the changing role of the embryo in evolutionary thought: roots of evo-devo

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Book Reviews Human Osteology & Skeletal Radiology: An Atlas and Guide . By E. Matshes, B. Burgridge, B. Sher, A. Mohamed, and B. Juurlink. Illustrated by C. Allen. 433 pp. Boca Raton, FL: CRC Press. 2005. $99.95 (cloth). Human Osteology & Skeletal Radiology: An Atlas and Guide is an exquisitely illu- strated atlas with clearly reproduced and well-labeled photographs and radiographs printed on high-quality glossy paper. This book follows in the fine tradition that we have come to expect from other CRC Press publications. The authors write in the pre- face that their goal was to ‘‘produce a clear and useable photographic atlas and accom- panying laboratory manual that would be of use to students at all levels of study.’’ I believe that they have succeeded. Most of the authors are faculty members at the College of Medicine, University of Saskatchewan, and have clinical degrees. I expected that this would be a good book for students in the health sciences, and I was not disappointed. You could tell by the succinct discussions, lists of defined terms and landmarks, short clin- ical concept sections, and bulleted points at the beginning of each of the seven chapters that these authors have had many years of gross anatomy teaching experience among them. However, I was also very surprised and impressed at the short sections on human skeletal variability and fragment identification for students interested in human osteology and forensics. These sec- tions are reminiscent of forensic works by Krogman and Iscan (1986), Bass (1995), and Schwartz (1995). The book is organized along a regional ana- tomical approach with an introductory chap- ter and two units with three chapters each on the axial and appendicular skeletons. Each chapter starts out with a short introductory text section describing the region, nomencla- ture, and age and population variability. This is followed by many large, glossy, and well- labeled photographs and drawings. In addi- tion, occasional plain film radiographs and computed tomographic (CT) and magnetic resonance (MR) scan images are used. The radiographic images are very clear and well- labeled, and their orientation is always described in the legends. The skeletal mate- rial used for the photographs was well pre- served, of high quality, and came from a number of different individuals. This gives the students a good idea of normal human skeletal variability, which is lacking in some of the artwork-only atlases. The photographs were well lit with a good depth of field. All of the structures of interest were clearly visi- ble, highlighted, and in focus, in contrast to some of the pictures seen in photographic atlases of cadaveric material. There are also occasional highlighted ‘‘boxes’’ in each chap- ter which display photographs, drawings, or radiographs of particularly important struc- tures or relationships and pedagogical descrip- tions of interest (e.g., Visualizing Pterion, Nutrient Foramina, Edentulous Mandible). There are also short sections on how function relates to particular skeletal morphologies (e.g., fibular, foot, and radial shapes). The last section in most chapters deals with indi- vidual bones with clear, higher-magnifica- tion photographs and drawings. Very complete sections on the tooth classes and the individual cranial bones, vertebrae, and carpal and tarsal bones are included. The text describing left-right side determination and fragment identification is succinct. The first unit on the axial skeletal section has an extensive chapter on the Skull (Chapter 2, 128 pages long), and shorter chapters on the Vertebral Column (Chapter 3, 84 pages long), and the Sternum and Ribs (Chapter 4, 20 pages long). The second unit on the appendicular skeleton has a chapter on the Shoulder and Upper Limbs (Chapter 5, 92 pages long), the Pelvis (Chapter 6, 16 pages long), and the Lower Extremities (Chapter 7, 82 pages long). I believe that this atlas will prove very helpful and popular to students in gross anatomy and human osteology courses, as was the goal of the authors. However, profes- sional anatomists and human osteologists might find this atlas a bit simplified and not complete enough for a personal reference text. Although the figures are clearly labeled, oriented, and well reproduced, many of the figures have only a few land- marks identified on them. In addition, the table of contents could have been expanded a little more with the page numbers of the various sections in each chapter listed. The sections in the text already have unique numerical headings, but their corresponding page numbers are not listed in the table of contents. I had to continually flip through ß 2005 Wiley-Liss, Inc. AMERICAN JOURNAL OF HUMAN BIOLOGY 17:665–672 (2005)

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Page 1: The changing role of the embryo in evolutionary thought: Roots of evo-devo

Book Reviews

Human Osteology & Skeletal Radiology: AnAtlas and Guide. By E. Matshes,B. Burgridge, B. Sher, A. Mohamed, andB. Juurlink. Illustrated by C. Allen. 433 pp.Boca Raton, FL: CRC Press. 2005.$99.95 (cloth).

Human Osteology & Skeletal Radiology:An Atlas and Guide is an exquisitely illu-strated atlas with clearly reproduced andwell-labeled photographs and radiographsprinted on high-quality glossy paper. Thisbook follows in the fine tradition that wehave come to expect from other CRC Presspublications. The authors write in the pre-face that their goal was to ‘‘produce a clearand useable photographic atlas and accom-panying laboratory manual that would be ofuse to students at all levels of study.’’ Ibelieve that they have succeeded. Most ofthe authors are faculty members at the Collegeof Medicine, University of Saskatchewan, andhave clinical degrees. I expected that thiswould be a good book for students in the healthsciences, and I was not disappointed. Youcould tell by the succinct discussions, listsof defined terms and landmarks, short clin-ical concept sections, and bulleted points atthe beginning of each of the seven chaptersthat these authors have had many years ofgross anatomy teaching experience amongthem. However, I was also very surprisedand impressed at the short sections onhuman skeletal variability and fragmentidentification for students interested inhuman osteology and forensics. These sec-tions are reminiscent of forensic works byKrogman and Iscan (1986), Bass (1995),and Schwartz (1995).

The book is organized along a regional ana-tomical approach with an introductory chap-ter and two units with three chapters each onthe axial and appendicular skeletons. Eachchapter starts out with a short introductorytext section describing the region, nomencla-ture, and age and population variability. Thisis followed by many large, glossy, and well-labeled photographs and drawings. In addi-tion, occasional plain film radiographs andcomputed tomographic (CT) and magneticresonance (MR) scan images are used. Theradiographic images are very clear and well-labeled, and their orientation is alwaysdescribed in the legends. The skeletal mate-

rial used for the photographs was well pre-served, of high quality, and came from anumber of different individuals. This givesthe students a good idea of normal humanskeletal variability, which is lacking in someof the artwork-only atlases. The photographswere well lit with a good depth of field. All ofthe structures of interest were clearly visi-ble, highlighted, and in focus, in contrast tosome of the pictures seen in photographicatlases of cadaveric material. There are alsooccasional highlighted ‘‘boxes’’ in each chap-ter which display photographs, drawings, orradiographs of particularly important struc-tures or relationships and pedagogical descrip-tions of interest (e.g., Visualizing Pterion,Nutrient Foramina, Edentulous Mandible).There are also short sections on how functionrelates to particular skeletal morphologies(e.g., fibular, foot, and radial shapes). Thelast section in most chapters deals with indi-vidual bones with clear, higher-magnifica-tion photographs and drawings. Verycomplete sections on the tooth classes andthe individual cranial bones, vertebrae, andcarpal and tarsal bones are included. Thetext describing left-right side determinationand fragment identification is succinct. Thefirst unit on the axial skeletal section has anextensive chapter on the Skull (Chapter 2,128 pages long), and shorter chapters on theVertebral Column (Chapter 3, 84 pageslong), and the Sternum and Ribs (Chapter4, 20 pages long). The second unit on theappendicular skeleton has a chapter on theShoulder and Upper Limbs (Chapter 5, 92pages long), the Pelvis (Chapter 6, 16 pageslong), and the Lower Extremities (Chapter 7,82 pages long).

I believe that this atlas will prove veryhelpful and popular to students in grossanatomy and human osteology courses, aswas the goal of the authors. However, profes-sional anatomists and human osteologistsmight find this atlas a bit simplified andnot complete enough for a personal referencetext. Although the figures are clearlylabeled, oriented, and well reproduced,many of the figures have only a few land-marks identified on them. In addition, thetable of contents could have been expandeda little more with the page numbers of thevarious sections in each chapter listed. Thesections in the text already have uniquenumerical headings, but their correspondingpage numbers are not listed in the table ofcontents. I had to continually flip through

� 2005 Wiley-Liss, Inc.

AMERICAN JOURNAL OF HUMAN BIOLOGY 17:665–672 (2005)

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the chapters to find the various chapters andsections. The book is also full sized (8.5inches � 11 inches) and hard-backed, whichwill work well in the gross anatomy andosteology laboratories but may be a bit cum-bersome out in the field. The authors andpublishers might consider releasing a soft-covered, field manual version of this bookat some time for students at archeologicalor forensic sites.

However, even with these very minor con-cerns, I believe that this atlas will make animportant addition to the anatomical atlasescurrently in use in gross anatomy and parti-cularly human osteology courses. I congratu-late the authors on this scholarly and well-produced piece of work, and I plan to recom-mend it to my own medical and dental schoolgross anatomy classes next year.

LITERATURE CITED

Bass, W. 1995. Human osteology: a laboratory and fieldmanual. Columbia, MO: Missouri Archeology Society.

Krogman W, Iscan M. 1986. The human skeleton inforensic medicine. Springfield, IL: Charles C. Thomas.

Schwartz J. 1995. Skeleton keys: an introduction tohuman skeletal morphology, development, and analy-sis. New York: Oxford University Press.

MARK P. MOONEY

Department of Oral Medicine& Pathology, Anthropology,and Department of Plasticand Reconstructive Surgery,OrthodonticsUniversity of PittsburghPittsburgh, PennsylvaniaDOI: 10.1002/ajhb.20412

Human Body Systems. Edited by MichaelWindelspecht. 10 volumes. Westport, CT:Greenwood Press. 2004. $399.00 (cloth).

Human Body Systems is a 10-volume series,with each volume focusing on onemajor organsystem in the body. An organ system is definedas ‘‘ . . . a group of organs that physiologicallyfunction together to conduct an activity for thebody’’ (Kelly, 2004: p. ix). What makes thisseries unique is the comprehensive approachthat is presented for learning about each organsystem. The series identifies 10 organ systems:circulatory, digestive, endocrine, lymphatic,muscular, nervous and sense organs, repro-ductive, respiratory, skeletal, and urinary.Each volume generally begins with an over-

view of the anatomy, physiology, and develop-ment of each organ system, starting at thecellular level. The next area of each volumehighlights important historical discoveries inthe areas of anatomy andmedicine, and futureareas of research are discussed in a number ofthe volumes. The last section of each volume isfocused on injuries, disorders, and diseases ofeach body system.

The stated intent of the series is to providea ‘‘ . . . one-stop reference source for anyonewith an interest in the human body’’ (Kelly,2004: p. ix). The volumes range in contentfrom 8 to 18 chapters and in length from 172to 231 pages. Each volume is written by pro-fessional authors who specialize in present-ing scientific topics to general audiences.While it is beyond the scope of this reviewto discuss each of the 10 volumes, this reviewwill briefly discuss two of the volumes, TheMuscular System (Adams, 2004) and TheSkeletal System (Kelly, 2004), as representa-tive examples for the layout of the volumesin this series.

The volume entitled The Muscular System(Adams, 2004) is organized into 10 chapterscovering a variety of topics. The book beginswith a detailed, although not overly techni-cal, description of the three basic types ofmuscle tissue in the human body and thebasics of muscle contraction in Chapter 1.The physiology of energy use by muscles isdiscussed in detail in Chapter 2, and Chapter3 provides a very thorough description ofhow skeletal muscles adapt physiologicallyto exercise and how muscles grow as a resultof exercise. A description of muscular devel-opment is presented in Chapter 4. Chapter 5discusses early historical discoveries of mus-cle anatomy and physiology, and Chapter 8 isdevoted to muscular dystrophy. Chapter 9presents an overview of various diseasesand disorders of skeletal muscles, and thelast chapter presents a description of dis-eases of cardiac muscle.

The volume entitled The Skeletal System(Kelly, 2004) is organized into 18 chaptersand is probably the most comprehensive ofall 10 books. This book is organized intothree sections. The first section provides anoverview of the skeletal system and othergeneral information, the second sectionfocuses on the history and highlights ofresearch pertaining to the skeleton andorthopedics, and the last section is focusedon select diseases and disorders of the skele-ton. The book begins with an overview of the

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functions of the skeletal system and generalterminology in Chapter 1. Chapters 2 and 3provide a general overview of the bones ofthe skeleton that includes descriptions ofbone groups and individual bones. A briefand relevant overview of joint classificationand joint anatomy is presented in Chapter 4.Chapter 5 provides a brief, but thorough dis-cussion of bone growth and renewal, andembryology of the skeleton is discussed inChapter 6.

The chapters that highlight important his-torical discoveries in the areas of skeletalresearch and orthopedics provide verydetailed accounts with relevant images.Each chapter covers a different time period.Chapter 7 begins with what was knownabout bones and the beginnings of orthope-dics during antiquity. This is followed byimportant discoveries and what was knownabout bones and orthopedics through theDark Ages (Chapter 8), the renaissance(Chapter 9), the seventeenth and eighteenthcenturies (Chapter 10), the nineteenth cen-turies (Chapter 11), the twentieth century(Chapter 12), and the twenty-first century,with a look toward future research (Chapter13). Chapter 14 provides a brief overview ofinjuries to the skeleton and highlights com-mon sports injuries. In Chapter 15, briefdescriptions of all of the relevant joint dis-eases and disorders are presented, andChapter 16 presents a brief overview of dis-orders of the hands, feet, and back. Finally,Chapter 17 provides an overview of diseasesand disorders of the skeleton, and Chapter18 provides an overview of genetic and con-genital disorders.

As a general overview text that includes avariety of topics, it is to be expected thatcertain topics will be more thoroughly dis-cussed than others. However, consideringthat the series deals specifically with suchtopics as anatomy, physiology, and growthand development, a section providing a gen-eral overview of the gross anatomy of eachorgan system should be included at thebeginning of each volume. Moreover, thevolume on the muscular system did notinclude a section on the gross anatomy ofthe skeletal muscles. Although the firstchapter is titled ‘‘Anatomy of the MuscularSystem,’’ the chapter is actually an overviewof the three different types of muscle tissue(Adams, 2004). Considering the varioustopics that are discussed in this book, at thevery least, there should have been a section

with images that presented skeletal musclesor muscle groups and discussed their origins,insertions, and actions.

In any text dealing with anatomy, physiol-ogy, and development, there should benumerous high-quality and descriptive pic-tures that accompany the text. In particular,with a reference that is described as a ‘‘one-stop reference source’’ (Kelly, 2004: p. ix),the images are just as important as thedescriptions. For example, in the text deal-ing with the skeletal system, there couldhave been additional gross anatomy picturesof the skeleton that accompanied thedescriptions in the text. Many of the anat-omy pictures of the skeleton are simple linedrawings that omit important morphologicallandmarks on individual bones. Also, a num-ber of relevant pictures were not included.Although Chapter 3 provides a thoroughdescription of the bones of the lower limb, alabeled picture of the articulated bones ofthe lower limb is not included.Furthermore, a labeled picture of the poster-ior surface of the femur is included inChapter 2; however, the distal epiphysis is apoor representation. Chapter 15 provides abrief description of the anatomy of a synovialjoint that is relevant to understanding jointdiseases and disorders, but a labeled pictureof the features of a synovial joint is notincluded in the book.

Overall, this series is a worthwhile contri-bution to the field of human biology. It willserve as an informative and practical refer-ence for its intended audience of publiclibraries, secondary school and undergradu-ate college libraries, medical professionals,and anyone that is interested in learningabout the history of anatomy and medicine.The volumes are not overly technical and arepresented in a manner that is easy for indi-viduals who are not familiar with this mate-rial to follow and comprehend it.

The strength of the series is in providing areference that combines multiple topics foreach organ system into one comprehensivevolume. In addition, the comprehensive his-torical overviews of important anatomicaland medical discoveries are a highlight ofthis series. Each of the chapters that focuson these historical topics has numerous rep-rinted pictures to supplement the text. Eachvolume summarizes the important research-ers and milestones or discoveries of medicaland anatomical research for each organ sys-tem and traces these developments from

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antiquity to future avenues of research.Anyone that teaches anatomy and physiol-ogy, or is interested in anatomy and physiol-ogy, would find these sections very usefuland interesting.

LITERATURE CITED

Adams A. 2004. The muscular system. Westport, CT:Greenwood Press.

Kelley E. 2004. The skeletal system. Westport, CT:Greenwood Press.

JOHN J. SCHULTZ

Department of Sociology andAnthropologyUniversity of Central FloridaOrlando, FloridaDOI: 10.1002/ajhb.20413

Molecular and Cellular Exercise Physiology.Edited by Frank C. Mooren and KlausVolker. xþ 454 pp. Champaign, IL: HumanKinetics. 2005. $89.00 (cloth).

This is an excellent text on exercise andcell biology edited by two authorities on thistopic (i.e., Dr. Frank Mooren and Dr. KlausVolker). In general, this is a well writtenbook that serves as a reference text for grad-uate students, post-graduate exercise phy-siologists, and sports physicians.

The content of this text is divided into twological sections. The first section addressesissues related to molecular physiology, suchas cell structure and regulation of geneexpression. These 10 chapters are succinctbut contain most of the essential informa-tion required to understand the basics ofmolecular physiology. As in most editedtexts, the writing style and the level of scien-tific detail varies across the chapters.Nonetheless, in general, the first section ofthe text is well done.

Section two of the book contains chaptersthat focus upon the impact of exercise on thecell. These well-written chapters discusssuch topics as exercise-induced changes inthe cardiac myocyte, endothelium, and theskeletal muscle fiber. These chapters serveas excellent references for the graduate stu-dent or post-graduate exercise scientist.Similar to the first segment of the text, thelevel of scientific detail and writing style isvariable across these later chapters.

In summary, I highly recommend thisreference book on molecular and cellular

exercise physiology. The book provides anup-to-date summary on many aspects of cel-lular and molecular exercise physiology andwould be a useful desk reference for gradu-ate students, post-graduate exercise physiol-ogists, and sports physicians.

SCOTT K. POWERS

Department of Applied Physiologyand KinesiologyUniversity of FloridaGainesville, FloridaDOI: 10.1002/ajhb.20414

Man the Hunted: Primates, Predators, andHuman Evolution. By Donna L. Hart andRobert W. Sussman. xviiiþ 312 pp.Boulder, CO: Westview Press. 2005.$29.95 (cloth).

The topic of human evolution is a dyn-amic and controversial research field.Multidisciplinary approaches integratingknowledge from various sources are neededto unravel the origins and evolutionary pathof our own species. That is what D. Hart andR. Sussman do in this book. Man the Huntedis organized into 10 chapters and is accessi-ble and interesting to the lay reader. It intro-duces a wide variety of present and extinctpredators that make primates part of theirdiet. The authors describe and debate thecommon view of Man as the evolving hunterand present their own view of Man’s evolu-tion as an adapting prey by integrating fossilrecords and behavioral data from living pre-dator–prey interactions involving humanand nonhuman primates.

Chapter 1, ‘‘Just another item on themenu,’’ introduces the reader to the idea of‘‘Man the Hunted,’’ where the evolution ofman is molded by pressures upon them asprey. The authors stress the need for a com-bination of fossil records with the study ofliving species of primates to assess humanevolution. Chapter 2, ‘‘Debunking ‘Man theHunter’,’’ follows with descriptions of some ofthe relevant paleontological findings of earlyhominids, taxonomic revisions, and how thesehave changed and keep changing the scienti-fic paradigms of human evolution. They layout the basis for the ‘‘Man the Hunter’’ theoryand review and challenge the evidence.

Chapter 3, ‘‘Who’s eating whom?’’ startswith a comparison of predation on humansversus apes and includes a brief descriptionof what constitutes predator–prey interac-

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tions, stating the major theories, and coevolu-tion phenomena. Presented also are primatediversity and the current defining features ofthe primate order aswell the author’s study ofpredation on primates.

Chapter 4, ‘‘Lions and tigers and bears, ohmy!’’ features examples of notorious cases ofpredation on humans by lions, tigers, leo-pards, cougars and bears, and includesattack and killing strategies and the ances-try of modern big cats. Chapter 5, ‘‘Coursinghyenas and hungry dogs,’’ begins withreports of human predation by wolves inEurasia, describes wolf ancestors, and callsattention to the fact that this does not occurin North America, speculating on why thismay be consistent with the theory of ‘‘Manthe Hunted.’’ Current and prehistoric hye-nas are also presented as important humanpredators, and they end with a brief descrip-tion of predation by hunting dogs, dholes,and jackals.

Chapter 6, ‘‘Missionary position,’’ describespredation by reptiles (snakes, komodo dra-gon, and crocodiles) of primate prey andincludes an interesting description of huntingstrategies and habitats. It finishes, however,with a reference to a shark attack that seemsto be out of place in this book. As the authorsthemselves acknowledge, although impress-ive predators, sharks cannot be consideredan evolutionarily relevant predator ofprimates due to significant habitat differ-ences.

Chapter 7, ‘‘Terror from the sky,’’ as thename suggests, refers to aerial predators andfocuses on three raptor species: the crowned-hawk eagle, the harpy eagle, and thePhilipine eagle. Nocturnal birds of prey arealso discussed. Again, descriptions of ecol-ogy, hunting strategies, extinct species andfossil records of hominid predation follow.Gil-da-Costa et al. (2003) report a huntingstrategy for harpies and subsequent primateadaptive behavior that differ from what isdescribed in the book, but they agree withHart and Sussman that eagle predation wasanother driving force in hominid evolution-ary adaptation. The chapter concludesemphasizing the importance of predationupon hominid evolution and moves next tothe ‘‘hominid prey perspective.’’

Chapter 8, ‘‘We weren’t just waiting aroundto be eaten,’’ refers to the difficulties in recon-structing behavior from fossil records andstresses the need for studying living nonhu-man primates for the comparative perspective.

They list the main features and behaviorspotentially used by early hominids to avoidpredation and develop bipedalism. Theauthors refer to different features by whichmankind is thought to be unique, mention‘‘Man the user of language,’’ and contrast itwith the example of learned American SignLanguage by chimpanzees, in a way thatcould be interpreted as contradictive of ouruniqueness for this capacity. Later, under thesection ‘‘Daytime talk shows,’’ they describethe alarm call vocalizations present in severalprimate species and briefly speculate about theorigins of speech and language in a perhapsmisleading way (see Hauser, Chomsky, andFitch 2002). The various stages of a predatorattack are presented and discussed. Theauthors suggest basic tool use by living pri-mates and that the ancestors served a protec-tive measure more than did hunting.

Chapter 9, ‘‘Gentle savage or bloodthirstybrute?’’ starts by briefly describing the com-mon views and main contributors to the ideaof ‘‘Man the Hunter.’’ They focus on theorigins of this theory and discuss the beha-vioral traits considered to be human univer-sals, relating them to social Darwinism andsociobiology theories, ideas which they dis-pute. They do agree that there is a carnivor-ous tendency and that hunting strategies areshared by primates, but the authors do notbelieve that violent behavior and violenceoriginated as inherited traits, a view ex-emplified by the ‘‘demonic males’’ of R.Wrangham and D. Peterson (1996) and the‘‘selfish gene’’ of R. Dawkins (1989). Here,however, the authors take a controversialstand by reducing these theories to ‘‘Euro-Christian beliefs-based interpretations’’ and‘‘less careful science’’. The position is argu-able, but what is discouraging is that they donot present a more convincing alternative.The chapter ends with a reference toA. Zihlman’s work, exploring the femalerole in both current and ancestral humanand nonhuman primate societies.

Chapter 10, ‘‘Man the Hunted,’’ presentsthe authors’ view of how our ancestors mayhave lived. Data on fossil dentition records,typical diets, and physical adaptations to dif-ferent ecological niches are used to assesshow the australopithecines survived. Theauthors stress how adaptive behaviors areinfluenced by habitat constraints, and mon-keys provide valuable comparative modelsdue to their social and ecological similaritiesto early hominids. A summary of relevant

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antipredator strategies follows. Finally, theauthors finish with their creative, entertain-ing and useful vision of ‘‘a day in the life’’ ofour hominid ancestors, leaving the readerwith a lively picture of their theory.

The opposition set up between the selec-tive pressures and influences of the ecologyon Man as the hunter and Man as the hunteddistracts from the fact that, most likely dur-ing hominid evolution, the two go hand-in-hand and are complementary. However,directing their comments mostly to themedia, the popular view of human originsfocuses on Man as the powerful dominatinghunter. This is per se a scientifically incor-rect view. This book promotes the ‘‘otherside of the coin,’’ with Man as prey, in theevolutionary context of the associated selec-tive pressures.

LITERATURE CITED

Dawkins R. 1989. The selfish gene. Oxford: OxfordUniversity Press.

Gil-da-Costa R, Palleroni A, Hauser MD, Touchton J,Kelley JP. 2003. Rapid acquisition of an alarmresponse by a neotropical primate to a newly intro-duced avian predator. Proc R Soc London B 270:605–610.

Hauser MD, Chomsky N, Fitch WT. 2002. The faculty oflanguage: what is it, who has it, and how did it evolve?Science 298:1569–1579.

Wrangham R, Peterson D. 1996. Demonic males: apesand the origins of human violence. New York:Houghton Mifflin Co.

Zihlman A (with N Tanner). 1976. Women in Evolution,Part I. Innovation and selection in human origins.Signs: J Women Culture Soc 13(1):558–608.

RICARDO GIL-DA-COSTA

Laboratory of Brain and CognitionNational Institute of Mental Health, NIHBethesda, MarylandDOI: 10.1002/ajhb.20415

The Changing Role of the Embryo inEvolutionary Thought: Roots of Evo-Devo.By Ron Amundson. xiv þ 280 pp. NewYork, NY: Cambridge University Press.2005. $75.00 (cloth).

Neo-Darwinism coined a particular pictureof evolution: the explanation of adaptationand speciation as the central goals of evolu-tionary theory and natural selection as theonly directing cause in evolution. Recently,evolutionary developmental biology (evo-devo) became an independent field, pointingout that that there are other important goalsof evolutionary biology: the explanation of the

origin of morphological novelties and bodyplans, and an account of evolvability. Neo-Darwinism has viewed the study of develop-ment as being irrelevant to evolutionary biol-ogy, but evo-devo biologists argue thatdevelopmental and morphological featuresrather than natural selection are crucial cau-sal factors in these new evolutionary explana-tions, yielding a more powerful explanatoryframework and a broader understanding ofthe mechanisms of evolution. Apart from con-temporary biology, neo-Darwinism has alsostrongly influenced how the history of biologyhas been viewed by biologists, philosophers,and historians. This led to a distorted pictureof 19th-century biology, and during the lasttwo decades historians of biology have startedto correct this view of history. Although theemergence of evo-devo and recent develop-ments in historical scholarship are largelyindependent, philosopher of biology RonAmundson brings these two trends togetherin this volume. Amundson distinguishesbetween functionalist and structuralistapproaches in the history of biology; the for-mer emphasizes the explanation of adaptationand the organism–environment relationship,and the latter stresses the explanation of formand the internal structure and development ofindividuals. Amundson’s central target iswhat he calls ‘‘Synthesis Historiography,’’i.e., the use of the conceptual framework ofneo-Darwinism to interpret the history of biol-ogy. The first half of Amundson’s book con-cerns the 19th century and debunks varioushistorical misinterpretations due to SynthesisHistoriography by using a structuralist or evo-devo perspective of history to uncover impor-tant historical facts that former approacheshave left out. The second half concerns the20th century, addressing how developmentdropped out of evolutionary theory and howSynthesis Historiography (SH) emerged in thefirst place and discusses the differencesbetween contemporary functionalist andstructuralist approaches.

A central element of SH is the distinctionbetween typological thinking and populationthinking. It forms the basis of the assumptionthat pre-Darwinian biologists were typologistsor essentialists who explained species fixitybased on the idea that species have immutableessences. As Amundson shows in line withrecent historical scholarship, this is simply amyth. Species fixism gained acceptance just acentury before Darwin, and its original accep-tance was scientifically progressive, because

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naturalists showed that species do not wildlytransmute across taxa. As it turns out, speciesfixity was not explained based onmetaphysicalideas, such as types or essences, but withrecourse to empirical assumptions. Pre-Darwinian, ‘‘idealistic’’ morphology has alsobeen viewed in the grip of typological thinking.However, the notion of types is actually scien-tifically legitimate and does not conflict withpopulation thinking. Population thinking isabout the variation within species, and thenotion of a type is about the relation betweenspecies. Higher taxa, in particular phyla, havea type, but species were not viewed as having atype. The advent of phylogenetic systematicspromoted taxa characterized by synapomor-phies. This is what pre-Darwinian, ‘‘typologi-cal’’ biology was getting at. SH commentatorsoften regard historical beliefs in continuitybetween groups as indicators of progressive-ness in biological thought, but Amundsonmakes clear that a crucial step toward evolu-tionary theory was the recognition of charac-teristic discontinuities between extant species.The 18th-century naturalists usually assumedthat the various extant species form a contin-uous or seamless fabric of nature and that thetaxonomic system was a mere human contri-vance, and it was an achievement of early19th-century taxonomy to view the hierarchi-cal structure of taxonomic groups as reflectingreal patterns existing in nature. Research incomparative anatomy and embryology sup-ported this view by introducing the homologyconcept and finding homologies across largertaxa. Morphological relations were viewed asreflecting true relations between biologicalgroups, preparing theway for viewing the hier-archical structure of taxa as reflecting the phy-logenetic tree of nature.

SH tended to lump together two differentpre-Darwinian traditions based on thenotion of typological thinking: natural theol-ogy and idealistic morphology. The naturaltheologians were functionalist, viewing spe-cies as independently created by God so as tobe perfectly adapted to its ecological niche.However, as Amundson discusses in detail,the idealistic morphologists were structural-ists who stressed morphological relationsbetween species and criticized the adapta-tionist natural theologians for not beingable to account for homologies. According toSH, idealistic morphologists viewed morpho-logical types as blueprints in the mind ofGod. Some pre-Darwinian morphologists,such as Richard Owen, clearly expressed evo-

lutionary ideas. In any case, morphologicalaccounts could use a nonphylogenetic notionof a type as a real explanatory concept with-out invoking divine creation. Idealistic mor-phologists viewed types as reflecting law-likerelationships between organismal forms.Amundson points out that it is legitimatethat morphologists did not yet have anaccount of the real (phylogenetic) nature oftypes because there are many episodes in thehistory of science where scientists are justi-fied in invoking a law while still lacking areal understanding of its underlying causalbasis. Darwin’s main enemies were, in fact,the creationist natural theologians, and inthe Origin of Species Darwin explicitly reliedon the well-known results of idealistic mor-phology. The existence of homologies andwhat Darwin called the ‘‘Unity of Type’’ sup-ported his crucial doctrine of common ances-try. Some SH commentators stressed Darwin’sideas of natural selection but ignored the phy-logenetic–morphological aspects of Darwin’swork or—being puzzled how Darwin couldmake favorable remarks about the allegedcreationist morphologists—blatantly misinter-preted these passages of the Origin.

Amundson discusses the influential tradi-tion of 19th-century evolutionary morphol-ogy (Gegenbaur, Haeckel), which was in thebusiness of establishing phylogenetic treesand studying morphological evolution (suchas the origin of tetrapod limbs) based on howmorphologies and ontogenies change in thecourse of phylogeny. Some SH commenta-tors have viewed this as mere descriptionand the telling of historical narratives.Contemporary phylogenetic approaches aremore likely to view 19th-century morphologyas real explanation, and many evo-devo biol-ogists view the tradition of evolutionarymorphology as an important forerunner oftheir approach; extant forms and the originof morphological novelties are explained onthe basis of how ontogenies have changedduring phylogeny.

In the 19th century, evolutionary considera-tions were tightly connected to development.Heredity was viewed as a part of development.In the second part of his book, Amundsondescribes how the emergence of classical genet-ics in the 20th century led to a redefinition of‘‘heredity’’ so that it referred to the transmis-sion of features but excluded development,leading to the split between embryology andgenetics, where the latter but not the formerbecame part of the Modern Synthesis.

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Page 8: The changing role of the embryo in evolutionary thought: Roots of evo-devo

Amundson also examines how the essentialiststory about pre-Darwinian biology originatedin the late 1950s. It is embarrassing that thesehistorical claims were made without a realstudy of the history. Apart from the distinctionbetween typological versus population think-ing, neo-Darwinians developed other dichoto-mies (e.g., proximate vs. ultimate causes).These conceptual schemes were used both tointerpret the history of evolutionary biology inthe sense of neo-Darwinism and to argueagainst contemporary critics of the ModernSynthesis. In the 1980s, they came to be usedto reject structuralist critiques by arguing thatdevelopmental considerations have no bearingon evolutionary theory. Amundson describesthese more recent debates. Neo-Darwinianshave a ‘‘residual’’ concept of homology thatviews homology as the remnants of ancestralfeatures that have not been modified by nat-ural selection. Evo-devo biologists have a‘‘developmental’’ concept of homology thatviews homologues as building blocks of mor-phological organization that causally figure indevelopment and influence morphological evo-lution, thereby reviving ideas from the 19th-century morphology. Neo-Darwinians focus onthe explanation of adaptation, viewing popula-tions as the entities that evolve, while evo-devobiologists focus on the explanation of the evo-lution of form, viewing ontogenies as what

evolves. Modern practitioners of evo-devoassume that selectionist and developmentalmodes of explanations are perfectly compati-ble. Amundson is, in fact, impressed by theseeming incommensurability between func-tionalist and structuralist perspectives, statingthat it has yet to be shown that bothapproaches can really be united.

This volume gives a masterful account andought to be read by everyone who is inter-ested in evolutionary biology. Amundsonreminds us that any contemporary pictureof biology and its history should never betaken as complete and unbiased. His discus-sion also indicates that certain moderndebates such as the debate about adaptation-ism versus developmental constraints hadforerunners (natural theology vs. idealisticmorphology), although they occurred in aseemingly different context (such as pre-Darwinian biology). Modern debates are notjust about what we take them to be; a con-sideration of the history of biology is rele-vant for our current situation.

INGO BRIGANDT

Department of Historyand Philosophy of ScienceUniversity of Pittsburgh

Pittsburgh, PennsylvaniaDOI: 10.1002/ajhb.20416

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