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Holger Preuschoft

Publications and source records attributed to Holger Preuschoft.

7 recordsLinked to original sources

Finite-element model construction for the virtual synthesis of the skulls in vertebrates: case study of Diplodocus.

The measurement of strains in real skulls is an inductive method that yields information about the stresses occurring in the a priori existing shape. In contrast, the approach taken here to determine the relationship between skull function and skull shape applies Wolff's law through a deductive technique of structure synthesis. This article describes the application of this method in the exact virtual synthesis of a sauropod skull, e.g., Diplodocus longus Marsh from Wyoming. An unspecific homogeneous solid is first constructed, giving the stresses ample volume to spread between points of force application and constraint. ANSYS 7.0 is used to form 10-noded tetrahedral finite elements with a maximum of 130,000 nodes. The initial conditions are the functional spaces for the eye openings, muscle forces, and the placement of the dental arcade, including assumed bite forces. Enforcing equilibrium of forces, the primary 3D stress flows in each load case are summarized by a physiological superposition, which accumulates the highest value of stress in each finite element. If the stress free parts are eliminated and the summarized stress flows are maintained, a reduced model appears, which is very similar to the real skull. This reduction of shape can be repeated iteratively and leads to a more exact form. Changes in the form of the dental arcade, its position relative to the braincase, the origins of muscles, or the height of the face lead to models that clearly resemble morphological differences between genera. The synthesis of a skull in this way demonstrates the direct correlation between functional loading and the biological structure and shape and can be used to test hypotheses regarding the relationship between structure and function during skull evolution.

Animals↗

Functional shape of the skull in vertebrates: which forces determine skull morphology in lower primates and ancestral synapsids?

In order to determine the extent to which the shape of the synapsid skull is adapted for resisting the mechanical loads to which it is subjected, block- or simple plate-shaped finite-element models were constructed and loaded with external muscle and bite forces in locations estimated to resemble points of application of these forces. These 2D or 3D finite-element models were iteratively loaded and modified by removing elements that experience only low stresses, and the resulting morphologies of the models were compared with fossil skulls of synapsids and the skulls of extant mammals. The results suggest that the stress flows in these unspecific models are very similar to the arrangement of bone material in real skulls. Morphological differences between taxa depend on a few a priori conditions: length and position of the tooth rows in relation to the braincase, arrangement of muscles, position of the orbits, and position of the nasal opening. Given these initial conditions, finite-element analysis consistently reveals the close similarity between stress flows and real skulls. The major difference between mammal-like reptiles and primates is the size of the braincase. This difference accounts for most of the morphological divergence. The postorbital bar seems to be a constructional element of the skull, rather than a means to protect the eyes. The skull shapes of higher primates are determined mainly by masticatory forces and less by external forces acting on the head. This study demonstrates the utility of finite-element modeling for testing hypotheses regarding relationships between form and function in vertebrate skulls.

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A biomechanical approach to craniofacial shape in primates, using FESA.

The head of a land-living vertebrate is exposed to the forces of acceleration, in particular the permanent earth acceleration (= gravity) and the muscle-generated bite and chewing forces. In mammals, at least, the latter seem to play the dominant role. Bite forces are applied to the teeth and close the circle of forces by passing through the facial skeleton to the insertions of the mandibular adductors. With the aid of three-dimensional Finite Element Systems Analysis (FESA), the stress flows in homogenous bodies are investigated, whereby the braincase, the orbits and the nasal channel are taken as preconditions. The muscle insertions are varied systematically. The resulting stress flows in all cases turn out to be very similar to the bony structures of a skull. Little or not stressed parts of the available homogenous body indicate the external surface or hollow spaces (= sinuses) inside the skull. The possible applications of forces (i.e., the forms and positions of the dental arcade in relation to the braincase) determine the pathways along which the forces are transmitted. It seems that the factors mentioned above as preconditions represent the selective pressures exerted by the lifestyle of the animal and its environment (= ecological conditions). The results that can be obtained by our deductive approach in comparison to inductive, experimental procedures are discussed briefly.

Animals↗

Biomimetic robotics should be based on functional morphology.

Due to technological improvements made during the last decade, bipedal robots today present a surprisingly high level of humanoid skill. Autonomy, with respect to the processing of information, is realized to a relatively high degree. What is mainly lacking in robotics, moving from purely anthropomorphic robots to 'anthropofunctional' machines, is energetic autonomy. In a previously published analysis, we showed that closer attention to the functional morphology of human walking could give robotic engineers the experiences of an at least 6 Myr beta test period on minimization of power requirements for biped locomotion. From our point of view, there are two main features that facilitate sustained walking in modern humans. The first main feature is the existence of 'energetically optimal velocities' provided by the systematic use of various resonance mechanisms: (a). suspended pendula (involving arms as well as legs in the swing phase of the gait cycle) and matching of the pendular length of the upper and lower limbs; (b). inverted pendula (involving the legs in the stance phase), driven by torsional springs around the ankle joints; and (c). torsional springs in the trunk. The second main feature is compensation for undesirable torques induced by the inertial properties of the swinging extremities: (a). mass distribution in the trunk characterized by maximized mass moments of inertia; (b). lever arms of joint forces at the hip and shoulder, which are inversely proportional to their amplitude; and (c). twisting of the trunk, especially torsion. Our qualitative conclusions are three-fold. (1). Human walking is an interplay between masses, gravity and elasticity, which is modulated by musculature. Rigid body mechanics is insufficient to describe human walking. Thus anthropomorphic robots completely following the rules of rigid body mechanics cannot be functionally humanoid. (2). Humans are vertebrates. Thus, anthropomorphic robots that do not use the trunk for purposes of motion are not truly humanoid. (3). The occurrence of a waist, especially characteristic of humans, implies the existence of rotations between the upper trunk (head, neck, pectoral girdle and thorax) and the lower trunk (pelvic girdle) via an elastic joint (spine, paravertebral and abdominal musculature). A torsional twist around longitudinal axes seems to be the most important.

Adult↗

Mechanisms for the acquisition of habitual bipedality: are there biomechanical reasons for the acquisition of upright bipedal posture?

Morphology and biomechanics are linked by causal morphogenesis ('Wolff's law') and the interplay of mutations and selection (Darwin's 'survival of the fittest'). Thus shape-based selective pressures can be determined. In both cases we need to know which biomechanical factors lead to skeletal adaptation, and which ones exert selective pressures on body shape. Each bone must be able to sustain the greatest regularly occurring loads. Smaller loads are unlikely to lead to adaptation of morphology. The highest loads occur primarily in posture and locomotion, simply because of the effect of body weight (or its multiple). In the skull, however, it is biting and chewing that result in the greatest loads. Body shape adapted for an arboreal lifestyle also smooths the way towards bipedality. Hindlimb dominance, length of the limbs in relation to the axial skeleton, grasping hands and feet, mass distribution (especially of the limb segments), thoracic shape, rib curvatures, and the position of the centre of gravity are the adaptations to arboreality that also pre-adapt for bipedality. Five divergent locomotor/morphological types have evolved from this base: arm-swinging in gibbons, forelimb-dominated slow climbing in orangutans, quadrupedalism/climbing in the African apes, an unknown mix of climbing and bipedal walking in australopithecines, and the remarkably endurant bipedal walking of humans. All other apes are also facultative bipeds, but it is the biomechanical characteristics of bipedalism in orangutans, the most arboreal great ape, which is closest to that in humans. If not evolutionary accident, what selective factor can explain why two forms adopted bipedality? Most authors tend to connect bipedal locomotion with some aspect of progressively increasing distance between trees because of climatic changes. More precise factors, in accordance with biomechanical requirements, include stone-throwing, thermoregulation or wading in shallow water. Once bipedality has been acquired, development of typical human morphology can readily be explained as adaptations for energy saving over long distances. A paper in this volume shows that load-carrying ability was enhanced from australopithecines to Homo ergaster (early African H. erectus), supporting an earlier proposition that load-carrying was an essential factor in human evolution.

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What does "arboreal locomotion" mean exactly and what are the relationships between "climbing", environment and morphology?

The characteristics of "climbing" in the sense of locomotion or posture on three-dimensional substrates are discussed from a biomechanical viewpoint. For this purpose, the mechanical conditions of the most widely spread modes of locomotion or gaits used in arboreal surroundings are reviewed. This allows precise identification of morphological characteristics of traits that are advantageous, and therefore have a positive selective value. Further, at least some of the environmental and substrate characteristics that need to be present for using a specific gait, are noted. It turns out that the extremity which is placed lower on the substrate, has to carry a higher load. If this extremity is consistently the hindlimb--which actually is the case in primates, because of understandable, though complex reasons--a division of labor is likely to occur between the limbs: the hindlimb becoming stronger and the forelimb weaker, but more versatile. A very specific, and advantageous feature of the primates is their possession of prehensile hands and feet. That means the autopodia are able (1) to produce by themselves, without the aid of body weight, very high frictional resistance, and (2) to transmit tensile forces as well as torsional moments on the substrate. The above-mentioned division of labor between fore- and hindlimbs implies that the former make the first contacts with and explore the properties of parts of the environment. As a next step, prehensile hands on long arms may easily replace length and mobility of the neck in getting hold of food items. So very characteristic traits of human body shape can be derived to a large extent from the necessities of arboreal locomotion: Prehensile hands, long arms, concentration of body weight on the hindlimbs, shortness of the trunk in comparison to limb length.

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The importance of the evolutionary heritage of locomotion on flat ground in small mammals for the development of arboreality.

The earliest representatives of the mammalian stem line were small. Recent small mammals preserving their morphology possess rather similar kinematic and dynamic locomotor patterns, even if they are not closely related. For a small animal, the mechanics of locomotion on a large branch is comparable to locomotion on flat ground. Combining these informations, it seems sensible to start a discussion on the origins of arboreality with a detailed analysis of the locomotion of small mammals on flat ground. For this purpose, the kinematics of twelve species of mammals were observed using cineradiography, a "general limb" of small mammals was derived as a principle, and its interactions with the trunk were analyzed. These data form the basis for a theoretical upscaling of the motion patterns in arboreal animals, revealing that the transfer of torques between animal and branch becomes unavoidable, thus making the use of prehensile hands advantageous, which by their tendency of distal concentration of muscle masses force the need to change the basic kinematic patterns.

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