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Part II - Organ structure, function, and behavior
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Book:
- Shaping Primate Evolution
- Published online:
- 10 August 2009
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- 20 May 2004, pp 97-98
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Summary
My own earliest studies tried to understand functional adaptation in the locomotor system, primarily of bones and joints, through morphometrics. As a preliminary, however, I did attempt to find out what I could of the muscles that moved the bone–joint unit. In those days all we did was dissect muscles and measure them through relative lengths, directions of pull, relative weights, and frequencies of (what were called in those days) muscular anomalies. Limited though such studies were, they were incredibly time-consuming. In all, I dissected 52 shoulders representing 28 primate species, 145 arms and forearms representing 27 primate species, and 167 hips and thighs in 33 primate species. That took many years. Yet even such primitive data provided useful initial information about muscles in relation to the respective bone–joint units.
But it was always clear to me, and I wrote about it without ever doing it, that better muscular studies would be necessary. Such studies would need to understand much more about muscular architecture than simply relative muscle weights, much more about muscle activity and functions through studies of living muscle than just anatomical inferences, much more about movements, postures, and overall behaviors of the living animals than just simple classifications of locomotion.
However, I certainly did not, in those early days, envisage the possibility of studies of the biomechanics of the jaws and teeth and, further, the biomechanics of food being masticated by them. And I knew nothing at all of functional adaptations in the brain.
A few examples of such investigations follow in this section. Again, my students and colleagues have gone so much further than was possible for me. Yet notwithstanding, stimulated by them, I have been able in these latter years to enter some of these areas myself.
20 - Postscript and acknowledgments
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- By Charles E. Oxnard, School of Anatomy and Human Biology, University of Western Australia, 35 Stirling Highway, Crawley, WA, 6009 Australia
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Book:
- Shaping Primate Evolution
- Published online:
- 10 August 2009
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- 20 May 2004, pp 415-419
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Summary
One usually thinks first of the well-known senior individuals who have been responsible for initiating one's career. My initial stimulus came, however, from the enthusiasm of a relatively unknown person. He was the headmaster of the tiny primary school I attended in a small village in Scotland at the outbreak of the 2nd World War. Knowing that education in Scotland was very classical: English and Mathematics, even Latin and Greek in those days, but no science to speak of, he understood, somehow, that this small boy was interested in science. He introduced me to the ideas of Wegener, Goethe, D'Arcy Thompson and Solly Zuckerman when I was nine years old, two of them, of course, not in the original.
As a result, I may be the only person in the world who knew about the movements of the continents but who did not know that Wegener's ideas were not accepted for almost half a century. By the time I reached university in 1952, the idea was center-stage as plate tectonics. I could not understand what all the excitement was about. I had always known it was so.
Likewise, I understood very well the idea that the skull was simply a series of fused vertebrae. It made sense to me. I did not know that Goethe had it wrong until I later came to read Gavin de Beer's tome on the vertebrate skull.
19 - Design, level, interface, and complexity: morphometric interpretation revisited
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- By Charles E. Oxnard, School of Anatomy and Human Biology, University of Western Australia, 35 Stirling Highway, Crawley, WA, 6009 Australia
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Book:
- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp 391-414
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Summary
Origins of morphometrics
The origin of morphometrics lies in the burst of methodological creativity that predates the availability of relevant datasets requiring such methods. A seminal decade saw developments from Hotelling (1931), Wilks (1935), Bartlett (1935), Fisher (1936), Mahalanobis (1936), and others. And the next 20 years provided the extensions of the methods by Rao (1948), Yates (1950), Kendall (1957), and others. Most of these investigations concentrated on developing the methods. The actual data examined during these developments were so few that, though they permitted analysis by manual techniques, they were too restricted to allow examination of real biological problems. One well-known dataset was Anderson's measurements of four variables, the lengths and breadths of sepals and petals, taken on 50 specimens each of three groups of iris. These data were used by Fisher (1936) in the development of the techniques, and by many other investigators since, including myself (e.g., Oxnard, 1973, 1983/84), both for development and for checking.
Of course, the antecedents of morphometrics came from even earlier times (e.g., Galton, 1889; Pearson, 1901) and even the nineteenth and late eighteenth centuries (e.g., Adanson, 1763; Quetelet, 1842). But it was not until the second half of the twentieth century that morphometric methods could be applied to large datasets (many variables, many specimens, many groups) and could be aimed at examining real rather than exemplar anthropological problems (e.g., Trevor, 1955; Ashton et al., 1957, 1975, 1976, 1981; Oxnard, 1967; Howells, 1973).
Part I - Craniofacial form and variation
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Book:
- Shaping Primate Evolution
- Published online:
- 10 August 2009
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- 20 May 2004, pp 9-10
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The study of craniofacial form and variation has always been one of the most important areas for those interested in shaping primate evolution. Skulls were the most frequently collected specimens in museums. Skull parts, especially teeth, are most frequently found in the fossil record. Skulls and teeth are easily examined in the living. The bones of the face allow some estimation of how their owners appeared. Appearance and change in appearance as produced by medical and dental technologies have profound effects upon individual well-being. All these are good reasons why this is one of the most critical of anatomical regions.
At the same time, however, skulls, faces, jaws, and teeth are the most complex region of the body. More, perhaps, than in any other region, do a number of completely different functions have to be integrated in its structure. The genetics underlying cranium, face, jaw, and teeth are even now not well known and clearly far more complicated than the postcranium. The development and growth of the head depends upon complex mechanisms and processes, many of which have only been elucidated in the last two decades. In evolutionary terms, the “head problem” in chordates, reflecting at the same time both very ancient and very recent elements, has always been more difficult to understand than, say, the equivalent trunk problem or limb problem (which problems do not even rate quotation marks).
Part IV - Theoretical models in evolutionary morphology
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Book:
- Shaping Primate Evolution
- Published online:
- 10 August 2009
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- 20 May 2004, pp 279-280
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Though I had always been interested in the use of mathematical methods of one kind or another, in the earlier days these interests were almost always for data analysis. The models of function with which we worked in the early days were simplistic in the extreme, scarcely deserving of the term “model” – in fact they were little more than functional anatomical inferences. At later stages I used models for assessing mechanical efficiency, but these mainly employed what is now a quite old-fashioned technique – photoelastic analysis – and they were extremely limited – e.g., to two dimensions only, and only to isotropic situations. Only much more recently, in collaboration with others (e.g., O'Higgins), have I come to use better modeling methods (such as finite elements).
However, in a completely surprising way, through a stimulus applied by Sydney Brenner and his invitation to present a model of mtDNA evolution (which I did not do) at a workshop hosted by the International Institute for Advanced Studies in Kyoto, I have come to be involved in true evolutionary modeling (mimicking species evolution and individual lineages) with Dr. Ken Wessen. Though not included as a section in this book, Dr. Wessen's work (in which I have been pleased to share) is reported in my own final chapter.
At this point, however, it is an especial pleasure to recognize, through the following sections, kinds of modeling that I scarcely envisaged in those earlier days. Thus the following chapters on modeling the origins and mechanics of bipedalism, and the mechanics of mastication, are extremely relevant.
Part V - Primate diversity and evolution
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Book:
- Shaping Primate Evolution
- Published online:
- 10 August 2009
- Print publication:
- 20 May 2004, pp 351-352
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Once again, though my earlier work was clearly aimed at understanding functional adaptations in specific bone–joint–muscle units, it also equally clearly led into interests in primate diversity and evolution. I was not, of course, a primate taxonomist, never having had the requisite training. I never worked in field situations (being allergic to high temperatures, heavy rainfall, high altitudes, mosquitoes, leeches, etc). I therefore never participated in field observations of living species or in field discoveries of fossils. And though I was never formally educated in mathematics and statistics, and never capable myself of making advances in them, I was always a user who was interested in how such methods could be applied to data. Especially was I interested in the kinds of questions that the above methods and data might answer.
I have thus remained enormously interested in all such studies carried out by others – indeed, the data of others were essential to some of the investigations that I myself made. Mainly working with colleagues, however, I may have been perhaps the first to apply full multivariate statistical analyses to the data of field observation, of the niche. Likewise, through colleagues and students, I may have been amongst the first to use morphometric methods as tools to go beyond the data themselves, to seek correlations with behavior, with the niche, with development, and with evolution. The following chapters on the niche, on cladistics, and on the development of morphometrics itself carry all this so much further. These studies (and many others not represented in this book) go so very far beyond what I originally envisaged.
Part III - In vivo organismal verification of functional models
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Book:
- Shaping Primate Evolution
- Published online:
- 10 August 2009
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- 20 May 2004, pp 227-228
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Though it would appear from my early work that I primarily used anatomical inference for the “functional” explanations for the morphological, especially morphometric, adaptations that I saw in my studies, I nevertheless was always interested in the “real” biomechanics, interested in testing, that is, the hypotheses resulting from anatomical inference. The best that we could do at the time for the shoulder was to rely on the pioneering electromyographic studies on the human shoulder carried out by Inman, Saunders, and Abbott during the Second World War. Doing electromyography on nonhuman primates seemed such a long shot in those days.
Yet in fact my first research grant (from the US Public Health Service as it was called then), awarded in 1962 while I was still in the UK, aimed in part to design an implantable telemetric device for recording electromyographic and strain-gauge information from freely moving primates. Stanley Salmons was the research fellow employed on that grant for that purpose. But of course, only a few weeks' reading was enough to demonstrate that we could not do it – it just was not possible to make the device small enough for that purpose in those days. Stanley Salmons, however, did go on to design the “buckle transducer'” a first device for measuring tension in tendons, so all was not lost. He is today Professor of Biomedical Engineering at the University of Liverpool and has been engaged in these latter years in studies making the latissimus dorsi muscle into an adjunct pump in cases of cardiac insufficiency.
17 - From optical to computational Fourier transforms: The natural history of an investigation of the cancellous structure of bone
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- By Charles E. Oxnard, The University of Western Australia
- Edited by Pete E. Lestrel, University of California, Los Angeles
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- Book:
- Fourier Descriptors and their Applications in Biology
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- 14 September 2009
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- 13 May 1997, pp 379-408
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Introduction
The remarkable patterns exhibited by cancellous bone, and the likelihood that it is related to mechanical stress and strain, have been known for a long time (e.g., Todd and Bowman, 1845; Wyman, 1867; Wolff, 1892). During most of that period, however, the nature of the cancellous patterns themselves remained based upon visual impressions of bone sections and later, bone radiographs (e.g., Thompson, 1917; Murray, 1936; Evans, 1957). It was not until after the middle of the present century that the idea of characterizing these complex bony patterns with Fourier analyses came about.
In 1968, J. C. Davis showed me how useful optical Fourier transforms could be in the analysis of thin sections of rocks (Davis, 1970). In so doing he put into my mind the idea that the method might be excellent for the study of sectional and radiographic information in bones. Using his specialized equipment, we produced the first transforms of cancellous architecture in sections of human vertebrae (Oxnard, 1970a; 1970b; 1972a; 1972b; 1973). The 1973 volume may have been the first time that a Fourier transform was figured on the cover of a book.
Later H. C. Pincus (e.g., Power and Pincus, 1974) demonstrated to me the Rank Image Analyser 2,000 used for similar purposes.