal walking and running 124 5.6 Pseudo?]elastic motion and true
elastic return in running gaits 130 5.7 Managing CoM motion in quadrupedal
gaits 131 5.7.1 Walk 132 5.7.2 Trot 133 5.7.3 Gallop 133 5.8 Conclusion 138
References 139 Chapter 6 Reductionist Models of Walking and Running 143
James R. Usherwood 6.1 Part 1: Bipedal locomotion and "the ultimate cost of
legged locomotion?" 143 6.1.1 Introduction 143 6.1.2 Reductionist models of
walking 144 6.1.3 The benefit of considering locomotion as inelastic 150
6.2 Part 2: quadrupedal locomotion 158 6.2.1 Introduction 158 6.2.2
Quadrupedal dynamic walking and collisions 158 6.2.3 Higher speed
quadrupedal gaits 161 6.2.4 Further success of reductionist mechanics 162
Appendix A: Analytical approximation for costs of transport including legs
and "guts and gonads" losses 166 6A.1 List of symbols 166 6A.2 Period
definitions for a symmetrically running biped 166 6A.3 Ideal work for the
leg 167 6A.4 Vertical work calculations for leg 168 6A.5 Horizontal work
calculations for leg 169 6A.6 Hysteresis costs of "guts and gonads"
deflections 169 6A.7 Cost of transport 170 References 170 Chapter 7
Whole?]Body Mechanics: How Leg Compliance Shapes the Way We Move 173 Andre
Seyfarth, Hartmut Geyer, Susanne Lipfert, J. Rummel, Yvonne Blum, M. Maus
and D. Maykranz 7.1 Introduction 173 7.2 Jumping for distance - a
goal?]directed movement 175 7.3 Running for distance - what is the goal?
177 7.4 Cyclic stability in running 178 7.5 The wheel in the leg - how leg
retraction enhances running stability 179 7.6 Walking with compliant legs
180 7.7 A dding an elastically coupled foot to the spring?]mass model 184
7.8 The segmented leg - how does joint function translate into leg
function? 185 7.9 Keeping the trunk upright during locomotion 187 7.10 The
challenge of setting up more complex models 188 Notes 190 References190
Chapter 8 The Most Important Feature of an Organism's Biology: Dimension,
Similarity and Scale 193 John E. A. Bertram 8.1 Introduction 193 8.2 The
most basic principle: surface area to volume relations 194 8.3 A ssessing
scale effects 197 8.4 Physiology and scaling 198 8.5 The allometric
equation: the power function of scaling 203 8.6 The standard scaling models
207 8.6.1 Geometric similarity 208 8.6.2 Static stress similarity 209 8.6.3
Elastic similarity 209 8.7 Differential scaling - where the limit may
change 210 8.7.1 A ssessing the assumptions 215 8.8 A fractal view of
scaling 215 8.9 Making valid comparisons: measurement, dimension and
functional criteria 217 8.9.1 Considering units 217 8.9.2 Fundamental and
derived units 219 8.9.3 Froude number: a dimensionless example 222
References 223 Chapter 9 Accounting for the Influence of Animal Size on
Biomechanical Variables: Concepts and Considerations 229 Sharon Bullimore
9.1 Introduction 229 9.2 Commonly used approaches to accounting for size
differences 230 9.2.1 Dividing by body mass 230 9.2.2 Dimensionless
parameters 232 9.3 Empirical scaling relationships 237 9.4 Selected
biomechanical parameters 238 9.4.1 Ground reaction force 238 9.4.2 Muscle
force 239 9.4.3 Muscle velocity 242 9.4.4 Running speed 242 9.4.5 Jump
height 244 9.4.6 Elastic energy storage 246 9.5 Conclusions 247
Acknowledgements 247 References 247 Chapter 10 Locomotion in Small
Tetrapods: Size?]Based Limitations to "Universal Rules" in Locomotion 251
Audrone R. Biknevicius, Stephen M. Reilly and Elvedin Kljuno 10.1
Introduction 251 10.2 A ctive mechanisms contributing to the high cost of
transport in small tetrapods 254 10.3 Limited passive mechanisms for
reducing cost of transport in small tetrapods 255 10.4 Gait transitions
from vaulting to bouncing mechanics 257 10.5 The "unsteadiness" of most
terrestrial locomotion 262 Appendix - a model of non?]steady speed walking
265 10A.1 Spring?]mass inverted pendulum model of walking 265 10A.2
Recovery ratio calculation 269 References 271 Chapter 11 Non?]Steady
Locomotion 277 Monica A. Daley 11.1 Introduction 277 11.1.1 Why study
non?]steady locomotion? 278 11.2 A pproaches to studying non?]steady
locomotion 279 11.2.1 Simple mechanical models 280 11.2.2 Research
approaches to non?]steady locomotion 281 11.3 Themes from recent studies of
non?]steady locomotion 282 11.3.1 Limits to maximal acceleration 282 11.3.2
Morphological and behavioral factors in turning mechanics 283 11.4 The role
of intrinsic mechanics for stability and robustness of locomotion 288
11.4.1 Some definitions 289 11.4.2 Measures of sensitivity and robustness
290 11.4.3 What do we learn about stability from simple models of running?
291 11.4.4 Limitations to stability analysis of simple models 295 11.4.5
The relationship between ground contact conditions and leg mechanics on
uneven terrain 296 11.4.6 Compromises among economy, robustness and injury
avoidance in uneven terrain 298 11.5 Proximal?]distal inter?]joint
coordination in non?]steady locomotion 299 References 302 Chapter 12 The
Evolution of Terrestrial Locomotion in Bats: the Bad, the Ugly, and the
Good 307 Daniel K. Riskin, John E. A. Bertram and John W. Hermanson 12.1
Bats on the ground: like fish out of water? 307 12.2 Species?]level
variation in walking ability 308 12.3 How does anatomy influence crawling
ability? 309 12.4 Hindlimbs and the evolution of flight 311 12.5 Moving a
bat's body on land: the kinematics of quadrupedal locomotion 315 12.6
Evolutionary pressures leading to capable terrestrial locomotion 318 12.7
Conclusions and future work 319 Acknowledgements 320 References 320 Chapter
13 The Fight or Flight Dichotomy: Functional Trade?]Off in Specialization
for Aggression Versus Locomotion 325 David R. Carrier 13.1 Introduction325
13.1.1 Why fighting is important 327 13.1.2 Size sexual dimorphism as an
indicator of male?]male aggression 328 13.2 Trade?]offs in specialization
for aggression versus locomotion 329 13.2.1 The evolution of short legs -
specialization for aggression? 329 13.2.2 Muscle architecture of limbs
specialized for running versus fighting 331 13.2.3 Mechanical properties of
limb bones that are specialized for running versus fighting 334 13.2.4 The
function of foot posture: aggression versus locomotor economy 334 13.3
Discussion 338 References 341 Chapter 14 Design for Prodigious Size without
Extreme Body Mass: Dwarf Elephants, Differential Scaling and Implications
for Functional Adaptation 349 John E. A. Bertram 14.1 Introduction 349 14.2
Elephant form, mammalian scaling and dwarfing 351 14.2.1 Measurements 356
14.2.2 Observations 356 14.3 Interpretation 357 Acknowledgements 364
References 364 Chapter 15 Basic Mechanisms of Bipedal Locomotion:
Head?]Supported Loads and Strategies to Reduce the Cost of Walking 369
James R. Usherwood and John E. A. Bertram 15.1 Introduction 369 15.2
Head?]supported loads in human?]mediated transport 370 15.2.1 Can the
evidence be depended upon? 371 15.3 Potential energy saving advantages 373
15.4 A simple alternative model 376 15.5 Conclusions 382 References 382
Chapter 16 Would a Horse on the Moon Gallop? Directions Available in
Locomotion Research (and How Not to Spend Too Much Time Exploring Blind
Alleys) 385 John E. A. Bertram 16.1 Introduction 385 References 392 Index
393
Historical Perspectives on Mammalian Locomotion 1 John E. A. Bertram 1.1
Introduction 1 1.2 The ancients and the contemplation of motion 2 1.3 The
European Renaissance and foundations of the age of discovery 3 1.4 The era
of technological observation 5 1.5 Physiology and mechanics of terrestrial
locomotion - cost and consequences 7 1.6 Comparative studies of gait 10 1.6
Re?]interpreting the mechanics: a fork in the road, or simply seeing the
other side of the coin? 13 1.7 The biological source of cost 13 1.8 The
physical source of cost (with biological consequences) - the road less
traveled 14 1.9 Conclusions 21 References 21 Chapter 2 Considering Gaits:
Descriptive Approaches 27 John E. A. Bertram 2.1 Introduction 27 2.2
Defining the fundamental gaits 28 2.3 Classifying and comparing the
fundamental gaits 30 2.4 Symmetric gaits 32 2.5 A symmetric gaits 34 2.6
Beyond "Hildebrand plots" 40 2.7 Statistical classification 43 2.8 Neural
regulation and emergent criteria 45 2.9 Mechanical measures as descriptions
of gaits 47 2.10 Conclusion 47 References 48 Chapter 3 Muscles as Actuators
51 Anne K. Gutmann and John E. A. Bertram 3.1 Introduction 51 3.2 Basic
muscle operation 52 3.2.1 Sliding filament theory - the basis for
cross?]bridge theory 52 3.2.2 Basic cross?]bridge theory 52 3.2.3
Multi?]state cross?]bridge models 57 3.3 Some alternatives to cross?]bridge
theory 59 3.4 Force production 60 3.4.1 Isometric force production 60 3.4.2
Non?]isometric force production 63 3.5 The Hill?]type model 66 3.6
Optimizing work, power, and efficiency 68 3.7 Muscle architecture 70 3.7.1
The sarcomere as the fundamental contractile unit 70 3.7.2 Muscle geometry
70 3.7.3 Elastic energy storage and return 72 3.7.4 Damping/energy
dissipation 72 3.8 Other factors that influence muscle performance 73 3.8.1
Fiber type 73 3.9 A ctivation and recruitment 75 3.10 What does muscle do
best? 76 References 76 Chapter 4 Concepts in Locomotion: Levers, Struts,
Pendula and Springs 79 John E. A. Bertram 4.1 Introduction 79 4.2 The limb:
How details can obscure functional role 83 4.3 Limb function in stability
and the concept of the "effective limb" 85 4.3.1 Considering the mechanisms
of stability 85 4.3.2 The role of the effective limb 88 4.4 Levers and
struts 89 4.5 Ground reaction force in gaits 92 4.5.1 Trot 94 4.5.2 Walk 96
4.5.3 Gallop 97 4.6 The consequence of applied force: CoM motion, pendula
and springs 98 4.7 Energy exchange in locomotion - valuable or inevitable?
102 4.8 Momentum and energy in locomotion: dynamic fundamentals 103 4.9
Energy - lost unless recovered, or available unless lost? 104 References
105 Chapter 5 Concepts in Locomotion: Wheels, Spokes, Collisions and
Insight from the Center of Mass 111 John E. A. Bertram 5.1 Introduction 111
5.2 Understanding brachiation: an analogy for terrestrial locomotion 112
5.3 Bipedal walking: inverted pendulum or inverted "collision?]limiting
brachiator analog"? 117 5.4 Basic dynamics of the step?]to?]step transition
in bipedal walking 120 5.5 Subtle dynamics of the step?]to?]step transition
in bipedal walking and running 124 5.6 Pseudo?]elastic motion and true
elastic return in running gaits 130 5.7 Managing CoM motion in quadrupedal
gaits 131 5.7.1 Walk 132 5.7.2 Trot 133 5.7.3 Gallop 133 5.8 Conclusion 138
References 139 Chapter 6 Reductionist Models of Walking and Running 143
James R. Usherwood 6.1 Part 1: Bipedal locomotion and "the ultimate cost of
legged locomotion?" 143 6.1.1 Introduction 143 6.1.2 Reductionist models of
walking 144 6.1.3 The benefit of considering locomotion as inelastic 150
6.2 Part 2: quadrupedal locomotion 158 6.2.1 Introduction 158 6.2.2
Quadrupedal dynamic walking and collisions 158 6.2.3 Higher speed
quadrupedal gaits 161 6.2.4 Further success of reductionist mechanics 162
Appendix A: Analytical approximation for costs of transport including legs
and "guts and gonads" losses 166 6A.1 List of symbols 166 6A.2 Period
definitions for a symmetrically running biped 166 6A.3 Ideal work for the
leg 167 6A.4 Vertical work calculations for leg 168 6A.5 Horizontal work
calculations for leg 169 6A.6 Hysteresis costs of "guts and gonads"
deflections 169 6A.7 Cost of transport 170 References 170 Chapter 7
Whole?]Body Mechanics: How Leg Compliance Shapes the Way We Move 173 Andre
Seyfarth, Hartmut Geyer, Susanne Lipfert, J. Rummel, Yvonne Blum, M. Maus
and D. Maykranz 7.1 Introduction 173 7.2 Jumping for distance - a
goal?]directed movement 175 7.3 Running for distance - what is the goal?
177 7.4 Cyclic stability in running 178 7.5 The wheel in the leg - how leg
retraction enhances running stability 179 7.6 Walking with compliant legs
180 7.7 A dding an elastically coupled foot to the spring?]mass model 184
7.8 The segmented leg - how does joint function translate into leg
function? 185 7.9 Keeping the trunk upright during locomotion 187 7.10 The
challenge of setting up more complex models 188 Notes 190 References190
Chapter 8 The Most Important Feature of an Organism's Biology: Dimension,
Similarity and Scale 193 John E. A. Bertram 8.1 Introduction 193 8.2 The
most basic principle: surface area to volume relations 194 8.3 A ssessing
scale effects 197 8.4 Physiology and scaling 198 8.5 The allometric
equation: the power function of scaling 203 8.6 The standard scaling models
207 8.6.1 Geometric similarity 208 8.6.2 Static stress similarity 209 8.6.3
Elastic similarity 209 8.7 Differential scaling - where the limit may
change 210 8.7.1 A ssessing the assumptions 215 8.8 A fractal view of
scaling 215 8.9 Making valid comparisons: measurement, dimension and
functional criteria 217 8.9.1 Considering units 217 8.9.2 Fundamental and
derived units 219 8.9.3 Froude number: a dimensionless example 222
References 223 Chapter 9 Accounting for the Influence of Animal Size on
Biomechanical Variables: Concepts and Considerations 229 Sharon Bullimore
9.1 Introduction 229 9.2 Commonly used approaches to accounting for size
differences 230 9.2.1 Dividing by body mass 230 9.2.2 Dimensionless
parameters 232 9.3 Empirical scaling relationships 237 9.4 Selected
biomechanical parameters 238 9.4.1 Ground reaction force 238 9.4.2 Muscle
force 239 9.4.3 Muscle velocity 242 9.4.4 Running speed 242 9.4.5 Jump
height 244 9.4.6 Elastic energy storage 246 9.5 Conclusions 247
Acknowledgements 247 References 247 Chapter 10 Locomotion in Small
Tetrapods: Size?]Based Limitations to "Universal Rules" in Locomotion 251
Audrone R. Biknevicius, Stephen M. Reilly and Elvedin Kljuno 10.1
Introduction 251 10.2 A ctive mechanisms contributing to the high cost of
transport in small tetrapods 254 10.3 Limited passive mechanisms for
reducing cost of transport in small tetrapods 255 10.4 Gait transitions
from vaulting to bouncing mechanics 257 10.5 The "unsteadiness" of most
terrestrial locomotion 262 Appendix - a model of non?]steady speed walking
265 10A.1 Spring?]mass inverted pendulum model of walking 265 10A.2
Recovery ratio calculation 269 References 271 Chapter 11 Non?]Steady
Locomotion 277 Monica A. Daley 11.1 Introduction 277 11.1.1 Why study
non?]steady locomotion? 278 11.2 A pproaches to studying non?]steady
locomotion 279 11.2.1 Simple mechanical models 280 11.2.2 Research
approaches to non?]steady locomotion 281 11.3 Themes from recent studies of
non?]steady locomotion 282 11.3.1 Limits to maximal acceleration 282 11.3.2
Morphological and behavioral factors in turning mechanics 283 11.4 The role
of intrinsic mechanics for stability and robustness of locomotion 288
11.4.1 Some definitions 289 11.4.2 Measures of sensitivity and robustness
290 11.4.3 What do we learn about stability from simple models of running?
291 11.4.4 Limitations to stability analysis of simple models 295 11.4.5
The relationship between ground contact conditions and leg mechanics on
uneven terrain 296 11.4.6 Compromises among economy, robustness and injury
avoidance in uneven terrain 298 11.5 Proximal?]distal inter?]joint
coordination in non?]steady locomotion 299 References 302 Chapter 12 The
Evolution of Terrestrial Locomotion in Bats: the Bad, the Ugly, and the
Good 307 Daniel K. Riskin, John E. A. Bertram and John W. Hermanson 12.1
Bats on the ground: like fish out of water? 307 12.2 Species?]level
variation in walking ability 308 12.3 How does anatomy influence crawling
ability? 309 12.4 Hindlimbs and the evolution of flight 311 12.5 Moving a
bat's body on land: the kinematics of quadrupedal locomotion 315 12.6
Evolutionary pressures leading to capable terrestrial locomotion 318 12.7
Conclusions and future work 319 Acknowledgements 320 References 320 Chapter
13 The Fight or Flight Dichotomy: Functional Trade?]Off in Specialization
for Aggression Versus Locomotion 325 David R. Carrier 13.1 Introduction325
13.1.1 Why fighting is important 327 13.1.2 Size sexual dimorphism as an
indicator of male?]male aggression 328 13.2 Trade?]offs in specialization
for aggression versus locomotion 329 13.2.1 The evolution of short legs -
specialization for aggression? 329 13.2.2 Muscle architecture of limbs
specialized for running versus fighting 331 13.2.3 Mechanical properties of
limb bones that are specialized for running versus fighting 334 13.2.4 The
function of foot posture: aggression versus locomotor economy 334 13.3
Discussion 338 References 341 Chapter 14 Design for Prodigious Size without
Extreme Body Mass: Dwarf Elephants, Differential Scaling and Implications
for Functional Adaptation 349 John E. A. Bertram 14.1 Introduction 349 14.2
Elephant form, mammalian scaling and dwarfing 351 14.2.1 Measurements 356
14.2.2 Observations 356 14.3 Interpretation 357 Acknowledgements 364
References 364 Chapter 15 Basic Mechanisms of Bipedal Locomotion:
Head?]Supported Loads and Strategies to Reduce the Cost of Walking 369
James R. Usherwood and John E. A. Bertram 15.1 Introduction 369 15.2
Head?]supported loads in human?]mediated transport 370 15.2.1 Can the
evidence be depended upon? 371 15.3 Potential energy saving advantages 373
15.4 A simple alternative model 376 15.5 Conclusions 382 References 382
Chapter 16 Would a Horse on the Moon Gallop? Directions Available in
Locomotion Research (and How Not to Spend Too Much Time Exploring Blind
Alleys) 385 John E. A. Bertram 16.1 Introduction 385 References 392 Index
393
List of Contributors xv Preface xvii Chapter 1 Concepts Through Time:
Historical Perspectives on Mammalian Locomotion 1 John E. A. Bertram 1.1
Introduction 1 1.2 The ancients and the contemplation of motion 2 1.3 The
European Renaissance and foundations of the age of discovery 3 1.4 The era
of technological observation 5 1.5 Physiology and mechanics of terrestrial
locomotion - cost and consequences 7 1.6 Comparative studies of gait 10 1.6
Re?]interpreting the mechanics: a fork in the road, or simply seeing the
other side of the coin? 13 1.7 The biological source of cost 13 1.8 The
physical source of cost (with biological consequences) - the road less
traveled 14 1.9 Conclusions 21 References 21 Chapter 2 Considering Gaits:
Descriptive Approaches 27 John E. A. Bertram 2.1 Introduction 27 2.2
Defining the fundamental gaits 28 2.3 Classifying and comparing the
fundamental gaits 30 2.4 Symmetric gaits 32 2.5 A symmetric gaits 34 2.6
Beyond "Hildebrand plots" 40 2.7 Statistical classification 43 2.8 Neural
regulation and emergent criteria 45 2.9 Mechanical measures as descriptions
of gaits 47 2.10 Conclusion 47 References 48 Chapter 3 Muscles as Actuators
51 Anne K. Gutmann and John E. A. Bertram 3.1 Introduction 51 3.2 Basic
muscle operation 52 3.2.1 Sliding filament theory - the basis for
cross?]bridge theory 52 3.2.2 Basic cross?]bridge theory 52 3.2.3
Multi?]state cross?]bridge models 57 3.3 Some alternatives to cross?]bridge
theory 59 3.4 Force production 60 3.4.1 Isometric force production 60 3.4.2
Non?]isometric force production 63 3.5 The Hill?]type model 66 3.6
Optimizing work, power, and efficiency 68 3.7 Muscle architecture 70 3.7.1
The sarcomere as the fundamental contractile unit 70 3.7.2 Muscle geometry
70 3.7.3 Elastic energy storage and return 72 3.7.4 Damping/energy
dissipation 72 3.8 Other factors that influence muscle performance 73 3.8.1
Fiber type 73 3.9 A ctivation and recruitment 75 3.10 What does muscle do
best? 76 References 76 Chapter 4 Concepts in Locomotion: Levers, Struts,
Pendula and Springs 79 John E. A. Bertram 4.1 Introduction 79 4.2 The limb:
How details can obscure functional role 83 4.3 Limb function in stability
and the concept of the "effective limb" 85 4.3.1 Considering the mechanisms
of stability 85 4.3.2 The role of the effective limb 88 4.4 Levers and
struts 89 4.5 Ground reaction force in gaits 92 4.5.1 Trot 94 4.5.2 Walk 96
4.5.3 Gallop 97 4.6 The consequence of applied force: CoM motion, pendula
and springs 98 4.7 Energy exchange in locomotion - valuable or inevitable?
102 4.8 Momentum and energy in locomotion: dynamic fundamentals 103 4.9
Energy - lost unless recovered, or available unless lost? 104 References
105 Chapter 5 Concepts in Locomotion: Wheels, Spokes, Collisions and
Insight from the Center of Mass 111 John E. A. Bertram 5.1 Introduction 111
5.2 Understanding brachiation: an analogy for terrestrial locomotion 112
5.3 Bipedal walking: inverted pendulum or inverted "collision?]limiting
brachiator analog"? 117 5.4 Basic dynamics of the step?]to?]step transition
in bipedal walking 120 5.5 Subtle dynamics of the step?]to?]step transition
in bipedal walking and running 124 5.6 Pseudo?]elastic motion and true
elastic return in running gaits 130 5.7 Managing CoM motion in quadrupedal
gaits 131 5.7.1 Walk 132 5.7.2 Trot 133 5.7.3 Gallop 133 5.8 Conclusion 138
References 139 Chapter 6 Reductionist Models of Walking and Running 143
James R. Usherwood 6.1 Part 1: Bipedal locomotion and "the ultimate cost of
legged locomotion?" 143 6.1.1 Introduction 143 6.1.2 Reductionist models of
walking 144 6.1.3 The benefit of considering locomotion as inelastic 150
6.2 Part 2: quadrupedal locomotion 158 6.2.1 Introduction 158 6.2.2
Quadrupedal dynamic walking and collisions 158 6.2.3 Higher speed
quadrupedal gaits 161 6.2.4 Further success of reductionist mechanics 162
Appendix A: Analytical approximation for costs of transport including legs
and "guts and gonads" losses 166 6A.1 List of symbols 166 6A.2 Period
definitions for a symmetrically running biped 166 6A.3 Ideal work for the
leg 167 6A.4 Vertical work calculations for leg 168 6A.5 Horizontal work
calculations for leg 169 6A.6 Hysteresis costs of "guts and gonads"
deflections 169 6A.7 Cost of transport 170 References 170 Chapter 7
Whole?]Body Mechanics: How Leg Compliance Shapes the Way We Move 173 Andre
Seyfarth, Hartmut Geyer, Susanne Lipfert, J. Rummel, Yvonne Blum, M. Maus
and D. Maykranz 7.1 Introduction 173 7.2 Jumping for distance - a
goal?]directed movement 175 7.3 Running for distance - what is the goal?
177 7.4 Cyclic stability in running 178 7.5 The wheel in the leg - how leg
retraction enhances running stability 179 7.6 Walking with compliant legs
180 7.7 A dding an elastically coupled foot to the spring?]mass model 184
7.8 The segmented leg - how does joint function translate into leg
function? 185 7.9 Keeping the trunk upright during locomotion 187 7.10 The
challenge of setting up more complex models 188 Notes 190 References190
Chapter 8 The Most Important Feature of an Organism's Biology: Dimension,
Similarity and Scale 193 John E. A. Bertram 8.1 Introduction 193 8.2 The
most basic principle: surface area to volume relations 194 8.3 A ssessing
scale effects 197 8.4 Physiology and scaling 198 8.5 The allometric
equation: the power function of scaling 203 8.6 The standard scaling models
207 8.6.1 Geometric similarity 208 8.6.2 Static stress similarity 209 8.6.3
Elastic similarity 209 8.7 Differential scaling - where the limit may
change 210 8.7.1 A ssessing the assumptions 215 8.8 A fractal view of
scaling 215 8.9 Making valid comparisons: measurement, dimension and
functional criteria 217 8.9.1 Considering units 217 8.9.2 Fundamental and
derived units 219 8.9.3 Froude number: a dimensionless example 222
References 223 Chapter 9 Accounting for the Influence of Animal Size on
Biomechanical Variables: Concepts and Considerations 229 Sharon Bullimore
9.1 Introduction 229 9.2 Commonly used approaches to accounting for size
differences 230 9.2.1 Dividing by body mass 230 9.2.2 Dimensionless
parameters 232 9.3 Empirical scaling relationships 237 9.4 Selected
biomechanical parameters 238 9.4.1 Ground reaction force 238 9.4.2 Muscle
force 239 9.4.3 Muscle velocity 242 9.4.4 Running speed 242 9.4.5 Jump
height 244 9.4.6 Elastic energy storage 246 9.5 Conclusions 247
Acknowledgements 247 References 247 Chapter 10 Locomotion in Small
Tetrapods: Size?]Based Limitations to "Universal Rules" in Locomotion 251
Audrone R. Biknevicius, Stephen M. Reilly and Elvedin Kljuno 10.1
Introduction 251 10.2 A ctive mechanisms contributing to the high cost of
transport in small tetrapods 254 10.3 Limited passive mechanisms for
reducing cost of transport in small tetrapods 255 10.4 Gait transitions
from vaulting to bouncing mechanics 257 10.5 The "unsteadiness" of most
terrestrial locomotion 262 Appendix - a model of non?]steady speed walking
265 10A.1 Spring?]mass inverted pendulum model of walking 265 10A.2
Recovery ratio calculation 269 References 271 Chapter 11 Non?]Steady
Locomotion 277 Monica A. Daley 11.1 Introduction 277 11.1.1 Why study
non?]steady locomotion? 278 11.2 A pproaches to studying non?]steady
locomotion 279 11.2.1 Simple mechanical models 280 11.2.2 Research
approaches to non?]steady locomotion 281 11.3 Themes from recent studies of
non?]steady locomotion 282 11.3.1 Limits to maximal acceleration 282 11.3.2
Morphological and behavioral factors in turning mechanics 283 11.4 The role
of intrinsic mechanics for stability and robustness of locomotion 288
11.4.1 Some definitions 289 11.4.2 Measures of sensitivity and robustness
290 11.4.3 What do we learn about stability from simple models of running?
291 11.4.4 Limitations to stability analysis of simple models 295 11.4.5
The relationship between ground contact conditions and leg mechanics on
uneven terrain 296 11.4.6 Compromises among economy, robustness and injury
avoidance in uneven terrain 298 11.5 Proximal?]distal inter?]joint
coordination in non?]steady locomotion 299 References 302 Chapter 12 The
Evolution of Terrestrial Locomotion in Bats: the Bad, the Ugly, and the
Good 307 Daniel K. Riskin, John E. A. Bertram and John W. Hermanson 12.1
Bats on the ground: like fish out of water? 307 12.2 Species?]level
variation in walking ability 308 12.3 How does anatomy influence crawling
ability? 309 12.4 Hindlimbs and the evolution of flight 311 12.5 Moving a
bat's body on land: the kinematics of quadrupedal locomotion 315 12.6
Evolutionary pressures leading to capable terrestrial locomotion 318 12.7
Conclusions and future work 319 Acknowledgements 320 References 320 Chapter
13 The Fight or Flight Dichotomy: Functional Trade?]Off in Specialization
for Aggression Versus Locomotion 325 David R. Carrier 13.1 Introduction325
13.1.1 Why fighting is important 327 13.1.2 Size sexual dimorphism as an
indicator of male?]male aggression 328 13.2 Trade?]offs in specialization
for aggression versus locomotion 329 13.2.1 The evolution of short legs -
specialization for aggression? 329 13.2.2 Muscle architecture of limbs
specialized for running versus fighting 331 13.2.3 Mechanical properties of
limb bones that are specialized for running versus fighting 334 13.2.4 The
function of foot posture: aggression versus locomotor economy 334 13.3
Discussion 338 References 341 Chapter 14 Design for Prodigious Size without
Extreme Body Mass: Dwarf Elephants, Differential Scaling and Implications
for Functional Adaptation 349 John E. A. Bertram 14.1 Introduction 349 14.2
Elephant form, mammalian scaling and dwarfing 351 14.2.1 Measurements 356
14.2.2 Observations 356 14.3 Interpretation 357 Acknowledgements 364
References 364 Chapter 15 Basic Mechanisms of Bipedal Locomotion:
Head?]Supported Loads and Strategies to Reduce the Cost of Walking 369
James R. Usherwood and John E. A. Bertram 15.1 Introduction 369 15.2
Head?]supported loads in human?]mediated transport 370 15.2.1 Can the
evidence be depended upon? 371 15.3 Potential energy saving advantages 373
15.4 A simple alternative model 376 15.5 Conclusions 382 References 382
Chapter 16 Would a Horse on the Moon Gallop? Directions Available in
Locomotion Research (and How Not to Spend Too Much Time Exploring Blind
Alleys) 385 John E. A. Bertram 16.1 Introduction 385 References 392 Index
393
Historical Perspectives on Mammalian Locomotion 1 John E. A. Bertram 1.1
Introduction 1 1.2 The ancients and the contemplation of motion 2 1.3 The
European Renaissance and foundations of the age of discovery 3 1.4 The era
of technological observation 5 1.5 Physiology and mechanics of terrestrial
locomotion - cost and consequences 7 1.6 Comparative studies of gait 10 1.6
Re?]interpreting the mechanics: a fork in the road, or simply seeing the
other side of the coin? 13 1.7 The biological source of cost 13 1.8 The
physical source of cost (with biological consequences) - the road less
traveled 14 1.9 Conclusions 21 References 21 Chapter 2 Considering Gaits:
Descriptive Approaches 27 John E. A. Bertram 2.1 Introduction 27 2.2
Defining the fundamental gaits 28 2.3 Classifying and comparing the
fundamental gaits 30 2.4 Symmetric gaits 32 2.5 A symmetric gaits 34 2.6
Beyond "Hildebrand plots" 40 2.7 Statistical classification 43 2.8 Neural
regulation and emergent criteria 45 2.9 Mechanical measures as descriptions
of gaits 47 2.10 Conclusion 47 References 48 Chapter 3 Muscles as Actuators
51 Anne K. Gutmann and John E. A. Bertram 3.1 Introduction 51 3.2 Basic
muscle operation 52 3.2.1 Sliding filament theory - the basis for
cross?]bridge theory 52 3.2.2 Basic cross?]bridge theory 52 3.2.3
Multi?]state cross?]bridge models 57 3.3 Some alternatives to cross?]bridge
theory 59 3.4 Force production 60 3.4.1 Isometric force production 60 3.4.2
Non?]isometric force production 63 3.5 The Hill?]type model 66 3.6
Optimizing work, power, and efficiency 68 3.7 Muscle architecture 70 3.7.1
The sarcomere as the fundamental contractile unit 70 3.7.2 Muscle geometry
70 3.7.3 Elastic energy storage and return 72 3.7.4 Damping/energy
dissipation 72 3.8 Other factors that influence muscle performance 73 3.8.1
Fiber type 73 3.9 A ctivation and recruitment 75 3.10 What does muscle do
best? 76 References 76 Chapter 4 Concepts in Locomotion: Levers, Struts,
Pendula and Springs 79 John E. A. Bertram 4.1 Introduction 79 4.2 The limb:
How details can obscure functional role 83 4.3 Limb function in stability
and the concept of the "effective limb" 85 4.3.1 Considering the mechanisms
of stability 85 4.3.2 The role of the effective limb 88 4.4 Levers and
struts 89 4.5 Ground reaction force in gaits 92 4.5.1 Trot 94 4.5.2 Walk 96
4.5.3 Gallop 97 4.6 The consequence of applied force: CoM motion, pendula
and springs 98 4.7 Energy exchange in locomotion - valuable or inevitable?
102 4.8 Momentum and energy in locomotion: dynamic fundamentals 103 4.9
Energy - lost unless recovered, or available unless lost? 104 References
105 Chapter 5 Concepts in Locomotion: Wheels, Spokes, Collisions and
Insight from the Center of Mass 111 John E. A. Bertram 5.1 Introduction 111
5.2 Understanding brachiation: an analogy for terrestrial locomotion 112
5.3 Bipedal walking: inverted pendulum or inverted "collision?]limiting
brachiator analog"? 117 5.4 Basic dynamics of the step?]to?]step transition
in bipedal walking 120 5.5 Subtle dynamics of the step?]to?]step transition
in bipedal walking and running 124 5.6 Pseudo?]elastic motion and true
elastic return in running gaits 130 5.7 Managing CoM motion in quadrupedal
gaits 131 5.7.1 Walk 132 5.7.2 Trot 133 5.7.3 Gallop 133 5.8 Conclusion 138
References 139 Chapter 6 Reductionist Models of Walking and Running 143
James R. Usherwood 6.1 Part 1: Bipedal locomotion and "the ultimate cost of
legged locomotion?" 143 6.1.1 Introduction 143 6.1.2 Reductionist models of
walking 144 6.1.3 The benefit of considering locomotion as inelastic 150
6.2 Part 2: quadrupedal locomotion 158 6.2.1 Introduction 158 6.2.2
Quadrupedal dynamic walking and collisions 158 6.2.3 Higher speed
quadrupedal gaits 161 6.2.4 Further success of reductionist mechanics 162
Appendix A: Analytical approximation for costs of transport including legs
and "guts and gonads" losses 166 6A.1 List of symbols 166 6A.2 Period
definitions for a symmetrically running biped 166 6A.3 Ideal work for the
leg 167 6A.4 Vertical work calculations for leg 168 6A.5 Horizontal work
calculations for leg 169 6A.6 Hysteresis costs of "guts and gonads"
deflections 169 6A.7 Cost of transport 170 References 170 Chapter 7
Whole?]Body Mechanics: How Leg Compliance Shapes the Way We Move 173 Andre
Seyfarth, Hartmut Geyer, Susanne Lipfert, J. Rummel, Yvonne Blum, M. Maus
and D. Maykranz 7.1 Introduction 173 7.2 Jumping for distance - a
goal?]directed movement 175 7.3 Running for distance - what is the goal?
177 7.4 Cyclic stability in running 178 7.5 The wheel in the leg - how leg
retraction enhances running stability 179 7.6 Walking with compliant legs
180 7.7 A dding an elastically coupled foot to the spring?]mass model 184
7.8 The segmented leg - how does joint function translate into leg
function? 185 7.9 Keeping the trunk upright during locomotion 187 7.10 The
challenge of setting up more complex models 188 Notes 190 References190
Chapter 8 The Most Important Feature of an Organism's Biology: Dimension,
Similarity and Scale 193 John E. A. Bertram 8.1 Introduction 193 8.2 The
most basic principle: surface area to volume relations 194 8.3 A ssessing
scale effects 197 8.4 Physiology and scaling 198 8.5 The allometric
equation: the power function of scaling 203 8.6 The standard scaling models
207 8.6.1 Geometric similarity 208 8.6.2 Static stress similarity 209 8.6.3
Elastic similarity 209 8.7 Differential scaling - where the limit may
change 210 8.7.1 A ssessing the assumptions 215 8.8 A fractal view of
scaling 215 8.9 Making valid comparisons: measurement, dimension and
functional criteria 217 8.9.1 Considering units 217 8.9.2 Fundamental and
derived units 219 8.9.3 Froude number: a dimensionless example 222
References 223 Chapter 9 Accounting for the Influence of Animal Size on
Biomechanical Variables: Concepts and Considerations 229 Sharon Bullimore
9.1 Introduction 229 9.2 Commonly used approaches to accounting for size
differences 230 9.2.1 Dividing by body mass 230 9.2.2 Dimensionless
parameters 232 9.3 Empirical scaling relationships 237 9.4 Selected
biomechanical parameters 238 9.4.1 Ground reaction force 238 9.4.2 Muscle
force 239 9.4.3 Muscle velocity 242 9.4.4 Running speed 242 9.4.5 Jump
height 244 9.4.6 Elastic energy storage 246 9.5 Conclusions 247
Acknowledgements 247 References 247 Chapter 10 Locomotion in Small
Tetrapods: Size?]Based Limitations to "Universal Rules" in Locomotion 251
Audrone R. Biknevicius, Stephen M. Reilly and Elvedin Kljuno 10.1
Introduction 251 10.2 A ctive mechanisms contributing to the high cost of
transport in small tetrapods 254 10.3 Limited passive mechanisms for
reducing cost of transport in small tetrapods 255 10.4 Gait transitions
from vaulting to bouncing mechanics 257 10.5 The "unsteadiness" of most
terrestrial locomotion 262 Appendix - a model of non?]steady speed walking
265 10A.1 Spring?]mass inverted pendulum model of walking 265 10A.2
Recovery ratio calculation 269 References 271 Chapter 11 Non?]Steady
Locomotion 277 Monica A. Daley 11.1 Introduction 277 11.1.1 Why study
non?]steady locomotion? 278 11.2 A pproaches to studying non?]steady
locomotion 279 11.2.1 Simple mechanical models 280 11.2.2 Research
approaches to non?]steady locomotion 281 11.3 Themes from recent studies of
non?]steady locomotion 282 11.3.1 Limits to maximal acceleration 282 11.3.2
Morphological and behavioral factors in turning mechanics 283 11.4 The role
of intrinsic mechanics for stability and robustness of locomotion 288
11.4.1 Some definitions 289 11.4.2 Measures of sensitivity and robustness
290 11.4.3 What do we learn about stability from simple models of running?
291 11.4.4 Limitations to stability analysis of simple models 295 11.4.5
The relationship between ground contact conditions and leg mechanics on
uneven terrain 296 11.4.6 Compromises among economy, robustness and injury
avoidance in uneven terrain 298 11.5 Proximal?]distal inter?]joint
coordination in non?]steady locomotion 299 References 302 Chapter 12 The
Evolution of Terrestrial Locomotion in Bats: the Bad, the Ugly, and the
Good 307 Daniel K. Riskin, John E. A. Bertram and John W. Hermanson 12.1
Bats on the ground: like fish out of water? 307 12.2 Species?]level
variation in walking ability 308 12.3 How does anatomy influence crawling
ability? 309 12.4 Hindlimbs and the evolution of flight 311 12.5 Moving a
bat's body on land: the kinematics of quadrupedal locomotion 315 12.6
Evolutionary pressures leading to capable terrestrial locomotion 318 12.7
Conclusions and future work 319 Acknowledgements 320 References 320 Chapter
13 The Fight or Flight Dichotomy: Functional Trade?]Off in Specialization
for Aggression Versus Locomotion 325 David R. Carrier 13.1 Introduction325
13.1.1 Why fighting is important 327 13.1.2 Size sexual dimorphism as an
indicator of male?]male aggression 328 13.2 Trade?]offs in specialization
for aggression versus locomotion 329 13.2.1 The evolution of short legs -
specialization for aggression? 329 13.2.2 Muscle architecture of limbs
specialized for running versus fighting 331 13.2.3 Mechanical properties of
limb bones that are specialized for running versus fighting 334 13.2.4 The
function of foot posture: aggression versus locomotor economy 334 13.3
Discussion 338 References 341 Chapter 14 Design for Prodigious Size without
Extreme Body Mass: Dwarf Elephants, Differential Scaling and Implications
for Functional Adaptation 349 John E. A. Bertram 14.1 Introduction 349 14.2
Elephant form, mammalian scaling and dwarfing 351 14.2.1 Measurements 356
14.2.2 Observations 356 14.3 Interpretation 357 Acknowledgements 364
References 364 Chapter 15 Basic Mechanisms of Bipedal Locomotion:
Head?]Supported Loads and Strategies to Reduce the Cost of Walking 369
James R. Usherwood and John E. A. Bertram 15.1 Introduction 369 15.2
Head?]supported loads in human?]mediated transport 370 15.2.1 Can the
evidence be depended upon? 371 15.3 Potential energy saving advantages 373
15.4 A simple alternative model 376 15.5 Conclusions 382 References 382
Chapter 16 Would a Horse on the Moon Gallop? Directions Available in
Locomotion Research (and How Not to Spend Too Much Time Exploring Blind
Alleys) 385 John E. A. Bertram 16.1 Introduction 385 References 392 Index
393