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Register a new userThermodynamics of animal locomotion
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Muscles are biological actuators extensively studied in the frame of Hills classic empirical model as isolated biomechanical entities, which hardly applies to a living organism subjected to physiological and environmental constraints. Here we elucidate the overarching principle of a emph{living} muscle action for locomotion, considering it from the thermodynamic viewpoint as an assembly of actuators (muscle units) connected in parallel, operating via chemical-to-mechanical energy conversion under mixed (potential and flux) boundary conditions. Introducing the energy cost of effort, $COE_-$, as the generalization of the well-known oxygen cost of transport, $COT$, in the frame of our compact locally linear non-equilibrium thermodynamics model, we analyze oxygen consumption measurement data from a documented experiment on energy cost management and optimization by horses moving at three different gaits. Horses adapt to a particular gait by mobilizing a nearly constant number of muscle units minimizing waste production per unit distance covered; this number significantly changes during transition between gaits. The mechanical function of the animal is therefore determined both by its own thermodynamic characteristics and by the metabolic operating point of the locomotor system.
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The deep connection between thermodynamics, computation, and information is now well established both theoretically and experimentally. Here, we extend these ideas to show that thermodynamics also places fundamental constraints on statistical estimation and learning. To do so, we investigate the constraints placed by (nonequilibrium) thermodynamics on the ability of biochemical signaling networks within cells to estimate the concentration of an external signal. We show that accuracy is limited by energy consumption, suggesting that there are fundamental thermodynamic constraints on statistical inference.
A model of coupled molecular oscillators is proposed to study nonequilibrium thermodynamics of synchronization. We find that synchronization of nonequilibrium oscillators costs energy even when the oscillator-oscillator coupling is conservative. By solving the steady state of the many-body system analytically, we show that the system goes through a nonequilibrium phase transition driven by energy dissipation, and the critical energy dissipation depends on both the frequency and strength of the exchange reactions. Moreover, our study reveals the optimal design for achieving maximum synchronization with a fixed energy budget. We apply our general theory to the Kai system in Cyanobacteria circadian clock and predict a relationship between the KaiC ATPase activity and synchronization of the KaiC hexamers. The theoretical framework can be extended to study thermodynamics of collective behaviors in other extended nonequilibrium active systems.
Compared to agile legged animals, wheeled and tracked vehicles often suffer large performance loss on granular surfaces like sand and gravel. Understanding the mechanics of legged locomotion on granular media can aid the development of legged robots with improved mobility on granular surfaces; however, no general force model yet exists for granular media to predict ground reaction forces during complex limb intrusions. Inspired by a recent study of sand-swimming, we develop a resistive force model in the vertical plane for legged locomotion on granular media. We divide an intruder of complex morphology and kinematics, e.g., a bio-inspired robot L-leg rotated through uniform granular media (loosely packed ~ 1 mm diameter poppy seeds), into small segments, and measure stresses as a function of depth, orientation, and direction of motion using a model leg segment. Summation of segmental forces over the intruder predicts the net forces on both an L-leg and a reversed L-leg rotated through granular media with better accuracy than using simple one-dimensional penetration and drag force models. A multi-body dynamic simulation using the resistive force model predicts the speeds of a small legged robot (15 cm, 150 g) moving on granular media using both L-legs and reversed L-legs.
Granular media (GM) present locomotor challenges for terrestrial and extraterrestrial devices because they can flow and solidify in response to localized intrusion of wheels, limbs, and bodies. While the development of airplanes and submarines is aided by understanding of hydrodynamics, fundamental theory does not yet exist to describe the complex interactions of locomotors with GM. In this paper, we use experimental, computational, and theoretical approaches to develop a terramechanics for bio-inspired locomotion in granular environments. We use a fluidized bed to prepare GM with a desired global packing fraction, and use empirical force measurements and the Discrete Element Method (DEM) to elucidate interaction mechanics during locomotion-relevant intrusions in GM such as vertical penetration and horizontal drag. We develop a resistive force theory (RFT) to account for more complex intrusions. We use these force models to understand the locomotor performance of two bio-inspired robots moving on and within GM.
We introduce the idea of weakly coherent collisional models, where the elements of an environment interacting with a system of interest are prepared in states that are approximately thermal, but have an amount of coherence proportional to a short system-environment interaction time in a scenario akin to well-known collisional models. We show that, in the continuous-time limit, the model allows for a clear formulation of the first and second laws of thermodynamics, which are modified to include a non-trivial contribution related to quantum coherence. Remarkably, we derive a bound showing that the degree of such coherence in the state of the elements of the environment represents a resource, which can be consumed to convert heat into an ordered (unitary-like) energy term in the system, even though no work is performed in the global dynamics. Our results therefore represent an instance where thermodynamics can be extended beyond thermal systems, opening the way for combining classical and quantum resources.
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