- Email: [email protected]
Vertebrate Biomechanics and Evolution
ARTICLE IN PRESS
Journal of Biomechanics 37 (2004) 1627–1629
Book reviews Vertebrate Biomechanics and Evolution Edited by V.L. Bels, J-P. Gasc and A Casinos; BIOS Scientific Publishers Limited, Oxford, UK This thought-provoking volume is an admirable attempt to present a wide range of vertebrate biomechanics investigations within the context of vertebrate evolution. It is comprised of selected papers that were presented in a symposium sponsored by the Society for Experimental Biology entitled ‘‘Biomechanics and Evolution’’ that was held at the University of Canterbury in 2001. The book covers many different biomechanical functions and organ systems in a broad range of animals. The book, however, goes beyond the rather straightforward question of ‘‘how do animals work?’’ The editors and authors of the various chapters extend their considerations to issues that are often addresses within a neo-Darwinian perspective. At the same time, the reader is repeatedly reminded of the ideas of Cuvier on the importance of form to function and the correlation of parts in vertebrate anatomy. More than four hundred million years of biomechanics is a lot to cover in 332 pages. The volume succeeds to a great extent as a result of the editors’ efforts to introduce the chapters within a framework of key concepts that are considered in evolution theory. They organize the book into two parts, with each part consisting of ten and nine chapters, respectively. Part I is entitled ‘‘Theory and Aquatic Animals’’ and Part II is ‘‘Terrestrial and Avian Animals’’. They have written the first chapter in each part to introduce us of some of the classic questions and issues that are relevant in the subsequent chapters. We are reminded of Earnst Mayr’s statement that nothing makes sense outside the perspective of evolution. Vertebrate ontogenetic and phylogenetic histories can be used with biomechanics to understand not only how animals work, but also why they work in that particular way and what other ways they could conceivably work. The introductory chapter for Part I is followed by a brief but interesting chapter by McNeill Alexander that reviews some of the early work in paleobiomechanics and points out the difficulties in investigating the mechanics of extinct animals. A 1943 study by Kermack is described wherein a model of a Devonian ostracoderm, one of the first vertebrates, was placed in a wind tunnel to better understand the swimming mechanics of this heavily armored fish. Other studies
ranging from the flight mechanics of pterosaurs to the running speeds of dinosaurs are introduced and the problems in making estimates of muscle function, bone material properties, body masses and other important mechanical parameters are discussed. These considerations lead us to appreciate both the benefits and limitations of using the morphology and biomechanics of extant animals as a basis for predicting the function of extinct species. The following chapter by Garland deals with selection experiments. He points out that selection experiments are almost non-existent in evolutionary biomechanics but have a long history in the development of agricultural products and the gradual domestication of plant and animals, especially dogs. He goes on to describe some of the considerations for selection experiments in a scientific setting. He shows interesting data from a mouse study in which breeding of successive generations was based on running abilities. In addition to being interesting in it’s own right, the chapter introduces some important concept for biomechanicians. He points out that some biomechanics researches assume that evolution by natural selection yields ‘‘optimum solutions’’ yet many now feel that evolution often yields merely ‘‘adequate’’ solutions. Garland also brings up questions of developmental and evolutionary constraints on vertebrate ‘‘design’’. He notes that all constraints are ultimately genetic in origin. Schulter is cited in arguing that ‘‘‘genetic constraints’ exist when alleles are absent from a population or when alleles affect multiple traits (pleiotropy) in ways that run counter to the prevailing selection.’’ He also considers the effect of random genetic drifts and the probability that these lead to multiple design solutions. These issues are fundamental in any study of biomechanics in evolution and resonate throughout the book with many of the authors. The ensuing chapters cover a wide range of interesting topics in vertebrate biomechanics including muscle plasticity, jaw mechanics and design, suction feeding in fish, respiration, swimming mechanics, and salamander feeding biomechanics. Each of the chapters touches on an aspect of evolution in some way. The content of Part II continues the themes of evolution onto land and into the air. Topics include feeding behavior and biomechanics of reptiles and birds, the biomechanics of the avian skull, terrestrial locomotion in tetrapods, lizards, and hominids, and the origins and diversity of the avian wing. The book ends with a chapter by Lauder who
ARTICLE IN PRESS 1628
Book reviews / Journal of Biomechanics 37 (2004) 1627–1629
attempts to put the current efforts in evolution research into focus and lays out areas in which biomechanics can play an important role in the future. I must say that I was hoping to see something more about the evolution of mechanobiology and the roles of mechanosensitive genes in morphogenesis. Although I realize that I may have a particular bias in this regard, it seems that this is an area of research that is growing rapidly. Historically, this area of biology has been referred to as ‘‘functional adaptation’’ and was thought by Roux, Wolff, and Darwin to have an important role in evolution. That being said, I feel that this book is a
fine contribution that lays out important issues and approaches. It is a ‘‘must read’’ for anyone wishing to understand vertebrate biomechanics within an evolutionary framework.
Dennis R. Carter Biomechanical Engineering Division, Departments of Mechanical Engineering and Bioengineering, Palo Alto VA Health Care System and Stanford University, Durand Bldg., Room 215, Stanford, CA 94305-4038, USA E-mail address: [email protected]
doi:10.1016/j.jbiomech.2004.01.016
Comparative biomechanics: life’s physical world: Steven Vogel; Princeton University Press, Princeton, NJ, 2003, 580pp., ISBN 0-691-11297-5 Steve Vogel is a great populariser of biomechanics. He has written many books over the years on the subject, for instance: ‘Life’s devices’ ‘Cats’ paws and catapults’ ‘Vital circuits’. His writing is characterised by the ability to give considerable physical insight along with humour and with a very broad knowledge of biology. The present book is intended ‘‘to be used as a textbook for undergraduates in biology at any level beyond an introductory course or undergraduates in any other area of science or engineering willing to refer to a basic textbook in biology, or graduate students or other scientists and engineers who need an entry point that gets them into the content and literature of biomechanics.’’ The level of physical and mathematical sophistication required is not high: ‘‘basic college physics and calculusyand ysome familiarity with algebra and with exponents and logarithms.’’ As Vogel writes, biomechanics covers ‘an awesome diversity of things’ and he has to be selective in what he has chosen to write about. In fact he has covered a very wide part of biomechanics as seen from the biological, though not the clinical, point of view. He starts with some introductory chapters on the concepts of quantities, units, scaling, dimensional analysis and various other physical concepts that will come in use later, such as moments of area, fractal dimensions, and velocity gradients. Even in these chapters there is constant reference of their subject matter to biological processes. The main part of the book has two blocks, on fluids and on solids and structures. The fluid block has chapters on fluids ‘at rest’, viscosity and flow, pressures in flowing fluids, events near surfaces, flow in tubes, circulatory systems, very low Reynolds number events, lift, thrust,
and moving at the liquid/gas interface. The second block has chapters on mechanics of biological materials, and their fitness for purpose, viscoelasticity, simple structures, and more complex structures, hydrostatic structures, structural systems (trees, skeletons etc.), muscles and other motile systems, and land locomotion. Finally a very interesting chapter tying up loose ends and putting biomechanics into perspective. There is a relentless tying of all the mechanical concepts to real biological situations. Plants are dealt with as well as animals, and there are passing references to fungi and bacteria (whose flagella are moved by the only true living wheel). Indeed the breadth of coverage is impressive, and is testament to Vogel’s career-long interest in the whole of biology. Vogel has intentionally not dealt in detail with some of the more physically tricky areas of biomechanics such as viscoelasticity and unsteady flows, even though they are of considerable importance to organisms. The book is large, about 230,000 words, with an attractive cover. It is copiously illustrated (I counted about 230 illustrations and graphs), somewhat less copiously equationed (about 160) and with about 35 tables. There are a very large number of references for a textbook, more than 600. These will certainly be a help for people coming to the subject from, say, engineering. In writing reviews, one should always leave until the end the impression that one wishes the reader to leave with. Good followed by bad is remembered mainly as bad, and vice versa. So, I shall start by mentioning a few things that I did not like about this book. First, Vogel’s extreme love of donnish periphrasis and puns does sometimes get in the way of the reader’s understanding. Take these examples: ‘‘the cost of making mucusymay preclude its routine use, so the trick may be reserved for times when
Journal of Biomechanics 37 (2004) 1627–1629
Book reviews Vertebrate Biomechanics and Evolution Edited by V.L. Bels, J-P. Gasc and A Casinos; BIOS Scientific Publishers Limited, Oxford, UK This thought-provoking volume is an admirable attempt to present a wide range of vertebrate biomechanics investigations within the context of vertebrate evolution. It is comprised of selected papers that were presented in a symposium sponsored by the Society for Experimental Biology entitled ‘‘Biomechanics and Evolution’’ that was held at the University of Canterbury in 2001. The book covers many different biomechanical functions and organ systems in a broad range of animals. The book, however, goes beyond the rather straightforward question of ‘‘how do animals work?’’ The editors and authors of the various chapters extend their considerations to issues that are often addresses within a neo-Darwinian perspective. At the same time, the reader is repeatedly reminded of the ideas of Cuvier on the importance of form to function and the correlation of parts in vertebrate anatomy. More than four hundred million years of biomechanics is a lot to cover in 332 pages. The volume succeeds to a great extent as a result of the editors’ efforts to introduce the chapters within a framework of key concepts that are considered in evolution theory. They organize the book into two parts, with each part consisting of ten and nine chapters, respectively. Part I is entitled ‘‘Theory and Aquatic Animals’’ and Part II is ‘‘Terrestrial and Avian Animals’’. They have written the first chapter in each part to introduce us of some of the classic questions and issues that are relevant in the subsequent chapters. We are reminded of Earnst Mayr’s statement that nothing makes sense outside the perspective of evolution. Vertebrate ontogenetic and phylogenetic histories can be used with biomechanics to understand not only how animals work, but also why they work in that particular way and what other ways they could conceivably work. The introductory chapter for Part I is followed by a brief but interesting chapter by McNeill Alexander that reviews some of the early work in paleobiomechanics and points out the difficulties in investigating the mechanics of extinct animals. A 1943 study by Kermack is described wherein a model of a Devonian ostracoderm, one of the first vertebrates, was placed in a wind tunnel to better understand the swimming mechanics of this heavily armored fish. Other studies
ranging from the flight mechanics of pterosaurs to the running speeds of dinosaurs are introduced and the problems in making estimates of muscle function, bone material properties, body masses and other important mechanical parameters are discussed. These considerations lead us to appreciate both the benefits and limitations of using the morphology and biomechanics of extant animals as a basis for predicting the function of extinct species. The following chapter by Garland deals with selection experiments. He points out that selection experiments are almost non-existent in evolutionary biomechanics but have a long history in the development of agricultural products and the gradual domestication of plant and animals, especially dogs. He goes on to describe some of the considerations for selection experiments in a scientific setting. He shows interesting data from a mouse study in which breeding of successive generations was based on running abilities. In addition to being interesting in it’s own right, the chapter introduces some important concept for biomechanicians. He points out that some biomechanics researches assume that evolution by natural selection yields ‘‘optimum solutions’’ yet many now feel that evolution often yields merely ‘‘adequate’’ solutions. Garland also brings up questions of developmental and evolutionary constraints on vertebrate ‘‘design’’. He notes that all constraints are ultimately genetic in origin. Schulter is cited in arguing that ‘‘‘genetic constraints’ exist when alleles are absent from a population or when alleles affect multiple traits (pleiotropy) in ways that run counter to the prevailing selection.’’ He also considers the effect of random genetic drifts and the probability that these lead to multiple design solutions. These issues are fundamental in any study of biomechanics in evolution and resonate throughout the book with many of the authors. The ensuing chapters cover a wide range of interesting topics in vertebrate biomechanics including muscle plasticity, jaw mechanics and design, suction feeding in fish, respiration, swimming mechanics, and salamander feeding biomechanics. Each of the chapters touches on an aspect of evolution in some way. The content of Part II continues the themes of evolution onto land and into the air. Topics include feeding behavior and biomechanics of reptiles and birds, the biomechanics of the avian skull, terrestrial locomotion in tetrapods, lizards, and hominids, and the origins and diversity of the avian wing. The book ends with a chapter by Lauder who
ARTICLE IN PRESS 1628
Book reviews / Journal of Biomechanics 37 (2004) 1627–1629
attempts to put the current efforts in evolution research into focus and lays out areas in which biomechanics can play an important role in the future. I must say that I was hoping to see something more about the evolution of mechanobiology and the roles of mechanosensitive genes in morphogenesis. Although I realize that I may have a particular bias in this regard, it seems that this is an area of research that is growing rapidly. Historically, this area of biology has been referred to as ‘‘functional adaptation’’ and was thought by Roux, Wolff, and Darwin to have an important role in evolution. That being said, I feel that this book is a
fine contribution that lays out important issues and approaches. It is a ‘‘must read’’ for anyone wishing to understand vertebrate biomechanics within an evolutionary framework.
Dennis R. Carter Biomechanical Engineering Division, Departments of Mechanical Engineering and Bioengineering, Palo Alto VA Health Care System and Stanford University, Durand Bldg., Room 215, Stanford, CA 94305-4038, USA E-mail address: [email protected]
doi:10.1016/j.jbiomech.2004.01.016
Comparative biomechanics: life’s physical world: Steven Vogel; Princeton University Press, Princeton, NJ, 2003, 580pp., ISBN 0-691-11297-5 Steve Vogel is a great populariser of biomechanics. He has written many books over the years on the subject, for instance: ‘Life’s devices’ ‘Cats’ paws and catapults’ ‘Vital circuits’. His writing is characterised by the ability to give considerable physical insight along with humour and with a very broad knowledge of biology. The present book is intended ‘‘to be used as a textbook for undergraduates in biology at any level beyond an introductory course or undergraduates in any other area of science or engineering willing to refer to a basic textbook in biology, or graduate students or other scientists and engineers who need an entry point that gets them into the content and literature of biomechanics.’’ The level of physical and mathematical sophistication required is not high: ‘‘basic college physics and calculusyand ysome familiarity with algebra and with exponents and logarithms.’’ As Vogel writes, biomechanics covers ‘an awesome diversity of things’ and he has to be selective in what he has chosen to write about. In fact he has covered a very wide part of biomechanics as seen from the biological, though not the clinical, point of view. He starts with some introductory chapters on the concepts of quantities, units, scaling, dimensional analysis and various other physical concepts that will come in use later, such as moments of area, fractal dimensions, and velocity gradients. Even in these chapters there is constant reference of their subject matter to biological processes. The main part of the book has two blocks, on fluids and on solids and structures. The fluid block has chapters on fluids ‘at rest’, viscosity and flow, pressures in flowing fluids, events near surfaces, flow in tubes, circulatory systems, very low Reynolds number events, lift, thrust,
and moving at the liquid/gas interface. The second block has chapters on mechanics of biological materials, and their fitness for purpose, viscoelasticity, simple structures, and more complex structures, hydrostatic structures, structural systems (trees, skeletons etc.), muscles and other motile systems, and land locomotion. Finally a very interesting chapter tying up loose ends and putting biomechanics into perspective. There is a relentless tying of all the mechanical concepts to real biological situations. Plants are dealt with as well as animals, and there are passing references to fungi and bacteria (whose flagella are moved by the only true living wheel). Indeed the breadth of coverage is impressive, and is testament to Vogel’s career-long interest in the whole of biology. Vogel has intentionally not dealt in detail with some of the more physically tricky areas of biomechanics such as viscoelasticity and unsteady flows, even though they are of considerable importance to organisms. The book is large, about 230,000 words, with an attractive cover. It is copiously illustrated (I counted about 230 illustrations and graphs), somewhat less copiously equationed (about 160) and with about 35 tables. There are a very large number of references for a textbook, more than 600. These will certainly be a help for people coming to the subject from, say, engineering. In writing reviews, one should always leave until the end the impression that one wishes the reader to leave with. Good followed by bad is remembered mainly as bad, and vice versa. So, I shall start by mentioning a few things that I did not like about this book. First, Vogel’s extreme love of donnish periphrasis and puns does sometimes get in the way of the reader’s understanding. Take these examples: ‘‘the cost of making mucusymay preclude its routine use, so the trick may be reserved for times when