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The Possibility of Zero Limb-Work Gaits in Sprawled and Parasagittal Quadrupeds: Insights from Linkages of the Industrial Revolution

Usherwood, J R


J R Usherwood


Animal legs are diverse, complex, and perform many roles. One defining requirement of legs is to facilitate terrestrial travel with some degree of economy. This could, theoretically, be achieved without loss of mechanical energy if the body could take a continuous horizontal path supported by vertical forces only—effectively a wheel-like translation, and a condition closely approximated by walking tortoises. If this is a potential strategy for zero mechanical work cost among quadrupeds, how might the structure, posture, and diversity of both sprawled and parasagittal legs be interpreted? In order to approach this question, various linkages described during the industrial revolution are considered. Watt’s linkage provides an analogue for sprawled vertebrates that uses diagonal limb support and shows how vertical-axis joints could enable approximately straight-line horizontal translation while demanding minimal mechanical power. An additional vertical-axis joint per leg results in the wall-mounted pull-out monitor arm and would enable translation with zero mechanical work due to weight support, without tipping or toppling. This is consistent with force profiles observed in tortoises. The Peaucellier linkage demonstrates that parasagittal limbs with lateral-axis joints could also achieve the zero-work strategy. Suitably tuned four-bar linkages indicate this is feasibly approximated for flexed, biologically realistic limbs. Where “walking” gaits typically show out of phase fluctuation in center of mass kinetic and gravitational potential energy, and running, hopping or trotting gaits are characterized by in-phase energy fluctuations, the zero limb-work strategy approximated by tortoises would show zero fluctuations in kinetic or potential energy. This highlights that some gaits, perhaps particularly those of animals with sprawled or crouched limbs, do not fit current kinetic gait definitions; an additional gait paradigm, the “zero limb-work strategy” is proposed.


Usherwood, J. R. (2020). The Possibility of Zero Limb-Work Gaits in Sprawled and Parasagittal Quadrupeds: Insights from Linkages of the Industrial Revolution. Integrative Organismal Biology, 2(1),

Journal Article Type Article
Acceptance Date Jun 3, 2020
Publication Date Jun 4, 2020
Deposit Date Oct 12, 2020
Publicly Available Date Oct 12, 2020
Journal Integrative Organismal Biology
Print ISSN 2517-4843
Publisher Oxford University Press
Peer Reviewed Peer Reviewed
Volume 2
Issue 1
Public URL
Publisher URL


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