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Architecture in Living Structure

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Architecture in Living Structure Architecture in Living Structure Edited by G.A. Zweers and P. Dullemeijer Reprinted from Acta Biotheoretica Volume 34, Nos. 2-4 1985 MARTINUS NIJHOFF/DR W. JUNK PUBLISHERS ,L a member of the KLUWER ACADEMIC PUBLISHERS GROUP II DORDRECHT I BOSTON I LANCASTER 'liliiii Distributors for the United States and Canada: Kluwer Academic Publishers, 190 Old Derby Street, Hingham, MA 02043, USA for the UK and Ireland: Kluwer Academic Publishers, MTP Press Limited, Falcon House, Queen Square, Lancaster LA1 1RN, UK for all other countries: Kluwer Academic Publishers Group, Distribution Center, P.O. Box 322, 3300 AH Dordrecht, The Netherlands Library of Congress Cataloging In Publication Data Main entry under title: Architecture 1n living structure. Proceedings of • symposium held 3/15-17/84 in LochmUhle, AU5senste lIe der Senckenbergischen Naturforschenden Gesellschaft, Frankfurt am Main. lncludes bibliographies. 1. Morphology (Animals)--Congresses. L Zweers. G. A. II. Dullemeijer, P. III. Senckenbergische Naturforschende Gesellschaft. Aussenstelle LochmUhle. QL799.A73 1985 591.4 85-21492 ISBN-13: 978-94-010-8787-2 ISBN-13: 978-94-010-8787-2 e-ISBN-13: 978-94-009-5169-3 001: 10.1007/978-94-009-5169-3 Copyright © 1985 by Martinus Nijhoff/Dr W. Junk Publishers, Dordrecht. Softcover reprint of the hardcover 1st edition 1985 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, mechanical, photocopying, recording, or otherwise, without the prior writ­ ten permission of the publishers, Martinus NijhofflDr W. Junk Publishers, P.O. Box 163, 3300 AD Dordrecht, The Netherlands. [3] 109 Proceedings of a symposium on Architecture in Living Structure The primary goal of the symposium was to arrive at a comparative analysis of various methodological pathways. The symposium was organized by W.F. Gutmann, P. Dullemeijer and G.A. Zweers and was held from March 15-17, 1984 in the Lochmuhle, Aussenstelle der Senckenbergischen Naturforschenden Gesellschaft, Frankfurt am Main. This issue is edited by G.A. Zweers and P. Dullemeijer. Contributions to the symposium R. McNeill Alexander: The legs of ostriches (Struthio) and moas [59] 165 (Pachyornis) . J. Bereiter-Hahn: Architecture of tissue ce11s. The structural basis [33] 139 which determines shape and locomotion of cells. Tj. de Cock Buning: Qualitative and quantitative explanation of the [87] 193 forms of heat sensitive organs in snakes. Tj. de Cock Buning, E. Otten & P. Dullemeijer: Some methodological [171] 227 remarks. P. Du11emeijer: Diversity of functional morphological explanation. [5] III M. Grasshoff: On the reconstruction of phylogenetic transformations: [43] 149 The origin of the arthropods. U. An der Heiden, G. Roth & H. Schwegler: Principles of self- [19] 125 generation and self-maintenance. J.W.M. Osse: Jaw protrusion, an optimization of the feeding apparatus [113] 119 of Teleosts? E. Otten: Proportions of the jaw mechanism of cichlid fishes: changes [101] 107 and their meaning. D. S. Peters: Mechanical constraints canalizing the evolutionary [51] 157 transformation of tetrapod limbs. W.E. Reif, R.D.K. Thomas & M.S. Fischer: Constructional morphology: [127] 233 the analysis of contraints in evolution. G. Roth & D.B. Wake: Trends in the functional morphology and sensori- [69] 175 motor control of feeding behavior in salamanders: an example of the role of internal dynamics in evolution. G.A. Zweers: Greek classicism in living structure? Some deductive [143] 249 pathways in animal morphology. Additional symposium presentations W.J. Bock: Parts and wholes in morphology: a reductionistic and inte­ grative approach to holism. W.J. Bock: Adaptations and other structural components to fitness. W.F. Gutmann: Organisms as energy converting systems. Symposium participants R. McNeill Alexander, J. Bereiter-Hahn, J.C. Vanden Berge, W.J. Bock, Tj. de Cock Buning, P. Dullemeijer, A. Grosse-Lohmann, M. Grasshoff, W.F. Gutmann, U. An der Heiden, W. Maier, J.W.M. Osse, E. Otten, D.S. Peters, G. Rehkamper, W.E. Reif, E. Reisinger, G. Roth, K. Vogel, M. Weingarten, E. Zeiske, U. Zeller, G.A. Zweers. Page numbers in breckets refer to pagination of book edition DIVERSITY OF FUNCTIONAL MORPHOLOGICAL EXPLANATION PIETER DULLEMEIJER Zoologisch Laboratorium, Rijksuniversiteit Leiden, The Netherlands An introduction to the 19th Lochmuhle conference on 'Architecture in living structure', organized by P. Dullemeijer, W.F. Gutmann and G.A. Zweers, held from March 15-17, 1984 in the Aussenstelle des Forschungs­ instituts Senckenberg der Senckenbergischen Naturforschenden Gesellschaft, Frankfurt am Main. 1. INTRODUCTION Although functional morphologists form a minority group amongst biologists, there is, among them, a wide diversity of opinion in approach, aim, attitude and procedure, resulting in controversial questions and answers. This Lochmuhle conference was organized to elucidate these differences with the aim to obtain a soil for a mutual understanding. A first pre­ requisite to reach this aim is an analysis of the various motives to do functional morphology and the applied methodologies. Therefore the participants were asked to emphasize the following aspects: 1. to indicate the aspects intended to be explained in a functional morphological case study; 2. to clarify the type of explanation; 3. to describe the subsequent methodological steps; 4. to focus on the following questions: a. how the conclusion depends on the first step of abstraction; b. how the conclusion changes if the selected part is considered a member of an integrated wider system; c. how the conclusion and the procedure changes when temporal aspects, such as ontogeny and evolution, are introduced. 112 [6] .-----:-------~--- ,7....:,,-------- , , , , \ ............ " ',-----'-, ....,::.---­ - ----;;? , " " ". Figure 1. Diagram of the bony elements in a viperid snake. Drawn lines: opened mouth; stippled lines: closed mouth. be, braincase; ec, ectopterygoid; lj, lower jaw; m, maxilla; pf, poison fang; pt, pterygoid; vc, vertebral column. 1.1. An example As an introduction I shall demonstrate the various possibilities of explanations on a very simple, almost classical example, rather than present an abstract discourse on philosophy, procedure and science theory [5]. There is a small bone in the upper jaw of a viperid snake (viper, rattlesnake), the ectopterygoid, which transfers the force and movement of the protractor and levator pterygoid muscles to the maxilla (fig. 1). As a result the latter is rotated so that the poison fang is erected. The ectopterygoid is retracted by the adductor pterygoideus muscle. This mediation of protractor and retractor movement during strike and swallowing is its function, its biological role, in a total chain and pattern of structures and movements. To fulfil this role the bone has to perform certain activities; one of them is to resist the strain put on it. Evenly distributed strain resistance of a specified bone is obtained by a specific shape, given the kind of material. This statement is reasonable in view of general experience from engineering science, but it needs of course an accurate analysis in each particular case. This analysis is provided by the deductive method, by which the shape is deduced from the functional [7) 113 FORCES IN A HORIZONTAL. PLANE • r===~:: u _____ - -j 1 FORCES IN A VERTICAL PLANE ----.-~ + ~- -----~ \;______ .-====\=---,_/=::::::::::::::J ~ ~L:=-==---:=---=-==----, \ / 7~~' ~~ J:,~ Figure 2. Mechanical model for the explanation of the shape of the ecto­ pterygoid in Vipera berus. Left row the construction of the model for the activities of the muscles in retraction (a) and protraction (b) in the horizontal plane. In (c) the necessary shape for resisting the bending and compression for the retractor (m.adductor pterygoideus), in (d) that for the protractors (m.protractor et levator). As both systems work alternately, in the addition both figures can overlap (d). In (f), (g) and (h) the same in the vertical plane. Combining (e) and (h) results in the three-dimensional model (i), which is compared to the actual shape represented by cross-sections. demands by means of a general theory (fig.2). In this deduction the following steps are taken (6). I. Of all functional demands the resistance against muscle force is qelected. 2. The position and the length of the ectopterygoid in the construction is taken as a given feature. 3. The type of construction is determined. 4. It is supposed that the elasticity of bone is known and that the material is evenly distributed. 5. It is supposed that the laws of static mechanics are applicable. 6. A model of the shape is made by applying the principle of optimal design. 7. The model of the shape is compared to the actual shape; the model is accepted or refuted. 8. If accepted it is concluded that the shape is explained from the functional demand (i.e., the activity of the bone) by means of the theory of statics under the mentioned boundary conditions. In formula S = f(A) in which S = shape, A = activity and f the explaining theory. This is the general deductive-nomological formula of Hempel and 114 [8] others [9]. In Popper's system the falsification of the models is emphasized [12] , whereas in other systems confirmation is accepted, as is common in the majority of natural sciences. 2. TYPES OF EXPLANATION If we now generalize this formula in Hempel's sense as y = f(x), in which y is the phenomenon to be explained (the explanandum) and f(x), the explanans, of which f
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